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      Conservation of Marine Birds of Northern North America, by James C. Bartonek &amp; David N. Nettleship&mdash;A Project Gutenberg eBook.
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<div>*** START OF THE PROJECT GUTENBERG EBOOK 49335 ***</div>

<div class="tnotes covernote">
  <p>The cover image was created by the transcriber and is placed in the public domain.</p>
</div>




<p class="ph2">WILDLIFE RESEARCH REPORTS</p>

<p>This series comprises reports of research relating to birds, mammals, and other
wildlife and their ecology, and specialized bibliographies on these, issued for wildlife
research and management specialists. The Service distributes these reports to official
agencies, to libraries, and to researchers in fields related to the Service's work.</p>


<p><strong>Library of Congress Cataloging in Publication Data</strong></p>

<p>Conservation of marine birds of northern North America.</p>

<p>(Wildlife research report: 11)</p>

<p>Supt. of Docs. no.: I 49.47/4:11</p>

<p>1. Sea birds&mdash;North America&mdash;Congresses. 2. Sea birds&mdash;Northwest,
Pacific&mdash;Congresses. 3. Birds, Protection
of&mdash;North America&mdash;Congresses.
4. Birds, Protection of&mdash;Northwest, Pacific&mdash;Congresses. 5.
Birds&mdash;North
America&mdash;Congresses. 6. Birds&mdash;Northwest, Pacific&mdash;Congresses. I.
Bartonek, James C. II. Natural Resources Council of America. III. United
States. Fish and Wildlife Service. IV. Series.</p>

<p class="center">QL681.C59<span style="margin-left:20%">333.9'5</span> <span style="margin-left:20%">79-607005</span></p>

<p class="center">Use of trade names does not imply U.S. Government endorsement of commercial products.
</p>

<p><span class="pagenum"><a name="Page_i" id="Page_i">[Pg i]</a></span></p>
<div id="halftitle">

<p class="ph1">CONSERVATION OF MARINE BIRDS OF
NORTHERN NORTH AMERICA</p>
</div>
<p><span class="pagenum"><a name="Page_ii" id="Page_ii">[Pg ii]</a></span></p>

<div class="figcenter" style="width: 399px;">
<img src="images/frontispiece.jpg" width="399" height="600" alt="" />
</div>

<p><span class="pagenum"><a name="Page_iii" id="Page_iii">[Pg iii]</a></span></p>
<div id="titlepage">
<div class="chapter"></div>
<hr class="chap" />




<h1>CONSERVATION OF MARINE BIRDS OF<br />
NORTHERN NORTH AMERICA<br />

<span class="large">Papers from the International Symposium<br />
held at the Seattle Hyatt House,<br />
Seattle, Washington, 13-15 May 1975</span></h1>


<p>Edited by</p>

<p>James C. Bartonek and David N. Nettleship</p>


<ul class="p6"><li>Sponsored by</li>
<li>Natural Resources Council of America</li>
<li>National Audubon Society</li>
<li>National Wildlife Federation</li>
<li>U.S. Department of the Interior, Fish and Wildlife Service</li>
</ul>


<p class="p6"><span class="xlarge">UNITED STATES DEPARTMENT OF THE INTERIOR<br /></span>
<span class="large">FISH AND WILDLIFE SERVICE<br /></span>
<strong>Wildlife Research Report 11<br />
Washington, D.C. &#8226; 1979<br /></strong>
</p>
</div>
<p><span class="pagenum"><a name="Page_iv" id="Page_iv">[Pg iv]</a></span></p>

<p class="center">Dedicated to the Memory<br />
of
</p>

<p>Robert D. Bergman, Leonard A. Boughton, and J. Larry Haddock,
Wildlife Biologists of the Fish and Wildlife Service, and Robert
Johnson, Pilot of the Office of Aircraft Services, all of the U.S. Department
of the Interior, who perished in the Gulf of Alaska on 30
September 1974 while conducting aerial surveys of marine birds,</p>

<p class="center">and to
</p>

<p>Einar Brun, Professor of Zoology in Troms&oslash; University and a contributor
to these proceedings, who perished in the Vega Sea on 13 July
1976 when returning from making aerial surveys of marine birds.</p>

<p><span class="pagenum"><a name="Page_v" id="Page_v">[Pg v]</a></span></p>
<div class="chapter"></div>
<hr class="chap" />




<h2>Contents</h2>


<div class="center">
<table class="tdb" border="0" cellpadding="4" cellspacing="0" summary="Contents">
  <tr>
    <td></td>
    <th class="tdb">Page</th>
  </tr>
  <tr>
    <td>Foreword, by Harvey K. Nelson</td>
    <td class="tdr"><a href="#Page_vii">vii</a></td>
  </tr>
  <tr>
    <td>Introduction, by Lynn A. Greenwalt</td>
    <td class="tdr"><a href="#Page_ix">ix</a></td>
  </tr>
  <tr>
    <td>Marine Environment of Birds</td>
    <td class="tdr"><a href="#Page_1">1</a></td>
  </tr>
  <tr>
    <td class="tdp">Long-term Climatic and Oceanographic Cycles Regulating Seabird Distributions and Numbers, by M. T. Myres</td>
    <td class="tdr"><a href="#Page_3">3</a></td>
  </tr>
  <tr>
    <td class="tdp">Sea Ice as a Factor in Seabird Distribution and Ecology in the Beaufort, Chukchi, and Bering Seas, by George J. Divoky</td>
    <td class="tdr"><a href="#Page_9">9</a></td>
  </tr>
  <tr>
    <td>Status of Marine Bird Populations</td>
    <td class="tdr"><a href="#Page_19">19</a></td>
  </tr>
  <tr>
    <td class="tdp">Distribution and Status of Marine Birds Breeding Along the Coasts of the Chukchi and Bering Seas, by James C. Bartonek and Spencer G. Sealy</td>
    <td class="tdr"><a href="#Page_21">21</a></td>
  </tr>
  <tr>
    <td class="tdp">Breeding Distribution and Status of Marine Birds in the Aleutian Islands, Alaska, by Palmer C. Sekora, G. Vernon Byrd, and Daniel D. Gibson</td>
    <td class="tdr"><a href="#Page_33">33</a></td>
  </tr>
  <tr>
    <td class="tdp">The Historical Status of Nesting Seabirds of the Northern and Western Gulf of Alaska, by LeRoy W. Sowl</td>
    <td class="tdr"><a href="#Page_47">47</a></td>
  </tr>
  <tr>
    <td class="tdp">Status and Distribution of Breeding Seabirds of Southeastern Alaska, British Columbia, and Washington, by David A. Manuwal and R. Wayne Campbell</td>
    <td class="tdr"><a href="#Page_73">73</a></td>
  </tr>
  <tr>
    <td>The Biology and Ecology of Marine Birds in the North</td>
    <td class="tdr"><a href="#Page_93">93</a></td>
  </tr>
  <tr>
    <td class="tdp">Trophic Relations of Seabirds in the Northeastern Pacific Ocean and Bering Sea, by David G. Ainley and Gerald A. Sanger</td>
    <td class="tdr"><a href="#Page_95">95</a></td>
  </tr>
  <tr>
    <td class="tdp">Population Dynamics in Northern Marine Birds, by William H. Drury</td>
    <td class="tdr"><a href="#Page_123">123</a></td>
  </tr>
  <tr>
    <td class="tdp">Time-energy Use and Life History Strategies of Northern Seabirds, by Erica H. Dunn</td>
    <td class="tdr"><a href="#Page_141">141</a></td>
  </tr>
  <tr>
    <td class="tdp">Zoogeography and Taxonomic Relationships of Seabirds in Northern North America, by M. D. F. Udvardy</td>
    <td class="tdr"><a href="#Page_167">167</a></td>
  </tr>
  <tr>
    <td>Conflicts Between the Conservation of Marine Birds and Uses of Other Resources</td>
    <td class="tdr"><a href="#Page_171">171</a></td>
  </tr>
  <tr>
    <td class="tdp">Social and Economic Values of Marine Birds, by David R. Cline, Cynthia Wentworth, and Thomas W. Barry</td>
    <td class="tdr"><a href="#Page_173">173</a></td>
  </tr>
  <tr>
    <td class="tdp">Resource Development Along Coasts and on the Ocean Floor: Potential Conflicts with Marine Bird Conservation, by Donald E. McKnight and C. Eugene Knoder</td>
    <td class="tdr"><a href="#Page_183">183</a></td>
  </tr>
  <tr>
    <td class="tdp">Mortality to Marine Birds Through Commercial Fishing, by Warren B. King, R. G. B. Brown, and Gerald A. Sanger</td>
    <td class="tdr"><a href="#Page_195">195</a></td>
  </tr>
  <tr>
    <td class="tdp">Interactions Among Marine Birds and Commercial Fish in the Eastern Bering Sea, by Richard R. Straty and Richard E. Haight</td>
    <td class="tdr"><a href="#Page_201">201</a></td>
  </tr>
  <tr>
    <td class="tdp">Interrelations Between Seabirds and Introduced Animals, by Robert D. Jones, Jr., and G. Vernon Byrd</td>
    <td class="tdr"><a href="#Page_221">221</a></td>
  </tr>
  <tr>
    <td class="tdp">Oil Vulnerability Index for Marine Oriented Birds, by James G. King and Gerald A. Sanger</td>
    <td class="tdr"><a href="#Page_227">227</a></td>
  </tr>
  <tr>
    <td>Programs and Authorities Related to Marine Bird Conservation</td>
    <td class="tdr"><a href="#Page_241">241</a></td>
  </tr>
  <tr>
    <td class="tdp">Programs and Authorities Related to Marine Bird Conservation in Washington State, by Ralph W. Larson</td>
    <td class="tdr"><a href="#Page_243">243</a></td>
  </tr>
  <tr>
    <td class="tdp">Programs and Authorities of the Province of British Columbia Related to Marine Bird Conservation, by W. T. Munro and R. Wayne Campbell</td>
    <td class="tdr"><a href="#Page_247">247</a></td>
  </tr>
  <tr>
    <td class="tdp">Petroleum Industry's Role in Marine Bird Conservation, by Keith G. Hay <span class="pagenum"><a name="Page_vi" id="Page_vi">[Pg vi]</a></span> </td>
    <td class="tdr"><a href="#Page_251">251</a></td>
  </tr>
  <tr>
    <td>Conservation of Marine Birds in Other Lands</td>
    <td class="tdr"><a href="#Page_259">259</a></td>
  </tr>
  <tr>
    <td class="tdp">Conservation of Marine Birds in New Zealand, by Gordon R. Williams</td>
    <td class="tdr"><a href="#Page_261">261</a></td>
  </tr>
  <tr>
    <td class="tdp">Marine Birds in the Danish Monarchy and Their Conservation, by Finn Salomonsen</td>
    <td class="tdr"><a href="#Page_267">267</a></td>
  </tr>
  <tr>
    <td class="tdp">Present Status and Trends in Population of Seabirds in Norway, by Einar Brun</td>
    <td class="tdr"><a href="#Page_289">289</a></td>
  </tr>
  <tr>
    <td>Symposium Summary</td>
    <td class="tdr"><a href="#Page_303">303</a></td>
  </tr>
  <tr>
    <td class="tdp">Conservation of Marine Birds of Northern North America&mdash;a Summary, by Ian C. T. Nisbet</td>
    <td class="tdr"><a href="#Page_305">305</a></td>
  </tr>
  <tr>
    <td>Appendix. Papers and oral summaries presented at the symposium but which do not appear in this publication</td>
    <td class="tdr"><a href="#Page_319">319</a></td>
  </tr>
</table><span class="pagenum"><a name="Page_vii" id="Page_vii">[Pg vii]</a></span></div>
<div class="chapter"></div>
<hr class="chap" />




<h2>Foreword</h2>


<p>The international symposium "Conservation of Marine Birds of Northern North America"
was convened because of a growing awareness that not all was well with our marine birds. The
symposium provided a forum for scientists, governmental administrators, conservationists,
and laypeople to discuss the diverse topics and issues that we must all understand if we are to
act both responsively and responsibly to assure that marine birds will not be lost through our
neglect.</p>

<p>The symposium was cosponsored by the Natural Resources Council of America, National
Audubon Society, National Wildlife Federation, and the U.S. Department of the Interior, Fish
and Wildlife Service; additional support was provided by the Canadian Wildlife Service, the
International Association of Game, Fish, and Conservation Commissioners, the Pacific Seabird
Group, the Sierra Club, the Smithsonian Institution, the Wildlife Management Institute, and
the Wildlife Society.</p>

<p>Persons interested and knowledgeable in the many and varied aspects of marine bird conservation
were invited to participate in this symposium. There were 139 registered and several
score of unregistered participants in attendance. Major topics treated were: (1) socioeconomic
considerations and conservation of marine birds; (2) the marine environment of birds; (3) status
of marine bird populations on land and sea; (4) the biology and ecology of marine birds in the
North; (5) conflicts between the conservation of marine birds and uses of other resources;
(6) programs and authorities related to the conservation of marine birds; and (7) conservation
of marine birds in other lands.</p>

<p>The objective of the symposium was to identify problems and the needed information and
programs necessary for the conservation of marine birds of northern North America. For the
purpose of this symposium the term "northern North America" referred to the coasts of Washington,
British Columbia, Alaska, Yukon Territory, and Northwest Territories and the adjacent
North Pacific and Arctic Oceans. "Marine bird" was defined as being any bird using
marine or estuarine waters. Speakers were asked to describe the status of information or the
state of the art as it pertained to their topic within the limitations set by the objective of the
symposium. Examples from other regions and of bird species not found in the regions of concern
were to be used for comparative purposes when little pertinent information was known for
regions or species of concern. Speakers were asked to identify the gaps in the knowledge and
methodology that are most critical to their topic.</p>

<p>I believe that this symposium was particularly successful in that it provided a timely forum
for many scientists who were about to embark on studies of marine birds in those areas of
Alaska and California being considered for outer continental shelf oil and gas exploration and
development. These published proceedings may be of lesser importance from that standpoint
because some data, particularly those on populations, are out of date. However, I believe that
the proceedings will long be of importance to biologists and administrators alike in charting
their respective courses to ultimately assure conservation of this valuable avian resource.</p>

<p>Many people from many organizations and agencies worked hard to put together the symposium
in the relatively short time of about 8 months. Nathaniel P. Reed was the person primarily
responsible for bringing this symposium to fruition. The Steering Committee was composed
of Daniel A. Poole, John S. Gottschalk, David N. Nettleship, Amos S. Eno, C. Eugene
Knoder, Warren G. King, Louis Clapper, Robert Hughes, Fred G. Evenden, James C. Bartonek,
and me. James C. Bartonek, Warren G. King, David N. Nettleship (Co-chairmen),
C. Eugene Knoder, David A. Manuwal, William H. Drury, and Spencer G. Sealy served on the
Program Committee. David A. Manuwal and Terence R. Wahl arranged trips for persons to
observe pelagic birds off the Washington coast and other birds on Skagit Flats. C. Eugene
Knoder handled financial matters. John A. Sayre and Richard Bauer made arrangements for
facilities and entertainment. Elaine Rhode prepared the program and abstracts for printing.
John Pitcher kindly contributed the artwork used in this publication as well as that used in the
program and abstracts.</p>

<p><span class="pagenum"><a name="Page_viii" id="Page_viii">[Pg viii]</a></span></p>

<p>George Reiger made general introductions to the symposium; Spencer G. Sealy, Daniel W.
Anderson, and I served as Session Chairmen; and James C. Bartonek served as General Chairman.
Elvis J. Stahr was guest speaker at the symposium banquet.</p>

<p>Most credit for the success of this symposium goes to the 52 persons who as authors,
coauthors, or summarizers of sessions presented much meaningful information in their presentations,
during recorded discussions, and during many informal occasions. I wish to make
special recognition of Ian C. T. Nisbet for his skillful summary of the symposium.</p>

<p>Editorial assistance in preparing the proceedings was provided by Judith Brogan.</p>

<p style="margin-left:50%">
Harvey K. Nelson<br />
<br />
<em>Chairman of Symposium and<br />
Director of Wildlife Resources</em><br />
</p>

<p><span class="pagenum"><a name="Page_ix" id="Page_ix">[Pg ix]</a></span></p>
<div class="chapter"></div>
<hr class="chap" />




<h2>Introduction</h2>


<p>Migratory birds make up a resource that is shared by many people of many
nations. Public awareness of marine birds&mdash;their manifold values, ecological
requirements, and problems&mdash;is prerequisite to their protection. I
am proud that the Fish and Wildlife Service can further this needed awareness
by publishing these proceedings of the international symposium
"Conservation of Marine Birds of Northern North America."</p>

<p class="right">Lynn A. Greenwalt, <em>Director<br />
Fish and Wildlife Service</em></p>
<p><span class="pagenum"><a name="Page_x" id="Page_x">[Pg x]</a><br /><a name="Page_1" id="Page_1">[Pg 1]</a></span></p>

<p><span class="pagenum"><a name="Page_2" id="Page_2">[Pg 2]</a><br /><a name="Page_3" id="Page_3">[Pg 3]</a></span></p>
<div class="chapter"></div>
<hr class="chap" />




<h2>MARINE ENVIRONMENT OF BIRDS</h2>
<div class="chapter"></div>
<hr class="chap" />




<h3><a name="Long-term_Climatic_and_Oceanographic_Cycles" id="Long-term_Climatic_and_Oceanographic_Cycles">Long-term Climatic and Oceanographic Cycles
Regulating Seabird Distributions and Numbers</a></h3>


<p class="center">by</p>

<p class="center">M. T. Myres</p>

<p class="center">Department of Biology, University of Calgary<br />
Calgary, Alberta, Canada T2N 1N4
</p>


<h4>Abstract</h4>

<div class="blockquot">

<p>Seabird ornithologists have generally paid little attention to the possible roles
played by long-term climatic cycles or air-ocean interactions on population
changes at established colonies or on the processes of colony establishment or
extinction. Yet, a rapidly expanding literature in the physical sciences suggests
that seabird numbers are not naturally stable at particular colonies for any great
length of time. It is suggested that the establishment of new colonies at one end
of the range may counter the decline of colonies at the other end. Perhaps these
changes in small marginal colonies are important, and they may be more indicative
and significant (when detected and explained) than are much larger changes
in numbers in bigger reproductive units in the center of a species' range. Fluctuations
in seabird numbers must in future be first considered as possible responses
either to short-term, or turnarounds in longer term, natural climatic or oceanographic
cycles, or to trends ranging in length from a few years to at least several
decades.</p></div>

<p>During the last 30 years extensive literature
in the fields of physical and biological
oceanography has accumulated that is not
readily accessible to the nonprofessional student
of seabirds and not as widely understood
by career seabird ornithologists as it should
be. This literature in oceanography and
marine fisheries is as extensive in Russian and
Japanese together as in the main languages of
Western Europe combined; this abundance
compounds the problem of becoming familiar
with it if, as a student of seabirds, one's interest
in the literature is initially somewhat
marginal. Nevertheless, to achieve the best
possible appreciation of the oceanographic influences
affecting seabirds, particularly in the
north Pacific Ocean and its adjacent embayment
seas, it is necessary to make the effort.</p>

<p>Because of the rigor of carrying out their
primary duties while at sea, only a very few
North American and European oceanographers
or fishery biologists have found time
to interest themselves in seabirds and then,
with a few notable individual exceptions, only
as an off-duty pastime. The reason is not far
to seek. It is far less important to examine the
ecology of organisms at the next highest level
of the food chain to the ones that are the primary
concern than it is to examine the next
lowest level (the food of the fishes or, in the
case of phytoplankton, the physical and
chemical environment in which the organisms
grow best).</p>

<p>Seabirds are at the very top of the marine
food chain, and they are not wholly aquatic in
any case since they mainly travel through the
air rather than the water and reproduce on
land rather than in the sea. Only with the relatively
recent recognition that seabirds contribute
to the recycling of nutrients back into
the ocean to an important degree, have seabirds
gained a new scientific constituency.</p>

<p>At about the same time, governments have
begun to recognize that seabirds are relatively
easily examined indicators of the presence
of unseen chemical pollutants in coastal
seas, perhaps primarily for the very same reasons
that they were previously so largely
ignored; namely, that they are at the top of
the food chains (and so accumulate the most-persistent
and least-degradable pollutants)
and that the on-land failures in their reproductive
biology are readily visible.</p>

<p><span class="pagenum"><a name="Page_4" id="Page_4">[Pg 4]</a></span></p>

<p>During the last 10 years, it has become evident
that yet another fundamental science is
even more basic to the achievement of a balanced
and in-depth understanding of the influence
of the environment upon seabirds&mdash;the
combined field of astrophysics, geophysics,
and climatology. New developments in this
field (when they are not published in <cite>Nature</cite> or
<cite>Science</cite>) appear in journals that are less
familiar to seabird ornithologists than those
in which the fishery biologists and biological
oceanographers publish their findings.</p>

<p>Unfortunately, important advances in
understanding the dynamics and energy
transport mechanisms of both the atmosphere
and the water masses of the oceans are
not being picked up by students of seabirds
because of the natural lag in communication
that occurs between disparate disciplines.
Only in the last few years have oceanographers
and climatologists been invited to
address gatherings of ornithologists, and the
modesty with which they have sometimes
done so has limited the impact of their offerings.</p>

<p>At this symposium, it was left to a biologist
with no pretentions in either physics or
mathematics to demonstrate the need for seabird
ornithologists to understand basic environmental
processes well beyond their usual
range of interests. I did so with a series of
slides taken from this "other" literature, and
I had intended to include in the published version
of this paper an extensive bibliography,
subdivided into category groupings, so that
seabird ornithologists could make their own
selection of the points in the spectrum at
which they most needed information.</p>

<p>Unfortunately, limitations upon space in
this volume, daily additions to the exploding
literature, and my own inability to keep up
with understanding this have forced me to
omit any references and not to attempt to expound
detailed specific physical mechanisms.</p>

<p>Thus unencumbered here, I shall briefly outline
instead what I perceive to be some of the
significance for seabird ornithology and conservation
of the rapidly expanding understanding
of the oceans, the air-sea interface,
atmospheric dynamics, and influences upon
the world's climate of extraterrestrial events.</p>


<h4>Small-scale or
Short-term Influences</h4>

<p>There is no need to dwell on the well-known
events that could be mentioned under this
heading. Seabird ornithologists are familiar
with the fact that the atmosphere is the
medium of seabirds both when searching the
ocean for feeding areas and when on migration,
and also a violent enemy, as when particular
storms cause occasional "wrecks" of
seabirds inland from coastlines. As a refinement
of the former, Manikowski of Poland
suggests that seabirds respond to the passage
of weather systems, so that their distribution
over the open ocean may be constantly changing.
Whereas some species may attempt to
avoid the stormy conditions of low-pressure
areas (cyclonic conditions), others more highly
specialized for exploiting the aerodynamic
properties of wind over a moving water surface
may possibly, instead, try to avoid large
high-pressure regions (anticyclonic conditions
with little or no wind). My student, Juan Guzman,
is attempting to determine whether this
may be so; if it is, it might be possible, for
example, to predict some things about the distribution
patterns and population structure of
southern hemisphere shearwaters while they
are visiting the oceans of the northern hemisphere
during the nonbreeding season.</p>

<p><span class="pagenum"><a name="Page_5" id="Page_5">[Pg 5]</a></span></p>

<p>In comparison with the "wrecks" brought
about by storms, which are of short duration
and not usually very serious, seabird ornithologists
are also familiar with relatively
brief and localized disasters caused by
changes in the ocean itself. The best-known
example is a slight change in the boundary of
an ocean current (or other shift in the position
of a distinctive water mass) that results in the
failure of food fishes to appear as they normally
would, close to breeding sites of conspicuous
colonial seabirds, such as the periodic
shift in the El Ni&ntilde;o off the west coast of
South America. A scarcely studied refinement
of this type of event would be the effects of
less-pronounced oceanic changes that might
reduce the planktonic food supply of nocturnally
active, burrow-nesting seabirds. In
such instances, the effects might also be a
breeding failure for only one or two seasons;
in all probability such events occur, but
whether they are as likely to be detected by us
is problematical. However, the populations of
most seabirds are probably already adapted
to survive short-term crises of this type because,
having long adult life spans, reproductive
adults that fail to raise young one year
may mostly live to succeed in doing so in the
next or succeeding year, when the oceanic
"anomaly" has disappeared. What constitutes
an "anomaly" will be considered again
shortly.</p>

<p>A third critical condition for seabirds may
be local or widespread, temporary or final, or
some combination of these. A single local
spill, or outfall, of a chemical pollutant will be
short term if we can take steps to alleviate the
consequences or stem the flow. Alternately,
we may consider it to be long term if we take
the view that it is one additional act of violence
resulting from the "progress" of Industrial
Man, and that it is never going to shift
into reverse gear. We may say that the effect
on seabird populations of spills of oil products
or chemical pollutants in coastal waters of a
region will be a "final solution" for any that
become wholly extinct before the oil wells go
dry or the industries fail. On the other hand,
the effect will have been merely a perturbation
of the population if the species survives
and outlives these activities. Recent upturns
in populations of peregrine falcons <em>(Falco
peregrinus)</em> and pelicans <em>(Pelecanus sp.)</em> in certain
places where environmental controls
have been enacted give us hope that crises of
several years' duration can be withstood by at
least those species that once were common in
relation to their respective food sources or
available safe breeding habitats. The really
critical features to document are the means
whereby abandoned breeding sites are reoccupied
and the time it takes.</p>

<p>It must never be forgotten that we know almost
nothing about the ecology of subadult or
nonreproductive adult seabirds during the
years they are at sea unconfined by membership
in a breeding unit and that we know almost
nothing about the activities of pelagic
seabirds in the nonbreeding season. These
birds may be far from land and hard to study,
but what happens during those phases of
their lives is basic to the composition of the
colony and condition of the birds when breeding.
A start would be to learn everything that
is known and is being discovered about the
oceans by oceanographers and, thus forearmed,
go looking for the seabirds with certain
questions clearly in mind.</p>


<h4>Detecting the Effects
of Long-term Cycles</h4>

<p>A scientist's working life lasts only a few
decades, and few studies of seabirds by a
single author or agency have been continued
for longer than 5-10 years on any one problem.
Further, while we as individuals may live
to be equally active in a certain field of research
20 years hence, our collective conscience
and collective muscle consist of
several levels of government that tend to exhibit
4- or 5-year changes of direction and
priorities. Certainly, the civil service may live
on as an inertial recorder of collective experience.
Certainly, too, those who live under one
form or another of dictatorship or, as in some
Canadian provinces, where conservative patterns
of voting occur, may experience a continuity
of research and development and conservation
policies that exceed the 4- to 5-year
turnaround pattern that is most common.
Yet, even these more continuous systems may
come to an end quite suddenly because of economic
or political happenstance.</p>

<p>The point of this digression is to show that
seabird ornithologists must not rely on government
programs to provide continuous data
over a long period of years&mdash;not, at least, in
most countries. Monitoring the biological circumstances
of seabirds is not the same as recording
the temperature regularly by machine
at a weather station, since this activity is unlikely
to be terminated unless the society collapses
altogether. We may know that in some
countries the amateur naturalist exists in
such numbers that records of seabirds will
continue to be made whatever the circumstances.
Nevertheless, planning of censuses
that will be repeated every 10 years is best assured
if government and career biologists
combine with the amateur element, so that
any one of them can continue the work if any
other element should be incapacitated. At any
one time, either the amateur or the government
or the university personnel may be the
prime mover, and each of these forms now
exists in various countries.</p>

<p><span class="pagenum"><a name="Page_6" id="Page_6">[Pg 6]</a></span></p>

<p>What the scientific literature in the fields of
the geophysical, atmospheric, and oceanographic
disciplines demonstrates is that natural
climatic oscillations probably range in
length from the 11-year sunspot cycle through
several decades (or a human lifetime) to
several hundred years. So, when our children
are the new trustees of seabird colonies 20 or
40 years hence, they must interpret their data
using the full range of physical as well as biological
data that we can leave for them. Indeed,
the information is, I believe, already
available over a long enough period (since
1940 at least) to allow some speculative interpretations
of what may have been happening
to our seabird populations, whether or not we
knew or had any evidence of it.</p>

<p>I have already suggested that extraterrestrial
events, particularly the 11-year sunspot
cycle, are increasingly believed to influence
the atmosphere of this planet. The Chinese
and Japanese have remarkably precise records
of the northern limits of certain agricultural
crops at particular times, the phenology
of flowering, and the freezing of lakes. These
demonstrate long-term trends in overall climate
in eastern Asia that extend over hundreds
of years. The climate of Japan is influenced
by the high-pressure area in winter over
mainland East Asia. There is evidence that
severe ice conditions in the Bering Sea during
the early 1970's may have been due to an eastward
shifting of this high-pressure area.
Again, the water mass of the Kuroshio Extension
and the West Wind Drift takes several
years to travel across the Pacific Ocean, and
there is an established temperature variation
that travels like a slow wave with it. Off
Japan, the Kuroshio Current periodically develops
meanders which slow the speed of the
eastward flow. Cold and warm "pools" of
water approach the west coast of Canada and
the western United States from time to time.</p>

<p>Ocean currents are driven by the atmospheric
motion above them, which consists of
several convective cells between the equator
and each pole. The outcome is zonal winds,
such as the trade winds and the westerlies.
However, as the influence of the sun on the atmosphere
is variable, the input of heat and the
extent of the major high-pressure areas vary,
as does the path of the jet stream. The recent
droughts in northern Africa and unusually
heavy rains in Australia are both linked to a
southward shift of the Intertropical Convergence
Zone in the atmosphere and a "corrugation"
of the wind circulation from a more normal
zonal (latitudinal) path. These shifts in
the atmospheric circulation are almost certainly
transmitted also to the ocean currents
and the marine ecosystem, with the influence
being felt for a long period of years.</p>

<p>One of the oceanic domains of the North
Pacific is the transitional domain, which lies
east-west where the West Wind Drift impinges
upon the coasts of British Columbia
and Washington State. It is precisely in this
sector that there was a well-documented
"temperature anomaly" in 1957-58. Since an
anomaly implies something completely out of
the ordinary, I seriously question the appropriateness
of the term for an event that may
or may not be recurrent (at the time it was a
pronounced variation from the oceanographic
records accumulated up to that time, but the
period had not been a very long one). It is no
coincidence that the numbers of albatrosses
recorded at Ocean Weather Station "Papa"
was higher during this warm-water "anomaly"
than subsequently (indeed, an 18-year record
of the seabirds recorded at "Papa" also exhibits
other interesting fluctuations from the
base-line data in certain years).</p>

<p>Recent analyses of sediments from off the
coast of California have demonstrated long-term
fluctuations in sardine populations extending
back at least 1,800 years, with increases
lasting 20-150 years and spaced
20-200 years apart. The number of anchovies
declined steadily. Yet until now, El Ni&ntilde;o
events have been treated as anomalies in that
region as well as off the coast of Peru. Just as
we recognize that different species of fish follow
the warm water north on such occasions,
we must also recognize the rather distinct seabird
species assemblage that is trapped, as it
were, in the Gulf of California. Clearly, like
the termination point of the West Wind Drift
at about the 45-55&deg; parallel, the coast of Baja
California and southern California State, from
the 25-35&deg; parallel where the California Current
begins to swing away from the coast to
the west as the North Equatorial Current, is
another zone of instability.</p>

<p><span class="pagenum"><a name="Page_7" id="Page_7">[Pg 7]</a></span></p>

<p>I think that it is no accident that the southern
limit of several northern species of North
Pacific seabirds ends in southeastern Alaska
or northern British Columbia, and that the
northern limit of the ranges of several other
species occurs in Washington State or southern
British Columbia. Indeed, the west coast
of Vancouver Island is not rich in species, and
several of those that exist are not present in
great numbers. This is a region of rather more
variable conditions than elsewhere, and
species evidently find that it is difficult to
colonize and it quickly becomes unsuitable
again. Since 1940, indeed, there has been a
parallel decline in the annual mean sea-surface
temperature at a number of coastal recording
stations in British Columbia, and this seems
to have been a rebound from a less well-documented
rise in sea-surface temperatures during
the 20 years before that, which culminated
in a peak around 1940. Salinity has likewise
trended downwards during the last 30 years.
The seabird colony size data before 1960 are
so nonquantitative that it is impossible to be
sure what changes in seabird populations and
breeding sites may have taken place in response
to these physical changes.</p>

<p>The lesson is that we must now examine all
future census and distribution data with
trends in sea-surface temperature and salinity
in mind as two of several likely factors influencing
them. We must no more ignore data
outside our own field than a salmon ecologist
might.</p>


<h4>Conclusions</h4>

<p>We know little of the accuracy of censuses
of seabird numbers made between 1850 and
1950. There has been a tendency to assume
that numbers of seabirds at long-established
colonies have been relatively unchanging,
even though the expansion of some species
into previously unrecorded breeding sites in
low numbers is well documented. Contraction
of breeding ranges, likewise, has most commonly
been attributed to the influence of man.
Recent literature from the physical sciences,
on the contrary, suggests that seabird numbers
at particular colonies are most unlikely
to have been stable for any great length of
time, at least at high or middle latitudes and
particularly at points where boundaries between
currents impinge on continental coasts.
Indeed, some early estimates of colony sizes
may not have been as much in error as we may
have assumed, neither when apparently too
large nor when apparently unlocated by previous
visitors.</p>

<p>The halving of a large colony over a period
of 20 to 50 years in the middle of the range of
a species and the establishment and disappearance
of smaller breeding groups at opposite
extremes of the range (both latitudinally
and longitudinally), may equally reflect natural
long-term climatic or oceanographic
changes and may naturally be reversed at
some time in the future, perhaps within half a
century. The implication for conservation of
seabird colonies that are at the contracting
end of a species' range is that cultural rather
than biological criteria may be the best
determinants.</p>

<p><span class="pagenum"><a name="Page_8" id="Page_8">[Pg 8]</a><br /><a name="Page_9" id="Page_9">[Pg 9]</a></span></p>
<div class="chapter"></div>
<hr class="chap" />




<h3><a name="Sea_Ice_as_a_Factor_in_Seabird_Distribution_and" id="Sea_Ice_as_a_Factor_in_Seabird_Distribution_and">Sea Ice as a Factor in Seabird Distribution and
Ecology in the Beaufort, Chukchi, and Bering Seas</a></h3>


<p class="center">by</p>

<p class="center">George J. Divoky<a name="FNanchor_1_1" id="FNanchor_1_1"></a><a href="#Footnote_1_1" class="fnanchor">[1]</a></p>

<p class="center">U.S. Fish and Wildlife Service<br />
Fairbanks, Alaska
</p>


<h4>Abstract</h4>

<div class="blockquot">

<p>Arctic sea ice has a variety of effects on seabirds. Although the decrease in surface
area available for feeding and roosting is probably the major restrictive
effect, also important are productivity of water covered by ice and the reduced
prey abundance in nearshore areas due to ice scour. The most important benefit
that sea ice provides to seabirds is the plankton bloom that occurs in the ice in
the spring. In the Beaufort and Chukchi seas this bloom supports an under-ice
fauna that is an important food source for seabirds.</p></div>

<p>Sea ice is a major factor in the distribution
and ecology of many of the birds treated in
this symposium. Sea ice is defined here as ice
formed by the freezing of seawater and includes
both free floating pack ice and the
more stable shorefast ice. Since icebergs are
composed of ice of land origin, they are not
discussed.</p>

<p>Before discussing the specific relationship
of birds and sea ice in the Beaufort, Chukchi,
and Bering seas, I list the general effects that
arctic ice can have on seabirds. For purposes
of discussion these effects can be divided into
negative effects, or disadvantages, and positive
effects, or advantages.</p>


<h4>General Effects of
Ice on Birds</h4>


<h5><em>Negative Effects</em></h5>


<h6>Sea Ice Decreases the Surface Area of Water</h6>

<p>The decrease in the surface area of water is
the simplest and most immediate effect that
sea ice has on birds. Ice acts as a barrier that
restricts the availability of food in the water.
Surface feeders are the most severely affected
since, in general, ice cover of 50% reduces the
possible feeding area by half. The effect on
diving species is not as severe since, if open
water is scattered throughout the ice, diving
species still have access to much of the prey in
the water column and benthos. When open
water is scarce, however, diving species can
become concentrated in the available water,
resulting in intense competition for available
prey. In certain situations the open water is
used only as a migratory pathway, but open
water is necessary for birds that must roost or
feed.</p>


<h6>Sea Ice Reduces Primary Productivity
in the Water Column</h6>

<p>Ice inhibits phytoplankton blooms in the
water column, thus decreasing the biological
productivity of ice-covered waters. This inhibition
occurs in two ways:</p>

<p>&#8226; <em>By decreasing light penetration of the
water column.</em>&mdash;Much of the sunlight reaching
the ice is reflected by the ice and by snow on
the ice. The amount of light reaching the
water depends on the angle of the light, thickness
of ice, and amount of snow cover. When
the layer of under-ice algae forms, it absorbs
light and further reduces the amount of light
reaching the water (Bunt 1963). This reduction
in light reduces the depth of the euphotic
zone.</p>
<p><span class="pagenum"><a name="Page_10" id="Page_10">[Pg 10]</a></span></p>
<p>&#8226; <em>By increasing the stability of the water
column.</em>&mdash;Increased stability of the water
column reduces the upwelling of nutrient-rich
waters into the euphotic zone. Ice stabilizes
the water column primarily by preventing
wind-driven movement of surface waters and
by forming a layer of meltwater at the surface
in the spring and summer (Dunbar 1968).</p>


<h6>Sea Ice Reduces Benthic and
Intertidal Biota</h6>

<p>Benthic flora and fauna can be reduced by
the presence of ice in two ways: In shallow
water ice can freeze to the bottom for much of
the year and prevent the establishment of
plant and animal populations; and when ice
floes are pushed together, they form underwater
ice keels that can scour the bottom
when the ice moves. Both of these events not
only act directly to decrease benthic populations
but also disturb the sediment, making it
less suitable for colonization. In areas with
heavy ice scour, sessile benthic populations
can be greatly reduced, although motile
species may move into scoured areas during
the ice-free period in summer. In addition to
preventing the establishment of sessile
benthic animal populations, ice scour also prevents
the establishment of beds of kelp and
eelgrass <em>(Zostera marina)</em>, thus decreasing the
diversity and productivity of arctic inshore
waters. Both kelp and eelgrass beds are important
feeding sites for birds in areas south
of the region affected by ice scour.</p>


<h6>Sea Ice Allows Terrestrial Predators
Access to Breeding Sites</h6>

<p>The formation of ice between the mainland
and offshore islands allows the arctic fox <em>(Alopex
lagopus)</em> and other predators access to the
islands used by breeding birds. Foxes can become
permanently established on islands that
have food sources during the period when
birds are absent from the island. Often, however,
there is little to attract foxes to the islands
other than breeding birds. Because
moats form around many islands before the
breeding birds arrive, foxes are primarily a
problem when moat formation is incomplete
or when the breakup of ice is late. Arctic foxes
are found on the pack ice throughout the summer
and thus can visit islands that are separated
from the mainland by open water but
are adjacent to the pack ice.</p>


<h5><em>Advantages</em></h5>


<h6>Sea Ice Provides a Matrix and
Substrate for an Ice-associated Plankton
Bloom and an Associated Under-ice Fauna</h6>

<p>The first detailed studies on the blooms of
diatoms that occur in the lower levels of ice
were done by Appollonio (1961). The importance
of this bloom in the energy budgets of
arctic and subarctic seas has only recently
been realized (Alexander 1974; McRoy and
Goering 1974). In areas where ice is present
throughout the year, the plankton bloom supports
a population of under-ice invertebrates.
These populations have been little studied but
apparently consist primarily of copepods and
amphipods (Mohr and Geiger 1968). Feeding
on the invertebrates associated with the ice
are two species of fish, polar cod <em>(Arctogadus
glacialis)</em> and arctic cod <em>(Boreogadus saida)</em>.
Andriashev (1968) used the term cryopelagic
to describe such fish, which are found in the
midwater zone but also are associated with
ice during some part of their life cycle.</p>

<p>The underside of multi-year ice has
numerous ridges and pockets that provide a
heterogeneous environment for the under-ice
fauna. This environment is protected from
disturbance from currents and wave action by
ice keels acting as barriers, which also provide
shelter from predators in the same manner as
a coral reef. The overall effect of the under-ice
flora and fauna is to increase the diversity of
surface waters in arctic seas by creating an inverted
benthic biota.</p>


<h6>Sea Ice Provides Hauling Out
Space for Marine Mammals</h6>

<p>The mammals that inhabit the ice in the
Chukchi and Bering seas and their adaptations
to the pack ice environment were discussed
by Fay (1974). Many of these species
frequently haul out on the ice, where they provide
food in the form of feces, placentas, and
carcasses.</p>


<h6>Sea Ice Provides Roosting Sites</h6>

<p>Ice provides a hard substrate that allows
seabirds to leave the water to roost. This
allows such species as the <em>Larus</em> gulls, which
typically roost on hard substrates, to occur in
large numbers well offshore.</p>

<p><span class="pagenum"><a name="Page_11" id="Page_11">[Pg 11]</a></span></p>


<h6>Sea Ice Reduces Wind Chill</h6>

<p>The unevenness of the upper surface of the
ice reduces the speed of winds directly over
the ice, thus providing a microhabitat and reducing
the amount of wind chill for birds sitting
on and next to the ice.</p>


<h6>Sea Ice Decreases Wave Action</h6>

<p>Ice floating on the water reduces the surface
disturbance of the water. Although
swells pass through areas with much ice
cover, waves do not. In addition, surface
waters on the lee side of ice floes and cakes
usually have little surface disturbance. Surface
feeders may be able to locate prey more
easily because of these reductions in surface
disturbance.</p>


<h4>Specific Effects of Ice
on Birds in the
Western Arctic</h4>

<p>The retreat of the pack ice each spring and
the formation of new ice each fall greatly affect
a large area of the Arctic Ocean off the
coast of Alaska and much of the Bering Sea.
Specific ways in which birds are affected by
ice in the western Arctic are discussed on a
seasonal basis. All observations are my own,
unless otherwise stated.</p>


<h5>Winter</h5>


<h6>Chukchi and Beaufort Seas</h6>

<p>From late November to mid-April, ice cover
of the Chukchi and Beaufort seas is almost
complete. The only areas where birds can be
expected to winter in these seas are the
chronic lead systems. Such lead systems are
found off Wainwright and Point Barrow and
south of the Point Hope-Cape Thompson
area (Shapiro and Burns 1975). Only the black
guillemot <em>(Cepphus grylle)</em> is known to regularly
winter offshore from Wainwright and
Point Barrow (Gabrielson and Lincoln 1959;
Nelson 1969). In the Point Hope-Cape Thompson
area, glaucous gulls <em>(Larus hyperboreus)</em>,
the common murre <em>(Uria aalge)</em>, and the thick-billed
murre <em>(U. lomvia)</em> occur throughout the
winter (Swartz 1967). It is likely that black
guillemots are also found in this area.</p>

<p>The lack of chronic lead systems in the
Beaufort Sea precludes the presence of wintering
seabirds. The one species that may be
found wintering in the Beaufort is the Ross'
gull <em>(Rhodostethia rosea)</em>. Ross' gull is believed
to winter primarily in the Arctic Ocean
(Bailey 1948). The number of sightings that
have been obtained in both the eastern and
western Arctic indicate that the species may
winter over much of the Arctic Ocean. It may
thus be expected to occur in both the Chukchi
and Beaufort seas during winter.</p>

<p>Ice cover&mdash;not prey abundance&mdash;plays the
major role in severely limiting bird numbers
in the Arctic Ocean in winter. Prey is known
to be abundant in parts of the Arctic Ocean
during the period of ice cover. In the Chukchi
Sea, Eskimos fishing through the ice can
catch 23 kg of arctic cod per person per day
(D.C. Foote, unpublished data). Eskimos jig
for the fish at considerable depths, and the
cod do not appear to be as common directly
below the ice as they are in summer. The effects
of new ice (which forms on the underside
of the ice during the winter) on the under-ice
fauna are not known. The abundance of amphipods
in ice-covered waters in winter is
demonstrated by the experience of the
Greeley Expedition in the eastern arctic. They
discovered that any scrap of food thrown into
a lead was quickly consumed by amphipods.
Nets were made to catch the amphipods and
the availability of this food source played a
major part in the survival of the expedition
(Schmitt 1965).</p>

<p>Aside from the food found in leads in the
ice, the only food available to birds in the
Beaufort and Chukchi seas in winter is carrion
and the feces of mammals found on the pack
ice. The presence of the arctic foxes on the
pack ice during the winter demonstrates the
availability of scavenging opportunities on
the ice. Arctic foxes on the pack ice live on
feces and the remains of seals killed by polar
bears <em>(Ursus maritimus)</em>. Polar bear and seals
are both common in the Beaufort and Chukchi
seas in winter, but no scavenging seabirds are
found there in the winter. It was thought that
the ivory gull <em>(Pagaphila eburnea)</em> was associated
with marine mammals during the winter,
but they are now known to winter at the
Bering Sea ice edge, where they feed on fish
and crustaceans (Divoky 1976). The only birds
associated with polar bear kills in the Chukchi
Sea in March are ravens, <em>Corvus corax</em> (T. J.
Ely, Jr., personal communication).</p>

<p><span class="pagenum"><a name="Page_12" id="Page_12">[Pg 12]</a></span></p>


<h6>Bering Sea</h6>

<p>Ice begins to cover the northern Bering Sea
in November and reaches its maximum by
February, when it usually extends as far
south as the edge of the continental shelf, and
covers nearly 75% of the surface of the Bering
Sea (Lisityn 1969). Coverage can vary greatly
from year to year. In certain years Bristol
Bay may be completely covered and in others
ice is found only in the northern part of the
Bay. Almost all ice in the Bering Sea is first-year
ice. This ice tends to be flat on the top
and underside and in general lacks the extensive
keels and pressure ridges found on multi-year
ice.</p>

<p>The Bering Sea ice has a number of large-scale
features of importance to birds. The
"front" is a zone of ice south of the consolidated
pack that is composed of small floes, ice
pans, and brash ice. This zone is prevented
from forming large floes by the action of
swells from the open water to the south. The
front continually changes in width. When
winds are from the south, it is compressed
into a narrow band; when winds are from the
north, it is a broad zone composed of bands of
ice interspersed with open water.</p>

<p>Polynias (areas of open water) are found immediately
south of the large islands in the
northern Bering Sea. They are formed by the
southward movement of ice caused by the prevailing
winds. This movement causes ice to be
pushed away from the south side of islands,
leaving areas of open water. Large polynias
are associated with St. Lawrence, St.
Matthew, and Nunivak islands and with the
south side of the Seward Peninsula (Shapiro
and Burns 1975).</p>

<p>The most biologically active area of the
Bering Sea in winter is the ice front. Studies
of primary productivity in April show that
production at the surface in the ice front is
high (1.98 mg C/m<sup>3</sup> per h). Surface waters directly
under the pack ice have much lower production
(0.29 mg C/m<sup>3</sup> per h), and that in the
water south of the ice is lower yet. At this
time production within the ice is very high
(more than 5 mg C/m<sup>3</sup> per h) (McRoy and
Goering 1974). Because this phytoplankton
bloom is trapped in the ice, it is not available
to grazers. Thus, before the spring melt the
ice front is the only area where a large quantity
of phytoplankton is available to higher
levels of the marine food chain.</p>

<p>The winter distribution of birds in the
Bering Sea correlates well with the findings
on primary productivity. Densities south of
the ice and the continental shelf average less
than 10 birds/km<sup>2</sup>. At the ice front during one
cruise in March, densities exceeded 500
birds/km<sup>2</sup>. Densities at the ice front increase
from south to north; they drop in the region
where the ice front grades into more consolidated
pack ice, and are less than 0.1 bird/km<sup>2</sup>
in the consolidated pack.</p>

<p>The most numerous species at the ice front
are common and thick-billed murres, which
constitute more than 90% of all birds seen.
Irving et al. (1970) were the first to report on
the large number of murres at the ice front.
Feeding flocks of 25,000 individuals have
been observed at the front, in which densities
were as high as 10,000 birds/km<sup>2</sup>. No other
diving species is common at the ice front. The
parakeet auklet <em>(Cyclorhynchus psittaculus)</em>
is seen on most cruises, but only during a
small percentage of observation periods and
always in low numbers. Black guillemots are
common north of the ice front and stragglers
are occasionally seen at the front. Pigeon
guillemots <em>(Cepphus columbus)</em>, least auklets
<em>(Aethia pusilla)</em>, and crested auklets <em>(A. cristatella)</em>
are irregular visitors to the front.</p>

<p>Surface feeding species commonly found at
the ice front include the northern fulmar <em>(Fulmarus
glacialis)</em> and five species of gulls. The
fulmar is common south of the ice and is
found only in the southern portion of the
front. Three species of <em>Larus</em> are found at the
ice front. The most common is the glaucous-winged
gull <em>(Larus glaucescens)</em>; the glaucous
gull is less frequently seen. The slaty-backed
gull <em>(L. schistisagus)</em>, a species that
breeds in Asia, is most common west of St.
Matthew Island (McRoy et al. 1971). The
black-legged kittiwake <em>(Rissa tridactyla)</em> is
common in open water south of the ice but is
also found throughout the entire width of the
front. The ivory gull is unique in that it is
found only at the ice front in winter. In addition
to these species, the fork-tailed storm-petrel
<em>(Oceanodroma furcata)</em> is a regular but
uncommon visitor to the ice front in winter.
Densities of surface feeding species at the ice
front are low when compared to the high densities
of murres, and do not regularly exceed
<span class="pagenum"><a name="Page_13" id="Page_13">[Pg 13]</a></span>10 birds/km<sup>2</sup>.</p>

<p>The primary food consumed by birds at the
ice front is pollock <em>(Theragra chalcogramma)</em>.
An amphipod <em>(Parathemisto libellula)</em> and the
euphausiids are less important. Examination
of the stomach contents of birds and fish
show that large feeding flocks are usually associated
with schools of pollock feeding on <em>P.
libellula</em> and euphausiids.</p>

<p>The habitat of the consolidated pack in the
Bering Sea is markedly different from that at
the ice front. Whereas the front is characterized
by bands of ice interspersed with open
water and ice coverage rarely exceeding 4
oktas (4/8), the consolidated pack consists primarily
of large expanses of unbroken ice.
Small leads are formed by the shifting of the
ice caused by currents and wind. Ice coverage
is usually 7 to 8 oktas. The southern part of
the consolidated pack, which grades into the
ice front, has frequent leads. Most of the
species found at the ice front can be found in
the southern part of the consolidated pack,
but murres are most common. Their numbers
decrease, however, in the more northerly
pack, where leads are less frequent. Black
guillemots, in contrast, increase with increasing
ice cover, and reach their greatest abundance
in the small leads constantly forming
and refreezing deep within the ice. Because
they exploit this habitat, they are dependent
on the formation of lead systems. I have often
seen leads a quarter mile wide refrozen to the
point where new ice covered all but a small
patch of open water; black guillemots were
frequently crowded into this open water. Before
the lead closes completely the guillemots
must fly to an open lead. When winds are light
and temperatures low, lead systems fail to
form as rapidly as usual, and when they do
they refreeze quickly, causing a loss of the
preferred habitat of wintering black guillemots.
A severe winter in the White Sea in
1965-66 decreased the amount of open water
and caused an increased black guillemot mortality
(Bianchi and Karpovitsch 1969). On a
windless day in March I conducted bird observations
in the Bering Sea ice where no leads or
open water were encountered. The only bird
seen was a black guillemot flying over the ice.
In situations such as this, where black guillemots
are prospecting for open water, they
may use the "water sky" and steam fog associated
with leads as visual aids. "Water sky"
is the reflection of the dark water in the clouds
over the lead, and contrasts sharply with the
"ice sky." The presence of "water sky" allows
birds to detect open water from a distance of
many miles.</p>

<p>Aside from birds found in and near island-associated
polynias, only murres and black
guillemots are regularly found on the consolidated
pack ice in winter.</p>

<p>The polynia associated with islands in the
consolidated pack provide refuge(s) for seabirds.
Fay and Cade (1959) found the polynias
south of St. Lawrence to be most important to
oldsquaws <em>(Clangula hyemalis)</em>. King eiders
<em>(Somateria spectabilis)</em>, common eiders <em>(S.
mollisima)</em>, and oldsquaws are common in the
St. Matthew Island polynias (McRoy et al.
1971). Because these polynias are in shallow-water
areas, they provide feeding opportunities
for benthic feeding species.</p>


<h5><em>Spring</em></h5>


<h6>Chukchi and Beaufort Seas</h6>

<p>In April and May a lead system develops
from the Bering Strait north to Cape Lisburne
and then northeast to Point Barrow. The lead
is a flaw lead that occurs between the shorefast
ice and the free-floating pack. It is a
major migration route for a number of species
of birds, primarily eiders. East of Point Barrow
in the Beaufort Sea, no similar well-defined
large lead exists. Consequently, there is
a greater chance of bird mortality occurring in
the Beaufort Sea than in the Chukchi Sea because
the early migrants are unable to find
open water. In 1960, 10% of all the king eiders
that migrate through the Beaufort Sea died
during a late freeze (Barry 1968). Additional
records of eider mortality due to late breakup
or sudden freezes were presented by Palmer
(1976).</p>

<p><span class="pagenum"><a name="Page_14" id="Page_14">[Pg 14]</a></span></p>

<p>In late May, rivers that empty into the
northern Chukchi and Beaufort seas begin to
flow. The shorefast ice is still present at this
time and the rivers flow over the ice. For large
rivers, such as the Colville and the Sagavanirktok,
the area of ice covered by water is
considerable. Openings in the ice develop
sometime after the river runoff starts and the
river water drains through the ice.</p>

<p>This river overflow plays an important role
in the breeding biology of certain island nesting
species, since the overflow surrounds islands
and prevents arctic foxes from reaching
the islands. The overflow also allows birds to
sit in the water near breeding sites. It is not
known whether river overflow contains prey
items available to birds. After the overflow
drains through the ice, the shorefast ice that
has been covered with river overflow decomposes
quickly, and patches of open water
occur early in areas just seaward of major
river deltas. For this reason the largest breeding
colonies on barrier islands in the northern
Chukchi and Beaufort seas are all found near
the mouths of large rivers. Islands away from
rivers become isolated from the pack ice by
moats, which are caused by the absorption of
solar radiation by the islands and the melting
of the ice immediately adjacent to them. Moat
formation is not as predictable and uniform as
river overflow.</p>


<h6>Bering Sea</h6>

<p>When the ice in the Bering Sea begins to
melt in April, the edge of the pack does not recede
northward as is frequently thought.
Rather, there is a general decomposition of ice
throughout the pack. The leads that are constantly
forming in the ice no longer freeze. As
melt continues and ice becomes rotten, leads
form with increasing frequency. This manner
of ice decomposition is important to birds.
The leads that form deep in the pack ice provide
feeding and roosting areas near the large
seabird colonies found north of the ice edge,
and are used by certain tundra-nesting ocean
migrants such as eiders, red phalaropes
<em>(Phalaropus fulicarius)</em>, and jaegers (<em>Stercorarius</em>
spp.). If ice decomposition is retarded
by persistent low temperatures, the initiation
of breeding may be delayed at northern
Bering Sea colonies and for some tundra
species.</p>

<p>At the time of decomposition the large
standing stock of phytoplankton present in
the pack ice is released into the water. No information
is available on fish and invertebrate
populations that are associated with the decomposing
ice. The quantity of organic carbon
released is considerable, although it is not
known what fish or invertebrate populations
are supported by this plankton as soon as it is
released. For birds breeding in areas where ice
is present in the initial stages of breeding, the
phytoplankton released by the disintegrating
ice could play an important part in the birds'
energy budgets.</p>


<h5><em>Summer</em></h5>


<h6>Chukchi and Beaufort Seas</h6>

<p>In the northern Chukchi and Beaufort seas
the nearshore marine environment is dominated
by sea ice in June and July. In June the
coastal areas are characterized by a snow-free
tundra teeming with nesting waterfowl and
shorebirds next to an expanse of sea ice almost
completely devoid of bird life. In areas
where river outflow does not occur, the use of
nearshore waters usually begins when a moat
forms along the shoreline. Amphipods and
other invertebrates are found in this moat,
especially at stream mouths. Limited but
regular use of the moat occurs, primarily by
loons (<em>Gavia</em> spp.), oldsquaws, and arctic
terns <em>(Sterna paradisaea)</em>.</p>

<p>As the snow on top of the shorefast ice begins
to melt, ponds form on top of the ice. As
melt proceeds, these melt ponds merge into
long, parallel channels and may cover well
over 50% of the ice surface. Only when thaw
holes form and the melt ponds are connected
to the water under the ice is food present in
the channels. Amphipods are then seen swimming
in these channels. Bird use of these
channels is not extensive.</p>

<p><span class="pagenum"><a name="Page_15" id="Page_15">[Pg 15]</a></span></p>

<p>It is usually late July before the nearshore
ice begins its rapid decomposition. Ice in the
lagoons is the first to melt. Ice seaward of the
barrier islands decomposes more slowly because
of the presence of keels and pressure
ridges. As the ice melts, the in-ice algal bloom
is released into the water. These algae are important
because they provide at least 25 to
30% of the productivity in coastal waters and
allow the biological growing season to begin
before the open-water plankton bloom occurs
(Alexander 1974). In nearshore areas close to
Barrow, large populations of mysids and amphipods
are associated with the decomposing
ice. At least in certain areas, these ice-associated
zooplankton populations are a major
food source for nearshore migrants, especially
red phalaropes, arctic terns, and Sabine's
gulls <em>(Xema sabini)</em>.</p>

<p>The effects of ice scour on the shoreline and
the nearshore bottom of the Chukchi and
Beaufort seas is demonstrated by the absence
of sessile benthic fauna and flora. The effect
this absence has on birds is seen in the feeding
habits of nearshore birds. Oldsquaws and
eiders, which frequently feed on molluscs, feed
instead on motile benthos species such as
mysids, amphipods, and isopods. The emperor
goose <em>(Philacte canagica)</em> is absent from the
northern Chukchi and Beaufort seas, apparently
due to the absence of eelgrass beds.
Ice scour is the major cause of the absence of
eelgrass in northern Alaska (C. P. McRoy,
personal communication).</p>

<p>The offshore ice in the Chukchi decomposes
more rapidly than that in the Beaufort,
largely because Bering Sea water enters the
Chukchi through the Bering Strait (Coachman
and Barnes 1961). By late July the Chukchi
is usually ice free as far north as Icy Cape.
In the Beaufort, however, ice decomposition
occurs slowly through June and July, and
only in August does a definite strip of open
water develop between the shore and the edge
of the pack ice. The amount of open water
varies greatly from year to year. In certain
years the Beaufort is not navigable due to the
lack of open water.</p>

<p>Aerial censusing in June and July shows
that bird densities on the offshore ice are extremely
low. In August and September, when
shipboard censusing can be conducted, densities
on the pack ice in both seas are about 10
birds/km<sup>2</sup>. Unlike the Bering Sea, where densities
south of the ice are much less than on the
ice, bird densities south of the ice in the Beaufort
and Chukchi seas are slightly higher in
the open water south of the ice, averaging
about 20 birds/km<sup>2</sup>. In the Chukchi the principal
species encountered on the ice are the
black-legged kittiwake and the thick-billed
murre. In the Beaufort, red phalaropes, oldsquaws,
and glaucous gulls are the most common
species.</p>

<p>Numerous arctic cod are associated with
the underside of the summer pack ice. Shipboard
censusing in the ice is complicated
when cod are stranded on ice floes, as the ice
shifts under the weight of the ship. Gulls, arctic
terns, and jaegers gather behind the ship
to feed on these fish; mixed flocks of more
than 100 birds are common. In the absence of
a ship to provide the disturbance needed to
make large numbers of cod available, these
birds are dependent on locating the fish in the
surface waters next to ice floes. Because cod
frequently swim over underwater ice shelves
they are highly visible from above and should
be easily accessible to aerial feeders.</p>


<h5><em>Fall</em></h5>


<h6>Chukchi and Beaufort Seas</h6>

<p>By the time ice formation begins in late September
or early October, most seabirds have
left the Arctic on their southward migration.
The principal exception is the oldsquaw,
which does not begin its migration until September.
Some oldsquaws remain in nearshore
waters until they are driven out by the formation
of new ice. In contrast to the spring mortality,
there are few records of extensive bird
mortality in the fall due to lack of open water.
One instance was reported for 1975, when
nearshore waters froze early and flightless
eiders were seen sitting on the ice near Pt. Lay
in the Chukchi Sea. The birds were in a
weakened condition, apparently due to their
inability to obtain food (W. J. Wiseman, personal
communication).</p>

<p>In the offshore waters the species associated
with the pack ice in September are the
same as those in August. In late September,
however, ivory and Ross' gulls become the
most common species at the ice edge in the
Chukchi. Glaucous gulls and black guillemots
are also associated with the advancing ice
edge (Watson and Divoky 1972). Except for
the Ross' gull, which apparently winters in
the arctic basin, these species remain with the
ice as it advances into the Bering Sea.</p>

<p><span class="pagenum"><a name="Page_16" id="Page_16">[Pg 16]</a></span></p>


<h6>Bering Sea</h6>

<p>Little is known about bird distribution in
the Bering Sea during ice formation because
cruises in rapidly forming ice are potentially
hazardous. It is not known if the large numbers
of birds found at the ice edge in March
are present in December and January.</p>


<h4>Discussion</h4>

<p>The principal effect of the arctic pack ice is
to lower biological productivity and bird densities
in the areas it covers. Unlike the antarctic
pack ice, which supports a large biomass of
pagophilic species, the number of pagophilic
species supported by the arctic pack ice is
small. Only the ivory gull, Ross' gull, and
black guillemot have specific adaptations to
the ice environment. The Ross' gull and guillemot
winter in the pack ice, and the ivory gull
is associated with ice throughout the year.
The total biomass of these species is low.
Other species which are regularly associated
with the arctic pack, such as murres and
black-legged kittiwakes, are also found in
large numbers away from the ice. In addition,
these species are usually associated with ice
for limited periods during the year&mdash;murres
primarily in winter and spring and kittiwakes
primarily in summer.</p>

<p>The difference in the antarctic and arctic
pack ice systems is largely due to the antarctic
pack ice being surrounded by ocean, whereas
the arctic pack ice is, in general, surrounded
by land. The high productivity associated
with the antarctic pack ice is due primarily
to the mixing that occurs at the edge
of the pack ice. There is little opportunity for
mixing to occur next to the arctic pack ice, except
where it is next to large expanses of
boreal waters. This occurs in the Bering Sea in
winter and spring, in the North Atlantic, and
to a minor extent in the Chukchi Sea in summer
and fall (Dunbar 1968). The limited geographic
range and seasonal nature of high productivity
at the arctic pack ice edge has been
a major factor in preventing a well-developed
pagophilic avifauna.</p>

<p>The importance of the in-ice algal bloom and
its associated under-ice fauna is not yet clear.
It is probably most important in areas such as
the Beaufort Sea, where productivity in the
water column is low. Although considerable
numbers of seabirds are regularly found in the
summer pack ice feeding on arctic cod and
zooplankton associated with the ice, bird densities
south of the ice are usually greater than
those in the ice. The only species that appear
to depend on the ice-associated fauna for
much of their food are the three pagophilic
species mentioned above.</p>


<h4>References</h4>

<div class="hangindent">

<p>Alexander, V. 1974. Primary productivity regimes
of the nearshore Beaufort Sea, with reference to
potential roles of ice biota. Pages 609-632 <em>in</em> J. C.
Reed and J. E. Sater, eds. The coast and shelf of
the Beaufort Sea. Arctic Institute of North
America, Arlington, Virginia.</p>

<p>Andriashev, A. P. 1968. The problem of the life
community associated with the antarctic fast ice.
Pages 147-155 <em>in</em> R. I. Currie, ed. Symposium on
antarctic oceanography. Scott Polar Research Institute,
Cambridge.</p>

<p>Appollonio, S. 1961. The chlorophyll content of arctic
sea ice. Arctic 14:197-200.</p>

<p>Bailey, A. M. 1948. Birds of arctic Alaska. Colo.
Mus. Nat. Hist. Popular Ser. 8. 317 pp.</p>

<p>Barry, T. W. 1968. Observations on natural mortality
and native use of eider ducks along the
Beaufort Sea coast. Can. Field-Nat. 82(2): 140-144.</p>

<p>Bianchi, V. V., and V. N. Karpovitsch. 1969. The influence
of abnormal ice-cover of the White Sea
and Murman in 1966 upon birds and mammals.
(In Russian, English summary.) Zool. Zh.
48(6):871-875.</p>

<p>Bunt, J. S. 1963. Diatoms of antarctic sea ice as
agents of primary production. Nature (Lond.)
199:1255-1257.</p>

<p>Coachman, L. K., and C. A. Barnes. 1961. The contribution
of Bering Sea water to the Arctic
Ocean. Arctic 15:147-161.</p>

<p>Divoky, G. J. 1976. The pelagic feeding habits of
Ivory and Ross' Gulls. Condor 78:85-90.</p>

<p>Dunbar, M. J. 1968. Ecological development in
polar regions: a study in evolution. Prentice-Hall,
Englewood Cliffs, N.J. 119 pp.</p>

<p>Fay, F. H. 1974. The role of ice in the ecology of
marine mammals of the Bering Sea. Pages 383-399
<em>in</em> D. W. Hood and E. J. Kelly, eds. Oceanography
of the Bering Sea. Univ. Alaska Inst. Mar.
Sci. Occas. Publ. 2.</p>

<p>Fay, F. H., and T. J. Cade. 1959. An ecological
analysis of the avifauna of St. Lawrence Island,
Alaska. Univ. Calif. Publ. Zool. 63:73-150.</p>

<p>Gabrielson, I. N., and F. C. Lincoln. 1959. The birds
of Alaska. The Stackpole Company, Harrisburg,
Pennsylvania, and Wildlife Management Institute,
Washington, D.C. 922 pp.</p>

<p>Irving, L., C. P. McRoy, and J. J. Burns. 1970.
Birds observed during a cruise in the ice-covered
Bering Sea in March 1968. Condor 72:110-112.</p>

<p><span class="pagenum"><a name="Page_17" id="Page_17">[Pg 17]</a></span></p>

<p>Lisityn, A. P. 1969. Recent sedimentation in the
Bering Sea (Transl. from Russian.) Israel Program
for Scientific Translations, Jerusalem.
614 pp.</p>

<p>McRoy, C. P., and S. R. Goering. 1974. The influence
of ice on the primary productivity of the
Bering Sea. Pages 403-421 <em>in</em> D. W. Hood and
E. J. Kelly, eds. Oceanography of the Bering Sea.
Univ. Alaska Inst. Mar. Sci. Occas. Publ. 2.</p>

<p>McRoy, C. P., S. W. Stoker, G. E. Hall, and E.
Muktoyuk. 1971. Winter observations of mammals
and birds, St. Matthew Island. Arctic 24:63-65.</p>

<p>Mohr, J. L., and S. R. Geiger. 1968. Arctic Basin
faunal precis-animals taken mainly from arctic
drifting stations and their significance for biogeography
and water-mass recognition. Pages
297-313 <em>in</em> J. E. Sater, coordinator. Arctic drifting
stations. Arctic Institute of North America.</p>

<p>Nelson, R. K. 1969. Hunters of the northern ice.
University of Chicago Press, Chicago, Ill. 429 pp.</p>

<p>Palmer, R. S. 1976. Handbook of North American
birds. Vol. 3. Yale University Press, New Haven,
Conn. 560 pp.</p>

<p>Schmitt, W. L. 1965. Crustaceans. University of
Michigan Press, Ann Arbor. 204 pp.</p>

<p>Shapiro, L. H., and J. J. Burns. 1975. Major late-winter
features of ice in northern Bering and
Chukchi seas as determined from satellite
imagery. Univ. Alaska Geophys. Inst. Rep. 75-78.</p>

<p>Swartz, L. G. 1967. Distribution and movements of
birds in the Bering and Chukchi seas. Pacific Sci.
21:332-347.</p>

<p>Watson, G. E., and G. J. Divoky. 1972. Pelagic bird
and mammal observations in the eastern Chukchi
Sea, early fall 1970. U.S. Coast Guard Oceanogr.
Rep. 50:111-172.</p></div>

<p><span class="pagenum"><a name="Page_18" id="Page_18">[Pg 18]</a></span></p>


<div class="footnotes"><h4>FOOTNOTES:</h4>

<div class="footnote">

<p><a name="Footnote_1_1" id="Footnote_1_1"></a><a href="#FNanchor_1_1"><span class="label">[1]</span></a> Present address: Point Reyes Bird Observatory,
4990 State Route 1, Stinson Beach, California
94970.</p>

<p><span class="pagenum"><a name="Page_19" id="Page_19">[Pg 19]</a></span></p></div></div>

<p><span class="pagenum"><a name="Page_20" id="Page_20">[Pg 20]</a><br /><a name="Page_21" id="Page_21">[Pg 21]</a></span></p>
<div class="chapter"></div>
<hr class="chap" />




<h2>STATUS OF MARINE BIRD POPULATIONS</h2>
<div class="chapter"></div>
<hr class="chap" />




<h3><a name="Distribution_and_Status_of_Marine_Birds_Breeding" id="Distribution_and_Status_of_Marine_Birds_Breeding">Distribution and Status of Marine Birds Breeding
Along the Coasts of the Chukchi and Bering Seas</a></h3>


<p class="center">by</p>

<p class="center">James C. Bartonek<a name="FNanchor_2_2" id="FNanchor_2_2"></a><a href="#Footnote_2_2" class="fnanchor">[2]</a></p>

<p class="center">U.S. Fish and Wildlife Service<br />
Fairbanks, Alaska</p>

<p class="center">and</p>

<p class="center">Spencer G. Sealy</p>

<p class="center">University of Manitoba<br />
Winnipeg, Manitoba, Canada
</p>


<h4>Abstract</h4>

<div class="blockquot">

<p>The Alaska coast fronting on the Chukchi and Bering seas, exclusive of the
Aleutian Islands, supports seven complexes of marine bird colonies numbering
more than 1 million birds each, nine colonies of 100,000 to almost 1 million birds,
and many smaller colonies. Colonies are found on most headlands and islands
and are dominated numerically by alcids and kittiwakes (<em>Rissa</em> sp.). Estuarine
habitats (mainly the lowlands of northern Seward Peninsula, Yukon-Kuskokwim
delta, and the north side of the Alaska Peninsula) are extremely important for
breeding and migrating marine waterfowl, shorebirds, gulls (<em>Larus</em> sp.), and terns
(<em>Sterna</em> sp.). Information on population size and distribution of breeding marine
birds within this area is extensive for only a few of the more heavily hunted
species of waterfowl. Except for the intensive and systematic censusing of a few
colonies in this region, population data on cliff-, burrow-, and crevice-nesting
birds are such that all but gross changes in numbers may go unnoticed, and if
noticed they could not be measured.</p></div>

<p>Habitats for breeding marine birds are
found along much of the 4,100-km coastline of
Alaska that fronts on the Chukchi and Bering
seas. Seasonal sea ice and an extensive outer
continental shelf are dominant features that
contribute to the productivity of these marine
waters, which sustain populations of fishes,
birds, and mammals that are of considerable
and diverse values to man (Kelley and Hood
1974).</p>

<p>Our purpose in this paper is to describe the
distribution, abundance, and relative status
of some of the nearly 100 species of marine
birds breeding within this region and the information
base from which the descriptions
are derived. Although the selection is admittedly
arbitrary, we discuss mainly the
colonial nesting species because they are
generally in greater jeopardy from lost breeding
habitat and from catastrophes than are
the species that are widely dispersed or solitary
in nesting. Because we believe matters
affecting the conservation of marine birds will
be geographically oriented, we discuss the
status and distribution of breeding birds on
that basis, rather than by the more traditional
taxonomic approach. We use the terms
"colony" and "colonies" somewhat loosely
and interchangeably to include any aggregation
of birds of the same or different species
nesting in proximity to each other, even those
on the same island or headland, although
populations may be miles apart and occupy
different kinds of habitats. The nature of this
paper and the scale of our maps do not allow
for detailed resolution of each colony's location
<span class="pagenum"><a name="Page_22" id="Page_22">[Pg 22]</a></span>(for the most part this information is not
available), but rather facilitates a general impression
of status.</p>

<p>Most place-names used by us are shown in
Fig. 1; the others may be located by referring
to Orth's (1967) gazetteer on Alaska.</p>


<h4>Information Base</h4>

<p>There is no adequate catalog of marine bird
colonies and other avian habitats for the
Bering-Chukchi region or for Alaska as a
whole. King and Lensink (1971) described the
waterfowl populations and major lowland
habitats of the State and listed only a few of
the many colonies of cliff-nesting birds.
LeResche and Hinman (1973) identified a few
additional colonies, provided fragmentary information
on composition and abundance at
some of these sites, and delineated areas of
wetland habitats on maps in their statewide
atlas on wildlife. General and occasionally
site-specific information on the location, but
rarely on population size and composition, of
colonies can be gleaned from the 321 species
accounts presented by Gabrielson and Lincoln
(1959) and from the general works by
Bent (1919, 1921, 1922, 1923, 1925, 1927,
1929), Dement'ev and Gladkov (1951),
Dement'ev et al. (1951, 1952), Palmer (1962),
Fisher (1952), Tuck (1960), and others. The
birds on the Asiatic side of these waters,
which are not treated in this paper, were described
by Portenko (1973).</p>

<p>Information on the status of waterfowl in
the region is generally more detailed than that
for most other groups of birds because waterfowl
have been the object of systematic surveys
since the late 1940's as part of the continent-wide
effort to manage populations for
sport hunting. Because the emphasis of these
surveys has been directed toward the species
of ducks important to hunters in the "lower
48" States, data are not adequate to measure
changes in populations for most sea ducks
and marine geese nesting in this region. These
surveys have, however, enabled biologists to
delineate waterfowl habitats and make reasonable
estimates of populations for some of
the more abundant and conspicuous species
(King and Lensink 1971; U.S. Fish and Wildlife
Service [FWS] 1973<em>c</em>; U.S. National Park
Service [NPS] 1973).</p>


<h5><em>Chukchi Sea Coast</em></h5>

<p>A disproportionate percentage of ornithological
investigations in arctic Alaska have
centered about Barrow, where ornithologists
were attracted because of the propensity of
vagrant birds to collect there and because of
the above average facilities, conveniences,
and transportation afforded first by the whaling
station, then by the military, and later by
a research laboratory. Recent petroleum development
near Prudhoe Bay has resulted in a
somewhat commensal eastward shift in ornithological
studies.</p>

<p>Bailey (1948), Gabrielson and Lincoln
(1959), and Pitelka (1974) reviewed much of
the published information on arctic avifauna,
including that of the Chukchi coast. Selkregg
[1975] mapped various avian habitats,
ascribed either relative or absolute values for
the population size of certain groups of birds,
and included a selected bibliography that did
not entirely duplicate those provided by the
other reviewers. Watson and Divoky (1975)
described the avifauna of Alaska's Beaufort
Sea coast, which is much the same as that of
the Chukchi coast from Point Barrow south to
Cape Lisburne (both coasts are of low relief).</p>

<p>Intensive studies near Barrow have done
much to characterize the behavior, productivity,
and ecological requirements of calidridine
sandpipers (Pitelka 1959; Pitelka et al.
1974; Holmes 1970, 1971) and, to partly explain
the cyclical relationships between
jaegers (<em>Stercorarius</em> spp.) and their prey (e.g.,
Pitelka et al. 1955; Maher 1974). Quantitative
estimates of certain bird populations at Cape
Thompson (Swartz 1966; Williamson et al.
1966), Little Diomede (Kenyon and Brooks
1960), and on the coastal lowlands of the
Seward Peninsula (King and Lensink 1971;
U.S. NPS 1973), and for black guillemots
<em>(Cepphus grylle)</em> throughout the region
(Divoky et al. 1974) are among the best data
on status of marine birds for any locality in
Alaska. Grinnell (1900<em>a</em>) described the birds
he observed in the Kotzebue Sound area.</p>

<p><span class="pagenum"><a name="Page_23" id="Page_23">[Pg 23]</a></span></p>

<div class="figcenter" style="width: 517px;">
<img src="images/p_023.jpg" width="517" height="700" alt="" />
<div class="caption"><p>Fig. 1. Place-names in the region of the Chukchi and Bering seas.</p></div>
</div>

<p><span class="pagenum"><a name="Page_24" id="Page_24">[Pg 24]</a></span></p>

<p>Cursory aerial surveys conducted by J. C.
Bartonek, J. G. King, and D. R. Cline (U.S.
FWS 1973<em>a</em>; U.S. NPS 1973; this paper) in
1972 and 1973 provided information on the
location and relative size of most, if not all,
colonies of cliff-nesting marine birds between
Point Barrow and the Bering Strait, including
those at Cape Lisburne, at Motherhood Point,
Nine-mile Point, Cape Deceit, Towalevic
Point, Sullivan Bluff, all on the northern base
of the Seward Peninsula, and at Fairway
Rock. The relative size of populations of most
species was probably underestimated because
the burrow-and crevice-nesting species were
largely unseen.</p>


<h5><em>Bering Sea</em></h5>

<p>Aside from work by Gabrielson and Lincoln
(1959) and the early but understandably incomplete
accounts by Nelson (1883, 1887) and
Turner (1886), no comprehensive description
of the avifauna of the Alaskan coast of the
Bering Sea exists. Many studies adequately
describe local avifauna, and some of them are
exemplary assessments of the status of
populations.</p>

<p>Most of the coastline suitable for cliff-nesting
marine birds and most of the smaller nearshore
islands from the Bering Strait south to
the tip of the Alaska Peninsula were reconnoitered
piecemeal from aircraft between 1970
and 1973 by J. C. Bartonek, J. G. King, D. R.
Cline, C. D. Evans, and M. L. Plenert (U.S.
FWS 1973<em>a</em>, 1973<em>b</em>; this paper). In late June
1973 Bartonek, Cline, and Plenert made brief
reconnaissances on foot of King, Besboro, and
Shaiak islands. Bartonek and J. G. Divoky,
traveling by boat and occasionally on foot, reconnoitered
colonies at Cape Seniavin, a portion
of the Walrus Islands group, Shaiak
Island, and the coastline from Cape Peirce
around Cape Newenham to Security Cove
(U.S. FWS 1973<em>a</em>, 1973<em>b</em>; this paper). Although
these cursory surveys (especially
those from aircraft) tended to identify nesting
sites of cliff-nesting birds while missing sites
used by burrow-and crevice-nesting species,
information was obtained on the location and
relative size of many previously unreported
colonies.</p>

<p>The mainland and island colonies in Norton
Sound have received little notice in the published
literature. Bailey (1943, 1948), although
working mainly at Little Diomede and in Arctic
and Lopp lagoons on the north side of the
Seward Peninsula, mentioned the birds at
Wales Mountain and Tin City. Nelson (1883,
1887) traveled throughout the region studying
the avifauna and the anthropology of
Eskimos. Grinnell (1900<em>b</em>) at Nome,
McGregor (1902) along the Koyuk River,
Hersey (1917) and Turner (1886) near St.
Michael, and Cade (1952) at Sledge Island provide
fragmentary examples of the area's
marine bird populations. Colonies at King,
Besboro, Egg, and Sledge islands, near York
Mountains, and at Bluff were described in
proposals for new National Wildlife Refuges
(U.S. FWS 1973<em>a</em>).</p>

<p>Sealy et al. (1971) reviewed the literature
and discussed the various zoogeographic relationships
among the avifauna of St. Lawrence
Island. Fay and Cade (1959) estimated numbers
and biomass of all birds on St. Lawrence
Island but did not identify locations and sizes
of particular populations; consequently, replication
of their estimates is precluded. An
exemplary study by B&eacute;dard (1969) identified
the locations and sizes of all populations of
crested auklets <em>(Aethia cristatella)</em>, least auklets
<em>(A. pusilla)</em>, and parakeet auklets <em>(Cyclorrhynchus
psittacula)</em> on the island. Sealy
(1973) identified breeding sites of horned
puffins <em>(Fratercula corniculata)</em> there and
throughout the species' range. Thompson
(1967) listed the birds observed at Northeast
Cape and on nearby Punuk Islands.</p>

<p>Annotated accounts have been published on
the breeding avifauna of St. Matthew, Hall,
and Pinnacle islands by Elliott (1882), Hanna
(1917), Bent (1919), and Gabrielson and
Lincoln (1959). Klein (1959) presented quantitative
data on the birds he observed incidental
to his study of reindeer <em>(Rangifer
tarandus)</em>.</p>

<p>The avifauna of the Yukon-Kuskokwim
delta, which is rich both in numbers and diversity,
has been treated extensively in the literature.
Nelson (1883, 1887), Turner (1886), Conover
(1926), Brandt (1943), Gabrielson and
Lincoln (1959), Williamson (1957), Kessel et
al. (1964), Harris (1966), Dau (1972), and
Holmes and Black (1973) all described the avifauna
in the same general area of the delta,
i.e., the eroding portion in the general vicinity
of Hooper and Hazen bays. The avifauna
of the aggrading portion of the Yukon
delta and of the Kuskokwim's mouth have not
been accorded similar attention. Populations
of waterfowl nesting on the delta and their
wintering affinities were described by King
and Lensink (1971) and U.S. FWS (1973<em>c</em>).</p>

<p><span class="pagenum"><a name="Page_25" id="Page_25">[Pg 25]</a></span></p>

<p>Studies of particular species of marine birds
on the delta (again, all in the general vicinity
of Hooper and Hazen bays) were reported by
Hansen and Nelson (1957) and Shepherd
(1960) for black brant <em>(Branta bernicla)</em>, by
Headley (1967) and Eisenhauer and Kirkpatrick
(1977) for emperor geese <em>(Anser canagica)</em>,
by Dau (1974) and Mickelson (1975) for
spectacled eiders <em>(Somateria fischeri)</em>, by
Petersen (1976) for red-throated loons <em>(Gavia
stellata)</em>, and by Holmes (1970, 1971, 1972) for
dunlins <em>(Calidris alpina)</em> and western sandpipers
<em>(C. mauri)</em>.</p>

<p>Birds of Nunivak Island were reported by
Swarth (1934), but the importance of the
island to marine birds was not put into proper
perspective until the Nunivak National Wildlife
Refuge was evaluated for designation as a
wilderness area (U.S. FWS 1972).</p>

<p>The Pribilof Islands have served as a focal
point for ornithological investigations of the
Bering Sea in much the same way that Barrow
has for the Arctic. The avifauna of the
Pribilofs has been described by Coues (1874),
Elliott (1882), Palmer (1899), Hanna (1918),
Preble and McAtee (1923), Gabrielson and
Lincoln (1959), Kenyon and Phillips (1965),
and a host of others that mainly added new
species to the record list. Although most of
these ornithologists marveled at the numbers
of birds, information is lacking from which
most changes in populations can be noted.
(An exception is the record of common and
thick-billed murres, <em>Uria aalge</em> and <em>U. lomvia</em>,
which formerly nested in such abundance on
Walrus Island that annually several tons of
eggs were gathered for consumption by residents
of the islands [Palmer 1899], but were
greatly reduced in numbers by the summer of
1973, when J. C. Bartonek, J. G. King, G. J.
Divoky, and D. T. Montgomery observed only
a few thousand murres on a small portion of
the island. Most of the suitable nesting sites,
especially the flat areas often used by common
murres, were occupied by Steller's sea
lions, <em>Eumetopias jubata</em>, which, apparently
because of reduced hunting pressure, occupied
the island and displaced the murres.)</p>

<p>For some unexplained reason the numerous
and large marine bird colonies along the north
side of Bristol Bay appear to have been
largely overlooked until recent years (Bartonek
and Gibson 1972). Gabrielson and
Lincoln (1959) summarized the few observations
by Osgood (1904) and Turner (1886) in
this area, but obviously were unaware that, in
aggregate, these colonies rival those of the
Pribilofs. Dick and Dick (1971) made an
exemplary study of marine birds and their
numbers at Cape Peirce and on nearby Shaiak
Island. Murie (1959) provided annotated remarks
on marine birds of Amak Island, but
not of nearby Sealion Rocks.</p>


<h4>Status and Distribution</h4>

<p>Seven groups of colonies of cliff-, burrow-,
and crevice-nesting birds are found on the
headlands and islands in the coastal region,
each numbering more than 1 million birds;
nine colonies range downward to 100,000
birds; and a host of others range downward to
1,000 birds (Fig. 2). Un-estimated numbers of
other marine birds nest on the lowlands about
Kotzebue Sound, the Yukon-Kuskokwim
delta, and Bristol Bay, but are not shown in
Fig. 2. The occurrence at colonies of 20 of the
nearly 100 species of marine birds is shown in
Fig. 3; their relative numbers at these sites
are not shown because data are generally
lacking.</p>

<div class="figcenter" style="width: 522px;">
<img src="images/p_025.jpg" width="522" height="700" alt="" />
<div class="caption"><p>Fig. 2. Relative numbers of marine birds at colonies
in different localities, without regard to species
composition or breeding status.</p></div>
</div>

<p><span class="pagenum"><a name="Page_26" id="Page_26">[Pg 26]</a></span></p>

<div class="figcenter" style="width: 503px;">
<img src="images/p_026.jpg" width="503" height="700" alt="" />
<div class="caption"><p>Fig. 3. Location of known breeding populations of some marine bird species without regard to size of
population.</p></div>
</div>

<p><span class="pagenum"><a name="Page_27" id="Page_27">[Pg 27]</a></span></p>

<div class="figcenter" style="width: 679px;">
<img src="images/p_027.jpg" width="679" height="633" alt="" />
</div>


<h5><em>Chukchi Sea</em></h5>

<p>The largest colonies of seabirds in the Chukchi
Sea are those on Little Diomede Island,
Cape Lisburne, Cape Thompson, and Fairway
Rock. Smaller colonies are in Kotzebue Sound
along the northern base of the Seward Peninsula.
These colonies are largely dominated by
thick-billed and common murres and black-legged
kittiwakes <em>(Rissa tridactyla)</em> and on
the islands in the Bering Strait also the
crested, least, and parakeet auklets. Horned
puffins, tufted puffins <em>(Lunda cirrhata)</em>,
pelagic cormorants <em>(Phalacrocorax pelagicus)</em>,
and glaucous gulls <em>(Larus hyperboreus)</em> make
up the remaining majority. For the whole area
there are probably fewer than a hundred birds
each of black guillemots and pigeon guillemots
<em>(Cepphus columba)</em> occupying colonies.
Dovekies <em>(Alle alle)</em> are occasionally sighted in
this area, but only as stragglers from their
normal range.</p>

<p>Part of the mystery surrounding the nesting
location of Kittlitz's murrelet <em>(Brachyramphus
brevirostris)</em> was solved when
Thompson et al. (1966) discovered a downy
chick in the Kukpuk River drainage nearly
45 km by river from salt water. Other nesting
sites of the Kittlitz's murrelets in this region<span class="pagenum"><a name="Page_28" id="Page_28">[Pg 28]</a></span>
were reported for Wales Mountain (Ford
1936; Bailey 1943, 1948) and the Cold Bay
area (Bailey 1973) (Fig. 3).</p>

<p>Only the colonies at Cape Thompson have
been censused systematically throughout a
breeding season. During one of three years of
varying census efforts, Swartz (1966) estimated
that about 400,000 birds of nine
species occupied the cliffs. Whereas the Cape
Thompson colonies received considerable attention
because of Swartz's efforts, the colonies
that extend along nearly 35 km of headlands
southward from, but mainly at, Cape
Lisburne have received little if any attention
by either early or recent ornithologists in the
Arctic, even though they support perhaps
twice the number of birds. Also perplexing is
why Chamisso and Puffin islands with their
several thousand nesting horned puffins and
lesser numbers of other seabirds were designated
as the Chamisso National Wildlife
Refuge in the early 1900's when none of the
many larger and more species-diverse colonies
in the area received comparable recognition
by and protection through refuge designation.</p>

<p>The lowlands on the north side of the
Seward Peninsula produce fall flights of sea
ducks that average 49,200 oldsquaws <em>(Clangula
hyemalis)</em>, 51,000 eiders (mostly common
eiders, <em>Somateria mollissima</em>), and 26,700
scoters (mostly black scoters, <em>Melanitta
nigra</em>) (King and Lensink 1971). Small populations
of black brant and emperor geese breed
in what outwardly appears to be excellent
habitat, and King and Lensink (1971) speculated
that subsistence hunting by local
Eskimos is responsible for suppressing these
populations.</p>


<h5><em>Bering Sea</em></h5>

<p>The largest concentration of nesting seabirds
in the Bering Sea and perhaps in the entire
North Pacific is that on St. George
Island. Colonies that rank somewhere below
that at St. George are along the coast from
Cape Newenham to Cape Peirce, in the Walrus
Islands (Round, High, Crooked, and Summit
islands, The Twins, and Black Rock), at Cape
Mohican on Nunivak Island, St. Matthew
Island, Southwest Cape of St. Lawrence
Island, and King Island.</p>

<p>The Pribilofs have the unique distinction of
being the primary nesting site of red-legged
kittiwakes <em>(Rissa brevirostris)</em>. They are also
interesting from the zoogeographic standpoint
in that they are the northernmost
stronghold of red-faced cormorants <em>(Phalacrocorax
urile)</em>; guillemots are conspicuous by
their absence, and larid gulls are conspicuously
scarce nesters.</p>

<p>St. Matthew Island and associated Hall and
Pinnacle islands, and all but Walrus Island of
the Pribilofs, are sites of nesting northern fulmars
<em>(Fulmarus glacialis)</em>. Nesting fork-tailed
or Leach's storm-petrels (<em>Oceanodroma furcata</em>
and <em>O. leucorhoa</em>) have been found nowhere
in this region, although both are commonly
observed at sea and both nest throughout
the Aleutians.</p>

<p>Most colony sites identified in Fig. 2 are
dominated by common or thick-billed murres
(or both) and black-legged kittiwakes. Glaucous
gulls (generally north of the Yukon-Kuskokwim
delta), glaucous-winged gulls
<em>(Larus glaucescens)</em> (generally to the south of
the delta), and pelagic cormorants occupy almost
every rocky prominence along the entire
coast (most of these sites are not shown in
Figs. 2 and 3). Double-crested cormorants
<em>(Phalacrocorax auritus)</em> nest at a few island
and inland locations in the Bristol Bay area.
The small auklets are largely restricted to
islands in the Bering Sea; the parakeet auklet
is the only one occasionally found in mainland
colonies.</p>

<p>The marine birds of the Yukon-Kuskokwim
delta lowlands, although largely uncounted,
in their aggregate probably exceed the numbers
at any individual site identified in Fig. 2.
This is not particularly surprising since the
delta has nearly 70,000 km<sup>2</sup> of habitat (King
and Lensink 1971) in contrast to the generally
small parcels of habitat occupied at the sea-cliff
and island sites.</p>

<p>King and Lensink (1971) estimated that fall
flights of sea ducks originating on the delta
averaged 292,300 oldsquaws, 51,000 eiders
(mostly common and spectacled eiders with
lesser numbers of Steller's eiders, <em>Polysticta
stelleri</em>), and 157,000 scoters (primarily black
scoters). They also estimated that half of the
150,000 black brant and most of the 150,000
emperor geese in Alaska's fall flight originate
there. Although no counts have been made,
we believe that the delta's lowlands support
easily more than half of Alaska's nesting dun<span class="pagenum"><a name="Page_29" id="Page_29">[Pg 29]</a></span>lins,
black turnstones <em>(Arenaria melanocephala)</em>,
rock sandpipers <em>(Calidris ptilocnemis)</em>,
western sandpipers, and substantial
percentages of red phalaropes <em>(Phalaropus
fulicarius)</em>, northern phalaropes <em>(Lobipes
lobatus)</em>, and red-throated loons.</p>

<p>The north side of the Alaska Peninsula (including
the wetlands, uplands, and estuaries)
is perhaps more important to marine birds as
a staging, feeding, and resting area than as a
nesting habitat. The importance of Izembek
Lagoon to black brant and emperor geese during
fall and spring is a classic example. King
and Lensink (1971) estimated that the fall
flight of sea ducks originating from the Peninsula
averages 53,400 oldsquaws, 1,700 eiders,
and 74,400 scoters. Breeding geese are scarce
throughout the area.</p>


<h4>Conclusions and
Recommendations</h4>

<p>Most of the major breeding habitats of
marine birds in the Chukchi and Bering seas
are known, but imprecisely identified as to
location and size. With few exceptions, the
populations of birds using these habitats are
described only by the subjective and ambiguous
descriptors of abundance such as
"abundant, common, occasional, and rare,"
which makes measurement of change impossible.</p>

<p>We recommend that first and foremost a
catalog of habitats used by birds be developed
to aid resource administrators, developers,
and biologists (all of whom should be "conservationists")
in identifying critical habitats.
We believe that such a catalog would preclude
many problems because birds and their habitats
could be considered at the planning stage
rather than only at the operational stage.
Such a catalog would also be useful to students
of ornithology who are seeking locations
suitable for particular studies.</p>

<p>Nowhere in this region have studies of
marine birds been of sufficient duration to enable
changes in populations (from whatever
cause) to be characterized. Since some species
of marine bird are known not to breed before
at least 3 or more years of age, meaningful information
on survival and recruitment in
populations cannot be obtained by studies of
less than 10 years. We therefore recommend
that long-term studies be initiated at as many
places as possible, but at least at one site on
the Yukon-Kuskokwim delta; at a mainland
colony site that has predominantly murres,
kittiwakes, puffins, and cormorants; and at
an island site that also has small auklets. Although
the nesting distribution of the Kittlitz's
murrelet remains an enigma, we regard
it less of a conservation issue and more of an
ornithological challenge. Consideration of
logistics and support facilities must, of
course, be included in the site selection process.
Most of the areas suggested for these
studies also merit recognition and protection
by being designated as a National Wildlife
Refuge, a National Park or Monument, or a
State Game Sanctuary.</p>


<h4>References</h4>

<div class="hangindent">

<p>Bailey, A. M. 1943. The birds of Cape Prince of
Wales, Alaska. Proc. Colo. Mus. Nat. Hist. 18.
113 pp.</p>

<p>Bailey, A. M. 1948. Birds of arctic Alaska. Colo.
Mus. Nat. Hist. Popular Ser. 8. 317 pp.</p>

<p>Bailey, E. P. 1973. Discovery of a Kittlitz's murrelet
nest. Condor 75(4):457.</p>

<p>Bartonek, J. C., and D. D Gibson. 1972. Summer
distribution of pelagic birds in Bristol Bay,
Alaska. Condor 74(4):416-422.</p>

<p>B&eacute;dard, J. 1969. The nesting of the least, crested,
and parakeet auklets around St. Lawrence
Island, Alaska. Can. J. Zool. 47(5):1025-1050.</p>

<p>Bent, A. C. 1919. Life histories of North American
diving birds. U.S. Natl. Mus. Bull. 107. 245 pp.</p>

<p>Bent, A. C. 1921. Life histories of North American
gulls and terns. U.S. Natl. Mus. Bull. 113.
345 pp.</p>

<p>Bent, A. C. 1922. Life histories of North American
petrels, pelicans, and their allies. U.S. Natl. Mus.
Bull. 121. 343 pp.</p>

<p>Bent, A. C. 1923. Life histories of North American
wild fowl. Part I. U.S. Natl. Mus. Bull. 126.
250 pp.</p>

<p>Bent, A. C. 1925. Life histories of North American
wild fowl. Part II. U.S. Natl. Mus. Bull. 130.
311 pp.</p>

<p>Bent, A. C. 1927. Life histories of North American
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<p>Holmes, R. T. 1972. Ecological factors influencing
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<p>Pitelka, F. A. 1959. Numbers, breeding schedules,
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<p>Pitelka, F. A. 1974. An avifaunal review for the Barrow
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<p>Pitelka, F. A., R. T. Holmes, and S. F. MacLean, Jr.
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<p>Pitelka, F. A., P. Q. Tomich, and G. W. Treichel.
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<p>Portenko, L. A. 1973. The birds of the Chukotsk
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<p>Sealy, S. G. 1973. Breeding biology of the horned
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<p>Sealy, S. G., F. H. Fay, J. B&eacute;dard, and M. D. F.
Udvardy. 1971. New records and zoogeographical
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<p>Selkregg, L. L., ed. [1975]. Alaska regional profiles:
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<p>Swarth, H. S. 1934. Birds on Nunivak Island,
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<p><span class="pagenum"><a name="Page_32" id="Page_32">[Pg 32]</a></span></p>


<div class="footnotes"><h4>FOOTNOTES:</h4>

<div class="footnote">

<p><a name="Footnote_2_2" id="Footnote_2_2"></a><a href="#FNanchor_2_2"><span class="label">[2]</span></a> Present address: U.S. Fish and Wildlife Service,
500 NE Multnomah Street, Portland, Oregon
97232.</p>

<p><span class="pagenum"><a name="Page_33" id="Page_33">[Pg 33]</a></span></p></div></div>
<div class="chapter"></div>
<hr class="chap" />




<h3><a name="Breeding_Distribution_and_Status_of_Marine_Birds" id="Breeding_Distribution_and_Status_of_Marine_Birds">Breeding Distribution and Status of Marine Birds
in the Aleutian Islands, Alaska</a></h3>


<p class="center">by<br />
Palmer C. Sekora<a name="FNanchor_3_3" id="FNanchor_3_3"></a><a href="#Footnote_3_3" class="fnanchor">[3]</a></p>

<p class="center">U.S. Fish and Wildlife Service<br />
Kailua, Hawaii</p>

<p class="center">G. Vernon Byrd<a name="FNanchor_4_4" id="FNanchor_4_4"></a><a href="#Footnote_4_4" class="fnanchor">[4]</a></p>

<p class="center">U.S. Fish and Wildlife Service<br />
Adak, Alaska</p>

<p class="center">and</p>

<p class="center">Daniel D. Gibson</p>

<p class="center"><em>University of Alaska<br />
Fairbanks, Alaska</em>
</p>


<h4>Abstract</h4>

<div class="blockquot">

<p>Seabird population estimates are generally lacking for the 1,800-km-long
Aleutian Islands. Only the locations of the larger colonies are known, and for
these there are only imprecise estimates of colony sizes and often even of species
composition. Changes in the status of several species and populations resulting
from geologic and marine actions and from human intrusions are evident. Accounts
are given for 25 species of marine birds breeding in these islands.</p></div>

<p>The 1,800-km-long chain of islands known
as the Aleutians provides nesting habitat for
various species of marine birds, including
three species of Procellariiformes and three of
cormorants (<em>Phalacrocorax</em> spp.), one species
of gull <em>(Larus glaucescens)</em>, both kittiwake
species (<em>Rissa</em> spp.), two species of terns
(<em>Sterna</em> spp.), and at least 13 species of alcids.</p>

<p>Seabird population estimates of known accuracy
are lacking for this isolated area. Locations
of larger colonies of breeding seabirds
are known, however, and sufficient data are
available to place colonies in broad size
ranges. Published information on the breeding
biology of marine birds is also lacking
from the Aleutians, but some studies are
under way. The distribution of nesting marine
birds away from the nesting cliffs is totally
unknown.</p>

<p>Introduced predators, primarily arctic
foxes <em>(Alopex lagopus)</em>, are now found on
nearly every island. Breeding marine bird
populations have suffered drastic reductions
as a result. They have probably also changed
because of natural habitat modifications
caused by earthquakes, volcanic eruptions,
tidal waves, and marine erosion.</p>

<p>The purpose of this paper is to summarize
the known present distribution and status of
breeding marine birds in the Aleutian Islands.</p>

<p><span class="pagenum"><a name="Page_34" id="Page_34">[Pg 34]</a></span></p>
<p>Description of the
Aleutian Islands</p>

<p>The Aleutian Islands form an arc that separates
the Bering Sea and the north Pacific
Ocean (Fig. 1). The island chain extends from
the tip of the Alaska Peninsula to within
483 km of the Commander Islands of Siberia.
The chain contains more than 200 islands&mdash;the
peaks of a submarine volcanic mountain
range. Volcanic activity and earthquakes occur
regularly.</p>

<p>Weather is characterized by perpetual overcast,
dense summer fog, high-velocity winds,
and mild temperatures with low annual and
diurnal variations. The sea is ice-free year-round
except in extremely cold winters, when
the arctic ice pack may reach the extreme
northern islands.</p>

<p>The Aleutians are treeless except for a few
introduced, stunted spruces. Woody shrubs
are restricted to the most northern islands on
each end of the Chain. Mosses, lichens, club
mosses, and heaths are common ground-cover
plants, and taller grasses, sedges, and umbellifers
constitute the overstory. Hulten (1960)
provided a list of terrestrial plants found in
the Aleutians. Amundsen and Clebsch (1971)
discussed terrestrial plant ecology at Amchitka,
central Aleutians. The marine plant
communities around the islands are fairly diverse.
Lebednik et al. (1971) described marine
algal communities at Amchitka.</p>

<p>The easternmost Aleutian island, Unimak,
has a mammalian fauna like that of the
Alaska Peninsula, including brown bear
<em>(Ursus arctos)</em>, caribou <em>(Rangifer tarandus)</em>,
wolf <em>(Canis lupus)</em>, and wolverine <em>(Gulo gulo)</em>.
West of Unimak, red foxes <em>(Vulpes fulva)</em> occurred
historically as far as Umnak, and arctic
foxes were apparently on Attu when the
Russians came in 1741 (Murie 1959). Except
for man and dog, no land mammals occurred
between Umnak and Aggatu islands. Arctic
foxes, introduced before 1930 for fur farming,
still roam almost every island. Norway rats
<em>(Rattus norvegicus)</em> were introduced on many
islands when ships were wrecked or as a result
of military activities during World War II.</p>

<p>Sea otters <em>(Enhydra lutris)</em> have repopulated
most of the Aleutians after being nearly
extirpated by 1900. Rookeries of Steller's sea
lion <em>(Eumetopias jubata)</em> are scattered
throughout the Aleutians during summer,
and numerous harbor seals <em>(Phoca vitulina)</em>
haul out on beaches and offshore rocks.</p>

<p>All five species of Pacific salmon (<em>Oncorhynchus</em>
spp.) occur near the islands, and at least
four of them (all but <em>O. tshawytscha</em>) spawn in
Aleutian streams. Dolly Varden <em>(Salvelinus
malma)</em> and three-spine sticklebacks <em>(Gasterosteus
aculeatus)</em> are found nearly everywhere
there is fresh water. The marine environment
provides habitat used by at least 77
species of fish (Isakson et al. 1971). O'Clair
and Chew (1971) furnished a recent reference
to littoral macrofauna at Amchitka.</p>

<p>About 200 species of birds have been recorded
in the Aleutians (Aleutian Islands National
Wildlife Refuge, unpublished data).
Many of these are windblown stragglers from
both North America and Asia; only 59 species
breed on the islands. Although seabirds make
up less than half (26 species or 44%) of the
breeding birds, they may compose more than
90% of the breeding avian biomass.</p>


<h4>Ornithological Investigations
in the Aleutians</h4>

<p>Published ornithological information from
the Aleutian Islands is relatively scarce.
G. W. Steller, naturalist on Vitus Bering's
1741 expedition to Alaska, was the first person
to record ornithological information in the
islands (Stejneger 1936). More than a century
passed before W. H. Dall (1873, 1874) published
the next papers dealing with birds in
the Aleutians. In 1878, the U.S. Army Signal
Corps sent L. M. Turner to the Aleutians to
set up weather stations at several locations.
Turner kept notes on birds at various locations
in the Aleutians and published two
papers (1885, 1886) on his observations.
Turner's data (1886) provided the first report
based on extended and widespread observations
in the area. E. W. Nelson, who replaced
Turner, also provided data on birds (Nelson
1887).</p>

<p>In 1906, A. C. Bent came to the Aleutians
specifically to look for birds, and he and
Alexander Wetmore recorded birds throughout
the island chain (Bent 1912). A. H. Clark<span class="pagenum"><a name="Page_35" id="Page_35">[Pg 35]</a></span>
(1910) provided a valuable record of his observations
in the Near Islands. All these workers
recorded birds in several locations, but none
provided data on more than a very few seabird
colonies.</p>

<div class="figcenter" style="width: 700px;">
<img src="images/p_035.jpg" width="700" height="227" alt="" />
<div class="caption"><p><strong>Fig. 1.</strong> The Aleutian Islands.</p></div>
</div>

<p>O. J. Murie, U.S. Biological Survey, made
the most complete survey of the Aleutians
(Murie 1959). He specifically recorded seabird
colonies, spending parts of four summers in
the area. Murie visited every large Aleutian
island and most small ones. He recorded
nearly every major colony of cliff-nesting or
talus-nesting seabirds known in the Aleutians,
but seldom gave sizes of colonies, and
separate colonies on a particular island were
often not differentiated.</p>

<p>World War II brought several ornithologists
to the Aleutians. Cahn (1947), Sutton
and Wilson (1946), Taber (1946), and Wilson
(1948) provided accounts of birds observed at
specific locations. After the war, Fish and
Wildlife Service personnel&mdash;including I. N.
Gabrielson (Gabrielson and Lincoln 1959),
K. W. Kenyon (Kenyon 1961), and R. D. Jones
(Refuge Narrative Reports 1949-1970)&mdash;recorded
observations of breeding seabirds at
several locations in the Aleutians. Investigations
associated with Atomic Energy Commission
nuclear testing at Amchitka Island
provided the first ecological study of avifauna
of an Aleutian island (White et al. 1977). Byrd
et al. (1974) provided a list of birds at Adak.</p>

<p>In 1971, the Near Islands were surveyed by
U.S. Fish and Wildlife personnel in a Cape
Cod dory. In 1972, the Aleutian Islands National
Wildlife Refuge obtained a vessel, the
<em>Aleutian Tern</em>, which allowed visits to all
parts of the island chain. That year, nearly
every large island as far west as Buldir was
visited, and seabird colonies were mapped.
Every island has been visited at least once
since 1972.</p>


<h4>Methods</h4>

<p>In estimating the current status of seabirds
in the Aleutians, all available data were considered.
Most of the information used, however,
is from surveys conducted by the U.S.
Fish and Wildlife Service (1970-75, unpublished
data). Because these surveys only incidentally
included Unimak, Akun, Akutan,
Unalaska, and Umnak islands, data for these
areas are almost totally lacking. Data for
Bogoslof, Adak, Amchitka, Buldir, Agattu,
Nizki, Alaid, and Attu are most accurate because
fairly intensive investigations have
been conducted there since 1970.</p>

<p>The available data are of unknown accuracy.
The method used by most investigators
who have surveyed areas in the Aleutians for
seabird colonies has been to circle islands in a
ship or small boat; when a colony was encountered,
they simply estimated the number
of birds they saw at the time. The accuracy of
the estimates is affected by weather, distance
from the colony, density of birds, ability and
experience of the observer, and other variables.
Estimates of kittiwakes and cormorants
should be the most accurate, since
nests were actually counted. Murres (<em>Uria</em>
spp.) are readily visible on the cliffs, but the
percentage of breeders on the cliffs at a particular
time of day during a particular part of
the breeding season is not known. Auklet
numbers are perhaps hardest to estimate,<span class="pagenum"><a name="Page_36" id="Page_36">[Pg 36]</a></span>
since swirling "clouds" of birds are encountered.</p>

<p>Even when the estimates of birds seen are
assumed to be accurate, data interpretation is
complex. Lack of information on diurnal
rhythms adds difficulty to data interpretation.
Counts of burrow-nesting birds (e.g.,
puffins) have been inaccurately interpreted
because of the lack of understanding of their
nesting ecology. Gulls (<em>Larus</em> spp.), terns, and
jaegers (<em>Stercorarius</em> spp.) are not well known
since shore parties have seldom investigated
island interiors. Nocturnal species (e.g., ancient
murrelet, <em>Synthliboramphus antiquus</em>,
and storm-petrels, <em>Oceanodroma</em> spp.) are perhaps
the least known. Since only crude estimates
of colony sizes are available, broad
limits are used in this paper to describe known
colonies.</p>


<h4>Status and Distribution
of Breeding Seabirds</h4>

<p>Even from the sparse literature available, it
is apparent that some seabird populations are
now drastically different from those in the
Aleutians around 1900. Changes in nesting
habitat due to volcanic eruptions, tidal waves,
marine erosion, and earthquakes have occurred
for centuries, and colonial nesting bird
populations have fluctuated accordingly. In
addition, native Aleuts used marine birds and
their eggs for food and their skins for clothing,
but the Aleuts were so diminished in
numbers by 1900 that they have had little recent
effect on the bird populations.</p>

<p>From about 1900 to 1936, arctic foxes were
introduced to most of the Aleutians for fur
farming. The foxes lived on birds in summer,
and some species (e.g., Aleutian Canada
geese, <em>Branta canadensis leucopareia</em>) were
wiped out wherever foxes were introduced.
Ground-nesting and some burrow-nesting seabirds
were also drastically reduced or extirpated
on many islands.</p>

<p>During World War II the thousands of
troops in the Aleutians brought dogs and cats
to some of the islands as pets, and many of
the animals were set free when the men departed.
The military also accidentally introduced
Norway rats to some of the islands.
Their role in seabird population reductions is
unknown.</p>

<p>Figures 2-15 (pages <a href="#Page_40">40</a>-46) present data on
the distribution of populations of birds that
have survived the foxes and other introduced
predators. An annotated list of seabirds
breeding in the Aleutians follows.</p>


<h5><em>Annotated List of Species</em></h5>


<h6>Northern fulmar <em>(Fulmarus glacialis)</em></h6>

<p>Northern fulmars breed on only three islands:
Buldir (200 pairs), Gareloi (1,500 pairs),
and Chagulak (more than 100,000 pairs).
Fulmars were apparently much more widespread
formerly (Murie 1959; Turner 1886).
Introduced foxes were probably involved in
the decline.</p>


<h6>Fork-tailed Storm-petrel and Leach's
Storm-petrel (<em>Oceanodroma furcata</em> and
<em>O. leucorhoa</em>)</h6>

<p>The distribution of storm-petrels is poorly
known due to their nocturnal behavior near
the nesting colonies. The presence of birds has
generally been noted by finding them aboard
ships anchored near islands after darkness.
Population estimates are not available for any
colonies, so symbols used in Fig. 3 indicate
probable numbers of breeding birds. In few
cases have active burrows or crevices been
discovered. Storm-petrels were formerly
much more common. Murie (1959) and John L.
Trapp (personal communication) found large
numbers of storm-petrel remains in fox dens.
Most present breeding colonies are probably
confined to offshore islets and fox-free
islands.</p>


<h6>Double-crested Cormorant, Pelagic
Cormorant, and Red-faced Cormorant
(<em>Phalacrocorax auritus</em>, <em>P. pelagicus</em>,
and <em>P. urile</em>)</h6>

<p>Double-crested cormorants breed as far
west as the Islands of Four Mountains. The
colonies vary in size from a few to 25 pairs.
Pelagic and red-faced cormorants nest from
Amak to Attu on nearly every island. Relative
abundance of the two in mixed colonies varies
between areas as well as from year to year.
Red-faced cormorants tend to nest in colonies
mixed with kittiwakes and murres, but pure
colonies also occur. Pelagic cormorants occupy
isolated, small colonies, but they also<span class="pagenum"><a name="Page_37" id="Page_37">[Pg 37]</a></span>
nest with kittiwakes and murres and are often
found with red-faced cormorants. By far the
densest concentration of cormorants occurs in
the Near Islands, especially at Attu, where an
estimated 77,000 birds were seen in 1970. In
the Aleutians as a whole, red-faced cormorants
outnumber pelagic cormorants, and
double-crested cormorants make up only a
very small percentage of the breeding
population.</p>

<h6>Parasitic Jaeger <em>(Stercorarius parasiticus)</em></h6>

<p>The distribution of jaegers is poorly known
because investigators have spent little time
ashore on most islands. Murie (1959) found
jaegers on a number of islands, and most of
the data in Fig. 5 are his. Population estimates
are available only for Amchitka (25
pairs; White et al. 1977) and Buldir (30-40
pairs; G. V. Byrd, unpublished data).</p>


<h6>Glaucous-winged Gull <em>(Larus glaucescens)</em></h6>

<p>Glaucous-winged gulls no longer nest on
islands where foxes occur except where
islands in lakes are available. Most colonies
are on offshore rocks or islets and range in
size from a few pairs to over 200 pairs, and
occasionally more. They are found throughout
the Aleutians, but the largest known colonies
are at Bogoslof (500 pairs) and Buldir (250
pairs).</p>


<h6>Black-legged Kittiwake and Red-legged
Kittiwake (<em>Rissa tridactyla</em> and
<em>R. brevirostris</em>)</h6>

<p>Black-legged kittiwakes breed locally in
every major island group, usually mixed with
murres and cormorants. The large colonies
contain over 25,000 birds, but colonies of less
than 50 pairs also occur. Red-legged kittiwakes
breed only on Buldir and Bogoslof.
They are remnants of a previously more widespread
population.</p>


<h6>Arctic Tern and Aleutian Tern (<em>Sterna
paradisaea</em> and <em>S. aleutica</em>)</h6>

<p>Terns breed locally in each island group.
Both species occur at Attu, Amchitka, Adak,
and Umnak, but only arctic terns are found at
Nizki. Factors limiting distribution are unknown.
Colonies vary in size from less than 10
pairs to 100 pairs.</p>


<h6>Common Murre and Thick-billed Murre
(<em>Uria aalge</em> and <em>U. lomvia</em>)</h6>

<p>Like kittiwakes, murres are abundant
locally. A pure colony of either species is almost
unknown, although one species often
makes up more than 90% of a colony. Common
murres may have been reduced by foxes,
since they tend to use sites with less slope
than those used by thick-billed murres. At
Bogoslof and the Baby islands, the birds use
inland, gently sloping areas because there are
no foxes. The presence of the lichen <em>(Caloplaca
spp.)</em>, which according to Tuck (1960) is indicative
of bird roosts, on several extensive
cliff areas suggests that either murres or kittiwakes,
or both, formerly used areas they do
not use now.</p>


<h6>Pigeon Guillemot <em>(Cepphus columba)</em></h6>

<p>This species has been noted near almost
every island that has been visited. Nesting
under beach boulders and driftwood, the birds
only occasionally are found in large concentrations
(near Great Sitkin more than 4,000 birds
were seen in 1971). Murie et al. (1937) summed
up the distribution of pigeon guillemot accurately:
"Each island has its meager quota of
these birds, nesting unobtrusively among the
rocks but never assembled in any really large
groups." Estimates of populations may be extremely
inaccurate because the diurnal
rhythm of the pigeon guillemot is unknown.</p>


<h6>Marbled Murrelet and Kittlitz's Murrelet
(<em>Brachyramphus marmoratus</em> and
<em>B. brevirostris</em>)</h6>

<p>Nests of neither species have been located
in the Aleutians, but nesting of both is suspected
at Adak, Unalaska, and Unimak,
where specimens of Kittlitz's with brood
patches or eggs in the oviduct have been collected
in nearshore waters. Courtship has
been recorded in marbled murrelets (Byrd et
al. 1974).</p>


<h6>Ancient Murrelet <em>(Synthliboramphus
antiquus)</em></h6>

<p>The distribution of this species is very
poorly known, since it is nocturnal near nesting
colonies. Murie (1959) wrote, "This is one
of the species that undoubtedly has greatly
declined in recent years, as a result of increase<span class="pagenum"><a name="Page_38" id="Page_38">[Pg 38]</a></span>
of the blue-fox industry." The leading of
downy young to sea by the adults is a very
noisy process and foxes could easily take
large numbers. Also, these murrelets nest in
fairly shallow burrows which foxes could dig
out easily. Birds were recorded near islands in
every group during surveys from 1972 to
1975, but workers seldom went ashore to determine
if they were nesting. In Fig. 12, the
only basis for designating most of the areas
marked as colonies is the presence of birds
during breeding season (15 May-1 July).</p>


<h6>Cassin's Auklet <em>(Ptychoramphus aleuticus)</em></h6>

<p>This is another species that was more common
before the fox was introduced. Cassin's
auklet now seems to occur only locally, but
these nocturnal birds are probably often overlooked.
They are known only from Buldir,
Umnak, and the vicinity of Oglodak.</p>


<h6>Parakeet Auklet <em>(Cyclorrhynchus psittacula)</em></h6>

<p>This auklet, which nests under beach
boulders, in burrows, and in rock crevices,
seems to use a greater variety of breeding
sites than do the other auklets. The largest
known colony is at Chagulak, where an estimated
10,000 were seen in 1972. Smaller colonies
are found as far west as Buldir.</p>


<h6>Crested Auklet, Least Auklet, and Whiskered
Auklet (<em>Aethia cristatella</em>, <em>A. pusilla</em>,
and <em>A. pygmaea</em>)</h6>

<p><em>Aethia</em> nest primarily in rock crevices of
talus slides. Such habitat occurs locally in
each major island group except the Near
Islands. Least auklets outnumber crested
auklets in the Aleutians, and whiskered auklets
are far less common than either. Estimates
of populations are probably grossly inaccurate
because of the difficulty both in estimating
the number of birds in the milling
flocks observed and in interpreting the estimates
after they are obtained.</p>


<h6>Horned Puffin and Tufted Puffin (<em>Fratercula
corniculata</em> and <em>Lunda cirrhata</em>)</h6>

<p>Horned puffins favor rock crevices in talus
slides and cliff faces for nesting, whereas
tufted puffins are primarily burrow nesters.
The historical distribution of the two species
was probably based on availability of nesting
sites, so tufted puffins were more widespread
and numerous. However, in areas where extensive
talus slopes are available, horned
puffins reached high densities. Predation by
introduced foxes may have altered the distribution
of tufted puffins, which now nest primarily
on fox-free islets just offshore from the
larger islands where foxes occur. The distribution
of horned puffins may not have been
altered significantly, since they are relatively
free from fox predation in their rock crevices.</p>


<h4>Recommendations</h4>

<p>A complete survey of the Aleutian Islands
has not been done. This should be done, by
methods that will provide accurate population
estimates. Life history information is needed
on almost all species, and data should be
gathered on selected populations to determine
trends. Information on winter distribution
should also be compiled. The effects of introduced
predators should be evaluated quantitatively,
and if control measures are needed,
effective, humane methods should be devised
and implemented.</p>


<h4>Acknowledgments</h4>

<p>The authors are grateful to the following
Fish and Wildlife Service personnel who
helped collect previously unpublished data
used in this paper: E. P. Bailey, C. S. Craighead,
C. P. Dau, M. H. Dick, G. J. Divoky, R.
Martin, J. L. Trapp, G. W. Watson, and C. M.
White. Most of the data were collected from
the deck of the Aleutian Islands National
Wildlife Refuge research vessel <em>Aleutian
Tern</em>. Captain George Putney is acknowledged
for his peerless seamanship and constant
encouragement; he also contributed observations
of birds.</p>

<p>The maps were adapted from a master supplied
by Elaine Rhode, Public Affairs Office,
U.S. Fish and Wildlife Service, Anchorage;
she also suggested the use of squares to display
data. C. M. White graciously made his in-press
manuscript available. W. B. Emison,
R. J. Gordon, and J. L. Trapp kindly made
their field notes available, and Trapp helped
compile data. Most of the funds for the surveys
in 1971-75 were provided by the U.S.
Fish and Wildlife Service.</p>

<p><span class="pagenum"><a name="Page_39" id="Page_39">[Pg 39]</a></span></p>


<h4>References</h4>

<div class="hangindent">

<p>Amundsen, C. C., and E. E. C. Clebsch. 1971. Dynamics
of the terrestrial ecosystem vegetation of
Amchitka Island, Alaska. Bioscience 21:619-623.</p>

<p>Bent, A. C. 1912. Notes on birds observed during a
brief visit to the Aleutian Islands and Bering Sea
in 1911. Smithsonian Misc. Coll. 56(2):1-29.</p>

<p>Byrd, G. V., D. D. Gibson, and D. L. Johnson. 1974.
The birds of Adak Island, Alaska. Condor 76:288-300.</p>

<p>Cahn, A. R. 1947. Notes on the birds of the Dutch
Harbor area of the Aleutian Islands. Condor
49:78-82.</p>

<p>Clark, A. H. 1910. The birds collected and observed
during the cruise of the United States Fisheries
steamer "Albatross" in the north Pacific Ocean
and in the Bering, Okhotsk, Japan, and Eastern
seas from April to December 1906. Proc. U.S.
Natl. Mus. 38:25-74.</p>

<p>Dall, W. H. 1873. Notes on the avifauna of the Aleutian
Islands, from Unalaska eastward. Proc.
Calif. Acad. Sci. 5:25-35.</p>

<p>Dall, W. H. 1874. Notes on the avifauna of the Aleutian
Islands, especially those west of Unalaska.
Proc. Calif. Acad. Sci. 5:270-281.</p>

<p>Gabrielson, I. N., and F. C. Lincoln. 1959. The birds
of Alaska. The Stackpole Co., Harrisburg, Penn.,
and Wildlife Management Institute, Washington,
D.C. 922 pp.</p>

<p>Hulten, E. 1960. Flora of the Aleutian Islands. J.
Kramer, Weinham/Bergstr., Sweden. 376 pp.</p>

<p>Isakson, J. S., C. A. Sinensted, and R. L. Burgner.
1971. Fish communities and food chains in the
Amchitka area. Bioscience 21:666-670.</p>

<p>Jones, R. D. 1949-1970. Annual refuge narrative reports,
Aleutian Islands N.W.R. Cold Bay,
Alaska. (Unpublished administrative report.)</p>

<p>Kenyon, K. W. 1961. Birds of Amchitka Island,
Alaska. Auk 78:305-326.</p>

<p>Lebednik, P. A., F. C. Weinmann, and R. E. Norris.
1971. Spatial and seasonal distributions of
marine algal communities at Amchitka Island,
Alaska. Bioscience 21:656-660.</p>

<p>Murie, O. J. 1959. Fauna of the Aleutian Islands
and Alaska Peninsula. U.S. Fish Wildl. Serv., N.
Am. Fauna 61. 364 pp.</p>

<p>Nelson, E. W. 1887. Report upon natural history
collections made in Alaska between the years
1877 and 1881. U.S. Army, Signal Serv. Arct.
Ser. Publ. 3. 337 pp.</p>

<p>O'Clair, C. E., and K. K. Chew. 1971. Transect
studies of littoral macrofauna, Amchitka Island,
Alaska. Bioscience 21:661-664.</p>

<p>Stejneger, L. 1936. George Wilhelm Steller&mdash;the
pioneer of Alaska natural history. Harvard University
Press, Cambridge, Mass. 623 pp.</p>

<p>Sutton, G. M., and R. S. Wilson. 1946. Notes on the
winter birds of Attu. Condor 48:83-91.</p>

<p>Taber, R. D. 1946. The winter birds of Adak,
Alaska. Condor 48:272-277.</p>

<p>Tuck, L. M. 1960. The murres. Ottawa. Canadian
Wildlife Series 1. 260 pp.</p>

<p>Turner, L. M. 1885. Notes on the birds of the Near
Islands, Alaska. Auk 2:154-159.</p>

<p>Turner, L. M. 1886. Contributions to the natural
history of Alaska. U.S. Army, Signal Serv. Arct.
Ser. Publ. 2. 226 pp.</p>

<p>White, C. M., F. S. L. Williamson, and W. B.
Emison. 1977. Avifaunal investigations. Pages
227-260 <em>in</em> M. L. Merritt and R. G. Fuller, eds.
Environments of Amchitka Island, Alaska. U.S.
Energy Research and Development Association,
Oak Ridge, Tenn. 682 pp.</p>

<p>Wilson, R. S. 1948. The summer bird life of Attu.
Condor 50:124-129.</p></div>

<p><span class="pagenum"><a name="Page_40" id="Page_40">[Pg 40]</a></span></p>

<div class="figcenter" style="width: 700px;">
<img src="images/p_040a.jpg" width="700" height="181" alt="" />
<div class="caption"><p>Fig. 2. Breeding distribution of northern fulmar.</p></div>
</div>

<div class="figcenter" style="width: 700px;">
<img src="images/p_040b.jpg" width="700" height="183" alt="" />
<div class="caption"><p>Fig. 3. Breeding distribution of storm-petrels.</p></div>
</div>

<p><span class="pagenum"><a name="Page_41" id="Page_41">[Pg 41]</a></span></p>

<div class="figcenter" style="width: 700px;">
<img src="images/p_041a.jpg" width="700" height="178" alt="" />
<div class="caption"><p>Fig. 4. Breeding distribution of cormorants.</p></div>
</div>

<div class="figcenter" style="width: 700px;">
<img src="images/p_041b.jpg" width="700" height="178" alt="" />
<div class="caption"><p>Fig. 5. Breeding distribution of parasitic jaeger.</p></div>
</div>

<p><span class="pagenum"><a name="Page_42" id="Page_42">[Pg 42]</a></span></p>

<div class="figcenter" style="width: 700px;">
<img src="images/p_042a.jpg" width="700" height="182" alt="" />
<div class="caption"><p>Fig. 6. Breeding distribution of glaucous-winged gull.</p></div>
</div>

<div class="figcenter" style="width: 700px;">
<img src="images/p_042b.jpg" width="700" height="181" alt="" />
<div class="caption"><p>Fig. 7. Breeding distribution of kittiwakes.</p></div>
</div>

<p><span class="pagenum"><a name="Page_43" id="Page_43">[Pg 43]</a></span></p>

<div class="figcenter" style="width: 700px;">
<img src="images/p_043a.jpg" width="700" height="180" alt="" />
<div class="caption"><p>Fig. 8. Breeding distribution of terns.</p></div>
</div>

<div class="figcenter" style="width: 700px;">
<img src="images/p_043b.jpg" width="700" height="182" alt="" />
<div class="caption"><p>Fig. 9. Breeding distribution of murres.</p></div>
</div>

<p><span class="pagenum"><a name="Page_44" id="Page_44">[Pg 44]</a></span></p>

<div class="figcenter" style="width: 700px;">
<img src="images/p_044a.jpg" width="700" height="181" alt="" />
<div class="caption"><p>Fig. 10. Breeding distribution of pigeon guillemot.</p></div>
</div>

<div class="figcenter" style="width: 700px;">
<img src="images/p_044b.jpg" width="700" height="181" alt="" />
<div class="caption"><p>Fig. 11. Breeding distribution of marbled and Kittlitz's murrelet.</p></div>
</div>

<p><span class="pagenum"><a name="Page_45" id="Page_45">[Pg 45]</a></span></p>

<div class="figcenter" style="width: 700px;">
<img src="images/p_045a.jpg" width="700" height="181" alt="" />
<div class="caption"><p>Fig. 12. Breeding distribution of ancient murrelet.</p></div>
</div>

<div class="figcenter" style="width: 700px;">
<img src="images/p_045b.jpg" width="700" height="179" alt="" />
<div class="caption"><p>Fig. 13. Breeding distribution of auklets.</p></div>
</div>

<p><span class="pagenum"><a name="Page_46" id="Page_46">[Pg 46]</a></span></p>

<div class="figcenter" style="width: 700px;">
<img src="images/p_046a.jpg" width="700" height="182" alt="" />
<div class="caption"><p>Fig. 14. Breeding distribution of horned puffin.</p></div>
</div>

<div class="figcenter" style="width: 700px;">
<img src="images/p_046b.jpg" width="700" height="182" alt="" />
<div class="caption"><p>Fig. 15. Breeding distribution of tufted puffin.</p></div>
</div>

<div class="footnotes"><h4>FOOTNOTES:</h4>

<div class="footnote">

<p><a name="Footnote_3_3" id="Footnote_3_3"></a><a href="#FNanchor_3_3"><span class="label">[3]</span></a> Present address: U.S. Fish and Wildlife Service,
William L. Finley National Wildlife Refuge, Route
2, Box 208, Corvallis, Oregon 97330.</p></div>

<div class="footnote">

<p><a name="Footnote_4_4" id="Footnote_4_4"></a><a href="#FNanchor_4_4"><span class="label">[4]</span></a> Present address: U.S. Fish and Wildlife Service,
Kilauea, Hawaii.</p>

<p><span class="pagenum"><a name="Page_47" id="Page_47">[Pg 47]</a></span></p></div></div>
<div class="chapter"></div>
<hr class="chap" />




<h3><a name="The_Historical_Status_of_Nesting_Seabirds_of_the" id="The_Historical_Status_of_Nesting_Seabirds_of_the">The Historical Status of Nesting Seabirds of the
Northern and Western Gulf of Alaska</a></h3>


<p class="center">by</p>

<p class="center">LeRoy W. Sowl</p>

<p class="center">U.S. Fish and Wildlife Service<br />
1011 East Tudor Road<br />
Anchorage, Alaska 99507
</p>


<h4>Abstract</h4>

<div class="blockquot">

<p>The history of ornithological field work in the Gulf of Alaska dates back to 20
July 1741 and Bering's discovery of Alaska. In spite of this long history, the
record is fragmentary and often seemingly contradictory. The coming of the
tanker terminal at Valdez and the pending development of oil and gas resources
on the outer continental shelf threaten massive change for seabirds in the Gulf
of Alaska. Often overlooked, however, is the fact that man has already effected a
change in status for many of these birds. In this paper I examine the scanty,
general record from the exploratory period, roughly 1741 to 1935, and the somewhat
more comprehensive record of the reconnaissance period, 1936-74, and
attempt to develop a basis for better understanding of the change in seabird
status that has already taken place. This paper should be treated as a verbal
model which can be improved as our knowledge of seabirds in the Gulf of Alaska
is expanded.</p></div>

<p>From the perspective of history, 1970
should prove to have been a momentous year
for Alaska and its seabirds. Two events, the
construction of the Trans-Alaska Pipeline and
the passage of the National Environmental
Policy Act (NEPA) merged head on in 1970
with the decision that Section 2c of NEPA applied
to the proposed pipeline. The systematic
appraisal of potential environmental impacts
required by Section 2c quickly exposed the inadequacy
of the existing data base in many
areas. With respect to seabirds in the Gulf of
Alaska, it was apparent that there had never
been any effort to develop a synthesis of the
information accumulated over 230 years. The
data gaps which were uncovered were appalling.</p>

<p>While the Trans-Alaska Pipeline impact
statement had provided shock therapy, it was
not the only influential event on the horizon.
Two local disturbances had already preceded
the pipeline. These were Project Chariot at
Cape Thompson and the Amchitka Island test
program. Now in quick succession the Wilderness
Act and native land claims added new
urgency to the need for solid resource information.
More recently, the outer continental
shelf minerals leasing program has made the
quick development of base-line information
even more essential.</p>

<p>All of the new activity in Alaska's coastal
waters has the potential to affect seabirds in
one way or another. We must remember, however,
that man's activities have been affecting
seabirds for a long time. We cannot accurately
assess the effect of a tanker terminal at
Valdez or offshore oil activity without first
developing some understanding of the current
status of seabirds in the context of the historical
record.</p>

<p>Seabird work in Alaska can be divided
roughly into three periods. The first is the
early historical or exploratory period; it extended
from Georg Steller's 1741 visit to
Kayak Island to 1935. This was literally a
period of exploration and the collection of in<span class="pagenum"><a name="Page_48" id="Page_48">[Pg 48]</a></span>formation
was dependent upon interest and
opportunity. The second is the reconnaissance
period; during this period investigators were
dispatched to a particular area to gather
general information for management application.
This period begins with Murie's extensive
investigations of the Alaska Peninsula
and the Aleutian Islands; I see it extending
from 1936 to 1975. In 1975 the need for data
became so acute that it was necessary to enter
the third period, one of intensive data gathering.
Knowing where the big seabird colonies
were located and knowing their general
species composition was no longer adequate.
The current intensive data-gathering effort in
the waters over oil and gas leasing areas is a
partial response to the recognition of this
inadequacy.</p>

<p>In this paper I draw some tentative conclusions
relative to the status of the 26 species of
primary seabirds (Fisher and Lockley 1954)
breeding in, or which may have bred in, the
northern and western Gulf of Alaska area.
This area extends from Cape Fairweather,
59&deg;N 138&deg;W, westerly along the coast to Ikatan
Bay, 55&deg;N 163&deg;W, at the end of the
Alaska Peninsula. These bird species tend to
be colonial, but not exclusively so. Two birds
which are primary seabirds, the mew gull
<em>(Larus canus)</em> and Bonaparte's gull <em>(L. philadelphia)</em>,
have not been included because they
tend to be more riverine than marine in habit.
Several marine ducks have been excluded because
they are secondary seabirds.</p>

<p>Information from the early exploratory
period is summarized under the next section.
The more detailed information from the reconnaissance
period is discussed in the species
accounts.</p>


<h4>Summary of the
Historical Record</h4>

<p>The history of ornithological field work in
the Gulf of Alaska goes back 235 years to 20
July 1741. On that day Bering's surgeon/naturalist,
Georg W. Steller, spent a
scant 10 h ashore on Kayak Island. He collected
a single bird. This bird, later named for
Steller, reminded him of a plate of the blue jay
by Make Catesby, the colonial-era predecessor
of Audubon, in Volume 1 of the <cite>Natural
History of Carolina, Florida, and the Bahama
Islands</cite> (Stejneger's annotated translation of
Steller's journal in Golder 1925). Collection of
the bird confirmed for Steller that the first
Russian Expedition had reached America.</p>

<p>Steller was an accomplished naturalist, but
his overbearing and superior manner had apparently
sorely irritated Bering and his officers
long before the expedition reached Kayak
Island. The seamen made little effort to go
ashore anywhere in Alaska and Steller was
blocked from doing so as well. In addition to
Kayak Island, he was able to go ashore only
on Nagai Island, first with a water party on
30 August and again the next day. He noted
that "all sorts of waterbirds in abundance
were seen." These included two kinds of cormorants,
auks, ducks, gulls, divers, pigeon
guillemots <em>(Cepphus columba)</em>, tufted puffins
<em>(Lunda cirrhata)</em>, and horned puffins <em>(Fratercula
corniculata)</em>.</p>

<p>Stejneger's comment on the identity of the
cormorants is interesting because, based on
his experience, he assumed them to be pelagic
and double-crested cormorants (<em>Phalacrocorax
pelagicus</em> and <em>P. auritus</em>). He gave no
thought to red-faced cormorants <em>(P. urile)</em>
which are now common there.</p>

<p>Steller noted on 6 September off Bird Island
in the Shumagin Islands, that "when we were
out to sea about half a mile we were especially
astonished at the untold numbers of seabirds
which we saw on the northern side of the island."
These birds were listed as cormorants,
auks, horned puffins, fulmars <em>(Fulmarus glacialis)</em>,
pigeon guillemots, black oystercatchers
<em>(Haematopus backmani)</em>, and a pied
diver which Stejneger assumed was an ancient
murrelet <em>(Synthliboramphus antiquus)</em>.</p>

<p>On 15 September when Bering's vessel, the
<em>St. Peter</em>, was south of Amukta Pass, Steller
recorded observing "river gulls." The observation
is not as interesting as Stejneger's
comment (Golder 1925) concerning it. Stejneger
stated that no true river gulls lived in
the Aleutians and these must, therefore, have
been another small gull with red feet. He
thought they must have been the red-legged
kittiwake <em>(Rissa brevirostris)</em>, which "inhabits
the Aleutian Islands from Bering Island
to Sannak."</p>

<p>Thirty-seven years after Bering's voyage,
Captain James Cook sailed into the Gulf of<span class="pagenum"><a name="Page_49" id="Page_49">[Pg 49]</a></span>
Alaska, arriving off Kayak Island on 11 May
1778. Cook was not accompanied by an able
naturalist. His surgeon, William Anderson,
did have some experience gained on earlier
voyages in preparing skins and taking notes,
but he had contracted tuberculosis and became
so ill that even his notes ceased after 8
June, while the expedition was in Cook Inlet.</p>

<p>Cook was under orders to keep a careful record
of everything he saw. One of the results
was that he had birds collected even though
he had no naturalist to do the work. Several
birds were collected in Prince William Sound
while Cook's vessels were at anchor in Port
Etches. These included two marbled murrelets
(<em>Brachyramphus marmoratus</em>&mdash;type
specimens), a black oystercatcher, a surfbird
<em>(Aphriza virgata)</em>, a surf scoter <em>(Melanitta perspicillata)</em>,
and a red-breasted merganser
(<em>Mergus serrator</em>&mdash;type specimen), along with
several forest birds (Stresemann 1949).</p>

<p>The watch journals of Cook and his officers
provide some additional information. Captain
Charles Clerke (Beaglehole 1974) remarked in
his log on the passage out of Prince William
Sound through Montague Strait on 20 May
that "it had almost become tautology to mention
whales and seals and innumerable sea
fowl that so confoundingly kept their
distance."</p>

<p>Between the Trinity Islands and Chirikof
Island on 18 June, Cook's men collected a
single tufted puffin. Later Cook passed close
to the Semidi Islands and the Shumagin Islands
and directly through the Sandman
Reefs. Beaglehole's version of this part of the
voyage makes no mention of seabirds.</p>

<p>There is a gap of 87 years during which
there is almost no hint of published material
bearing on the status of seabirds in the Gulf
of Alaska. In 1865 the Russo-American Telegraph
Expedition touched this area. Dall and
Bannister (1869) provide us with a few scraps
garnered during that expedition, primarily by
Bischoff. The glaucous-winged gull <em>(Larus
glaucescens)</em> was described as the most common
species from California northward.
Bischoff's collections at Kodiak indicate that
the horned and tufted puffins were collected
with ease. He was able also to collect an Aleutian
tern (<em>Sterna aleutica</em>&mdash;type specimen)
along with an egg.</p>

<p>Dall (1873) noted in 1872 that the black-legged
kittiwake <em>(Rissa tridactyla)</em> was common
at Round Island and Delarof Harbor,
Unga Island, in the Shumagins. The inference
is that it was more common at these two
places than elsewhere. The Arctic tern <em>(Sterna
paradisaea)</em> was abundant in the Shumagin
Islands and particularly at Range Island in
Popoff Strait. Dall expressed the opinion that
the horned puffin was very abundant in the
Shumagins and appeared to fill the niche of
the tufted puffin, which he did not see there.
The only other bird which he thought to be
very common was the pigeon guillemot. He
did not note the common murre <em>(Uria aalge)</em> at
all.</p>

<p>In 1908 the second of three Alexander Expeditions
conducted field work in the Prince
William Sound area. From Dixon (1908) and
Grinnell (1910) we can derive some basis for
assessing status in a very general way. The
most common seabird noted was the marbled
murrelet. Glaucous-winged gulls and black-legged
kittiwakes were common; the glaucous-winged
gull was the more common.
Horned puffins were judged to be slightly
more common than tufted puffins by both
authors. The northern end of Montague Strait
appears to have been the center of abundance
for puffins. Dixon noted that on 16 July 1908
there were swarms of puffins in the channel
along Green Island. Pigeon guillemots were
common along the rocky coasts. Parakeet
auklets <em>(Cyclorrhyncus psittacula)</em>, common
murres, and ancient murrelets were noted
only in very small numbers.</p>

<p>After the Alexander Expeditions there was
another doldrum in which little was done.
During this lull in activity, a note by Townsend
(1913) appeared which compared the
numbers of crested auklets <em>(Aethia cristatella)</em>
at Yukon Harbor, Big Koniuji Island, to the
least auklets <em>(A. pusilla)</em> of St. George Island,
stating that the crested auklets were more
numerous. He sailed into the Yukon Harbor
anchorage on the evening of 1 August and observed
that crested auklets "were present in
myriads. The surface of the water was covered
with them, and the air was filled with them."</p>

<p>The formal record available to researchers
is very shallow for this exploratory period.
With a few exceptions it was compiled by non-scientists,
primarily explorers and egg and
skin collectors.</p>

<p><span class="pagenum"><a name="Page_50" id="Page_50">[Pg 50]</a></span></p>


<h4>Current Status</h4>


<h5><em>Setting the Stage</em></h5>

<p>This paper should be viewed as a conceptual
model. While I attempted to be as objective as
possible, subjectivity was unavoidable. Many
of the tentative conclusions are based on very
little data. Each improvement will make it a
better management tool. Because of the space
limitations, it is not possible to go into a detailed
tracking of my reasoning for each
species. In an attempt to overcome this handicap,
I am including some examples of the
sorts of reasoning that went into the process.</p>

<p>In 1973 I led a Fish and Wildlife Service
(FWS) reconnaissance survey team that was
delineating seabird colonies along the Alaska
Peninsula. In the Shumagin Islands we entered
or crossed Koniuji Strait twice (on 11
and 12 June) without even suspecting the
presence of a horned puffin colony. A third
passage through the strait (13 June) was not
so uneventful. The water and the air were
filled with horned puffins. This led to the discovery
that the 430-m mountain on the southeastern
corner of Big Koniuji was also
covered with horned puffins, clear to its top.
The minimum estimate of the birds that were
visible was 140,000. Even this number of
birds would make this the largest horned
puffin colony ever discovered. David Spencer
(personal communication) had noted similar
swarms of horned puffins in this strait in 1956
while flying sea otter surveys in the area. In
1975 a field camp was established at Yukon
Harbor, with study of this colony as one of the
prime objectives of the investigators. As far
as these investigators could tell no such large
colony existed there, even though the nesting
habitat was still there, unaltered. This sort of
event, one of the banes and vagaries of estimating
seabird numbers, is not rare.</p>

<p>In 1973, when FWS personnel delineated
the colony on the southwestern end of Bird Island
in the Shumagins, there were estimated
to be 43,000 kittiwakes, 24,000 murres, and
6,000 cormorants present; no tufted puffins
were seen about the colony. The last time (in
1970) one of the observers, Edgar Bailey, had
visited the colony with Robert Jones, there
was an extremely large colony of tufted
puffins which Jones (E. Bailey, personal communication)
estimated at more than 1 million
birds. We made a particular effort to visit
Jude Island, between the Shumagin Islands
and the Pavlof Islands, because David
Spencer (personal communication) had reported
once having seen the air over the island
filled with an extremely large number of
tufted puffins. However, there were no puffins
at this colony either.</p>

<p>Let us examine the facts in context. On 8
June we had visited High Island where we had
attempted to collect puffin eggs for pesticide
analysis, but had been able to find only one
egg. Also, there were only 6,000 tufted puffins
where George Putney, master/engineer of the
<em>Aleutian Tern</em>, had seen much larger numbers
in 1972. These two facts could easily be related
to explain the current situation because
it was still early in the breeding season. The
horned puffin observations in Koniuji Strait
(11-13 June) were in keeping with this conclusion
also&mdash;an indication that these birds had
not yet settled down to a full breeding effort.
The erratic comings and goings of common
puffins <em>(Fratercula arctica)</em> early in the season
have been well documented (Lockley 1962). It
is an easy step to extend this reasoning to the
absence of birds at Bird Island on 11 June,
even though fresh signs of the characteristic
evidence of tufted puffin occupancy were
missing. Jude Island provides a different clue,
however. There were 3,000 pigeon guillemots,
an unheard-of concentration, apparently
occupying abandoned tufted puffin burrows
on 15 June. Also, on 7 June we had made a
very interesting observation that had no
special significance at the time: murres on
Spitz Island were occupying little parapets
created by mashing down the mouths of
puffin burrows which filled the slope above
the cliff portion of their colony.</p>

<p>After looking at all of the observations
cited above, I conclude that tufted puffins
were greatly reduced in numbers on these
sites in 1973 and that they had been absent
from the burrows used by the murres and
pigeon guillemots for more than the current
breeding season. What causes these sorts of
changes? I do not know.</p>

<p>One reason for year-to-year change may be
local movements of colonies. Black-legged
kittiwakes nest at several places in lower Orca
Inlet, Prince William Sound. Counts made at
these sites in 1972 and 1974 yielded almost<span class="pagenum"><a name="Page_51" id="Page_51">[Pg 51]</a></span>
identical totals but the numbers of birds
varied between individual sites. This may be
an indication that all of these sites are part of
one large composite colony and that, at least
in this colony and for this species, the birds
shift at will.</p>

<p>The best record of population flux involving
two species has been summarized by Peterson
and Fisher (1955). In 1872 and 1873 the
murres observed on Walrus Island in the
Pribilofs were almost entirely common
murres. In 1890 common and thick-billed
murres <em>(Uria lomvia)</em> were evenly matched in
number. By 1901 the colony was almost exclusively
dominated by thick-billed murres. In
1911 and 1914 the few thick-billed murres
present were almost lost among the then
dominant common murres. In 1940 thick-billed
murres dominated again. When Peterson
and Fisher visited the island in 1953, the
situation was again reversed and common
murres had almost completely replaced the
thick-billed murres. These changes are even
more impressive because of the number of
birds involved, between 1 and 2 million in
1953. There are more tenuous indications that
somewhat the same thing may occur between
two other congener pairs, the pelagic and red-faced
cormorants and the black-legged and
red-legged kittiwakes. The causative factor,
or factors, is not readily apparent. One possibility
is long-term climatic fluctuation.</p>

<p>Dement'ev and Gladkov (1966) provide an
example of abrupt and massive change. Before
1876, the pelagic cormorant abounded on
the Commander Islands. During the winter of
1876-77, the birds were decimated by an unknown
epizootic disease. By spring only a few
individuals remained alive. The record shows
that by 1882 they were already becoming
common again. Red-faced cormorants were
apparently not reduced in number because
Dement'ev and Gladkov (1966) state that
they were common in "the second half of the
last century and the beginning of this." Did
they flourish only while the pelagic cormorants
were reduced in number?</p>

<p>Bowles (1908) gives another indication of
naturally induced population impact. He
noted large numbers of dead seabirds on
Washington beaches and the ocean "rather
plentifully dotted with sick birds ..." He
examined some birds and found "many hundreds"
of tapeworms in every bird. His conclusion
was that their intestines were so
solidly packed with tapeworms that starvation
was "an absolute certainty."</p>

<p>Some apparent disruptions are long term.
In the Gulf of Alaska there is a hiatus in the
distributions of a number of small seabirds
that are active around their colonies only at
night. Repeatedly, the northern Gulf of
Alaska shows up as an area of reduced population,
as a boundary between subspecies, or as
a limit to a range. This same area has a noticeable
lack of total darkness during a substantial
portion of the breeding season.</p>

<p>The nocturnal habit no doubt evolved because
it was advantageous to concentrate on
the breeding grounds only under the cover of
darkness, when diurnal predators were at a
great disadvantage. Cody (1973) states that
Cassin's auklet <em>(Ptychoramphus aleuticus)</em>,
which is strictly nocturnal around its colonies,
avoids these colonies on brightly moonlit
nights. He sees this as an apparent response
to gull predation. At higher latitudes the
small alcids have overcome this disadvantage
by swamping predators through their sheer
numbers. In the Gulf of Alaska I suspect that
few of the small seabirds, except possibly the
fork-tailed storm-petrel <em>(Oceanodroma furcata)</em>,
have ever achieved great enough numbers
to offset the impact of extended daylight.</p>

<p>Past disruptions of seabird populations are
both natural and man-induced; however, the
documentary record is much too fragmentary
to allow us to fully appreciate what has occurred
or what the long-term effect has been.
To give some perspective to the problems associated
with assessing change and attempting
to understand it, some of the indicators of
natural and unnatural change and flux in seabird
populations are reviewed here.</p>

<p>The flux in bird numbers can be related to
the time of day, season of the year, and atmospheric
conditions on a short-term basis. This
sort of flux or apparent flux can easily be explained.
The underlying cause of some of the
longer term flux is not so easily arrived at.
Murie (1959), Gabrielson and Lincoln (1959),
and Sowl and Bartonek (1974) have noted
some of the man-induced changes. These are
also explored to some extent in the species accounts
as they are found to apply.</p>

<p>I sometimes refer to a colony size class<span class="pagenum"><a name="Page_52" id="Page_52">[Pg 52]</a></span>
when discussing the existing data rather than
to an actual population estimate. The size
classes used are defined as follows:</p>


<ul><li>Class I&mdash;less than 100 birds</li>
<li>Class II&mdash;100-1,000</li>
<li>Class III&mdash;1,000-10,000</li>
<li>Class IV&mdash;10,000-100,000</li>
<li>Class V&mdash;100,000-1,000,000</li>
<li>Class VI&mdash;more than 1,000,000</li>
</ul>

<p>The <cite>Dictionary of Alaska Place Names</cite>
(Orth 1967) is the reference for those who wish
to locate some of the less obvious sites. The
<cite>Coast Pilot, No. 9</cite> (U.S. Department of Commerce
1964) is another useful reference.</p>


<h5><em>Species Accounts</em></h5>


<h6>Northern Fulmar <em>(Fulmarus glacialis)</em></h6>

<p>Petrels of a number of species can be found
in the Gulf of Alaska, some of them in great
numbers. Only the northern fulmar breeds
there.</p>

<p>The fulmar is common in the offshore
waters of the northern Gulf of Alaska
throughout most of the year (Isleib and
Kessel 1973). Most authors, including Clark
(1911), one of the earlier ones, who commented
on the distribution of fulmars farther out in
the Gulf, have considered them to be abundant.
Nichols (1927) raised one of the few
voices of apparent dissent; he noted that in
1926 he encountered the largest number of
fulmars (about 800) on 11 July in Shelikof
Strait after he had left the Gulf. During the
summer, fulmars are very common seaward of
Montague Island, particularly to the northeast
of Patton Bay and in the approaches to
Montague Strait. Data derived from FWS
surveys in July and August 1972 showed an
estimated 10,000 fulmars in a stretch of
waters 19 km wide along the east side of Montague
Island (Isleib and Kessel 1973).</p>

<p>Over the Portlock Banks and in Stevenson
Entrance, fulmars sometimes concentrate in
very large numbers, either by themselves or in
company with sooty shearwaters <em>(Puffinus
griseus)</em>. In August 1973, FWS observers
crossing Perenosa Bay saw large numbers of
tube-nosed birds moving northeastward
across the Bay. Although these appeared to
be predominantly shearwaters, there were
also many fulmars. There was a general movement
of birds through Shuyak Strait from
Shelikof Strait into the Gulf of Alaska. It was
not determined whether the fulmars were
moving with the shearwaters or on a regular
feeding flight. Fulmars are often found close
to Afognak Island in the area between Sea
Lion Rocks and Sea Otter Island. Gabrielson
and Lincoln (1959) reported seeing swarms of
fulmars in Marmot Strait and around the
small islands on the north side of Afognak in
early August. Murie (1959) noted fulmars in
Shelikof Strait and again around the Shumagin
Islands. There is nothing in this record
to indicate any change in their distribution at
sea recently.</p>

<p>The Semidi Islands support the Gulf of
Alaska's largest fulmar breeding population,
a Class V colony (U.S. Bureau of Sport
Fisheries and Wildlife 1973). Gabrielson and
Lincoln (1959) considered it to be one of the
four largest colonies in Alaska.</p>

<p>Gabrielson (1940) was told by Captain Sellevold
of the marine vessel <em>Brown Bear</em> that he
thought the birds nested on Sea Otter Island
in Perenosa Bay. Gabrielson also learned that
they probably nested on Sea Lion Rock at the
head of Marmot Strait. In August 1973 I observed
fulmars in close proximity to Sea Lion
Rock. More recently, small numbers of apparently
breeding fulmars have been found in
the Barren Islands (L. W. Sowl, personal observation
and Edgar Bailey, unpublished
FWS report, Anchorage, Alaska). Although
no other colonies are known or suspected, the
evidence suggests the possible existence of
some.</p>

<p>Peterson and Fisher (1955), on noting dark
fulmars between St. Paul and St. George
when only the light morph was present on any
of the colonies in the Pribilofs, expressed no
surprise. They offered the opinion that a
round trip of 960 km to one of the dark morph
colonies in the Aleutians just might be within
the operating range of a fulmar on a 4-day
vacation from nest-tending duties. Using this
as a general yardstick, it appears that the rich
foraging grounds over the Portlock Banks
might also be within the range of breeding fulmars
from the Semidis. The trip up Shelikof
Strait and on to Portlock Bank by way of
Shuyak Strait is only slightly longer than the
one from Chagulak to St. Paul. The feeding
grounds off Montague Island would require a<span class="pagenum"><a name="Page_53" id="Page_53">[Pg 53]</a></span>
1,600-km round trip from the colonies in the
Semidi Islands. Birds from the Barren Islands
and any colonies around Shuyak Island
could easily reach the Montague Island
grounds, but why would they cross the Portlock
Banks to do so?</p>

<p>Fulmar colonies may be found in the Chiswell
Islands. It is also a possibility that the
existence of colonies on islands along the
north coast of Afognak Island will be verified
and that others will be found in the vicinity of
Shuyak Island. Gabrielson and Lincoln (1959)
expressed the opinion that there is almost certainly
a colony on Sutwik Island. If there is
one, however, I did not see it on one quick trip
around the island in 1973.</p>

<p>Gabrielson (1940) expressed surprise at the
size of the Semidi Island breeding colony.
Gabrielson and Lincoln (1959) considered
1911 to be the first time breeding fulmars
were found in the Shumagins. They apparently
based this on two eggs collected
there that year and documented in a plate in
Bent (1964). Other than Gabrielson's opinion,
there is nothing to indicate a major change in
fulmar status during this century. If there has
been a change in status, it has probably been
in the direction of increasing populations.</p>


<h6>Fork-tailed Storm-petrel <em>(Oceanodroma
furcata)</em></h6>

<p>The fork-tailed storm-petrel probably
breeds throughout the Gulf of Alaska. It is
abundant at sea during the summer in most
offshore waters. Murie (1959) described it as
the dominant petrel in the Bering Sea and the
North Pacific.</p>

<p>In view of its wide distribution and apparent
abundance very little is known about
the fork-tailed storm-petrel's breeding colonies.
Friedmann (1935) recorded specimens
and eggs from Kodiak dating back to 1843.
Murie (1959) noted them as nesting on Sanak
Island and stated that they almost certainly
nested in the Shumagins and on other islands
along the Alaska Peninsula. David Roseneau
(Isleib and Kessel 1973) found this storm-petrel
"breeding by the 10,000's" on East
Amatuli Island in the Barren Islands in June
1965. This was subsequently verified in 1974
by Edgar P. Bailey (unpublished report, FWS,
Anchorage, Alaska).</p>

<p>On 2 July 1972, responding to a tip by
James W. Brooks (personal communication),
M. E. Isleib and I anchored at Fish Island in
the Wooded Islands. We did not locate any
storm-petrel burrows, but a steady flow of
storm-petrels passed over the boat throughout
the darkest part of the night. Surveys
conducted at about that time provided an estimate
of 19,000 fork-tailed storm-petrels in
Prince William Sound, primarily in or close to
Montague Strait, and in coastal waters on the
east side of the Sound's outer islands. In this
area Isleib (personal communication) has
noted a general movement of fork-tailed
storm-petrels westward around Montague Island
and into Prince William Sound through
Montague Strait each morning and a corresponding
countermovement each evening. I
conclude that in 1972 there was a Class IV
colony in the Wooded Islands, numbering between
19,000 and 38,000 birds. Additional
colonies will be discovered in a similar manner
as more systematic searches are made.</p>

<p>No colonies were discovered during the 1973
reconnaissance survey of the islands south of
Alaska Peninsula. Working primarily inshore,
FWS investigators encountered very few
storm-petrels during the day. On the night of
14 June, the FWS vessel, <em>Aleutian Tern</em>, responded
to a Mayday call and was either in
transit or participating in rescue operations
from 2245 to 0420 h on the morning of 15
June. During this period numerous fork-tailed
storm-petrels were encountered, particularly
off Cape Wedge on Nagai Island. After we
anchored in Eagle Harbor on Nagai, more
storm-petrels were heard about the vessel.</p>

<p>At about this same date, National Marine
Fisheries Service enforcement officers flying
fisheries patrols observed storm-petrels in
abundance south of the Shumagin Islands
(James Branson, personal communication).
These observations support the belief that
there are probably substantial undiscovered
colonies in the Shumagin Islands.</p>

<p>Fork-tailed storm-petrels are abundant
summer residents in the northern Gulf of
Alaska and the estimate by Isleib and Kessel
(1973) is that populations using the waters off
the North Gulf Coast probably number in the
millions. Certainly the same estimate is valid
for the rest of the Gulf area west of the Chugach
Islands.</p>

<p>The status of these birds relative to their<span class="pagenum"><a name="Page_54" id="Page_54">[Pg 54]</a></span>
historical abundance cannot be derived from
the existing information. There is strong suspicion
that the introduction of fox on many of
the islands in the area during the early part of
this century probably caused a reduction in
their numbers. Murie (1959) said that experience
taught him that wings left from fox kills
or remains of storm-petrels in fox droppings
could be accepted as evidence of the presence
of a colony. Gabrielson and Lincoln (1959) reported
that E. P. Walker visited the Wooded
Islands in 1922 searching for a storm-petrel
colony that had been reported to exist there in
1918. He could not find it even though he
searched diligently. This apparent disappearance
was attributed to the introduction of
fox.</p>

<p>There is another factor to consider, however.
The limited number of specimens now
available from the Gulf of Alaska indicates
that separate subspecies occupy the eastern
and western Gulf of Alaska. The accepted
boundary is somewhere in the vicinity of
Prince William Sound. This is an indication
that there has been a hiatus in this area of
rather long duration. I have speculated that
this sort of break may be in some way related
to the length of day and a period during the
summer when there is little darkness to cover
activities near the colony. Thoresen (1964)
and Cody (1973) have both reported that western
gulls <em>(Larus occidentalis)</em> assemble in
Cassin's auklet colonies on moonlit nights to
prey on arriving adults. It is likely that other
nocturnal species would provoke the same
sort of hunting tactic. A light-related predation
factor implies that the predators rely on
sight. Avian predators are indicated.</p>


<h6>Leach's Storm-petrel
<em>(Oceanodroma leucorhoa)</em></h6>

<p>Even less well understood than the breeding
distribution of the fork-tailed storm-petrel is
that of Leach's storm-petrel.</p>

<p>Bendire (1895) quotes notes from Chase
Littlejohn, who found Leach's storm-petrel to
be an abundant breeder on unspecified small
islands near Sanak in 1894. It greatly outnumbered
the fork-tailed storm-petrel. On his
visit in 1937 Murie (1959) learned that all of
the large colonies of seabirds that had once
existed there were gone. He attributed this to
overfishing and associated perturbation and
to the introduction of fox. No systematic assessment
of seabirds on Sanak has been attempted
since Littlejohn's time.</p>

<p>No Leach's storm-petrel colonies have been
encountered during reconnaissance surveys of
the Gulf of Alaska. Small numbers have been
reported from time to time and while it is very
much less abundant than the fork-tailed
storm-petrel, I expect that it will be found in
small numbers at various places in the Gulf of
Alaska when it becomes possible to make
more thorough searches. It may occur in remote
areas like the smaller islands scattered
throughout the Sandman Reefs&mdash;possibly
even in large numbers. On the basis of the
Sanak record, we must assume that this
storm-petrel has been greatly reduced in numbers,
at least in the western portion of the
Gulf.</p>


<h6>Double-crested Cormorant <em>(Phalacrocorax
auritus)</em></h6>

<p>The white-crested cormorant, the race of
the double-crested cormorant residing in the
Gulf of Alaska, is principally an inhabitant of
the marine environment. This cormorant is a
common, but apparently patchily distributed,
resident throughout the northern and western
Gulf of Alaska.</p>

<p>Gabrielson and Lincoln (1959) thought that
it nested only from Kodiak Island westward
into the Aleutians. However, it probably
breeds from Yakutat Bay westward. Isleib
and Kessel (1973) estimated the abundance of
the double-crested cormorant along the North
Gulf Coast as several thousands, about one-tenth
as common as the pelagic cormorant. It
is the third most abundant of the four cormorant
species nesting in the area. It occurs
as scattered inclusions in many colonies
throughout the area, and at least in the Shumagin
Islands, even occurs in some colonies
by itself.</p>

<p>There are no data on which to base an estimate
of any change in status. It probably is
not much affected by many of the naturally
occurring perturbations.</p>


<h6>Brandt's Cormorant <em>(Phalacrocorax
penicillatus)</em></h6>

<p>On 22 July 1972, 13 Brandt's cormorants (4
sitting on nests) were found at Seal Rocks in<span class="pagenum"><a name="Page_55" id="Page_55">[Pg 55]</a></span>
Hinchinbrook Entrance, Prince William
Sound (Isleib and Kessel 1973). Two years
later I positively identified two individuals in
breeding plumage among a mixed group of
cormorants in the Chiswell Islands west of
Seward. Are these recent range extensions?
Possibly, but I propose an alternative explanation.</p>

<p>Palmer (1962) showed the distribution of
this cormorant as breeding north to Puget
Sound and as a straggler north to Forrester
Island, Alaska. This viewpoint is shared by
the American Ornithologists' Union (1957),
which regards the bird as casual as far north
as Forrester Island, where this species was
collected by Willet (1918).</p>

<p>Let us look at the other record, the one that
is not supported by specimens. Bent (1964)
thought of Brandt's cormorant as a breeding
resident of Forrester Island. Gabrielson and
Lincoln (1959) admonished bird observers to
be on the lookout for this particular cormorant
in the vicinity of Ketchikan and
Prince of Wales Island. Brandt's cormorant
also appears on the bird list for the Kodiak
National Wildlife Refuge as an accidental
visitor.</p>

<p>Early observers like Bent were explorers.
They carefully examined and made notes on
all the birds they saw because there was always
a chance of a new discovery. It is also
very probable that Bent paid particular attention
to the cormorants when he was at a place
like Forrester Island. He would have undoubtedly
been very interested in trying to
confirm the presence of the now extinct
Palla's cormorant <em>(P. perspiculatus)</em>, as he
must have been aware of Schlegel's (1862-64)
list of the birds in the Dresden Museum since
Willet (1914) had recently referred to it. The
staffs for the Kodiak and Aleutian Islands
National Wildlife refuges have included some
very careful observers, such as Frank Beals.
These men would have noticed the difference
if a new bird such as Brandt's cormorant was
seen, verified the sighting visually, and then
noted it in their field diaries. They would not
have bothered to develop the type of proof
needed for an undisputable record, but the
bird would have appeared in the refuge bird
list (as it does).</p>

<p>The outside coasts of the Alexander Archipelago,
Kenai Peninsula, and the Islands of
the Kodiak Archipelago impose some logistical
requirements which discourage all but
the most determined birders. Not many have
been able to reach more than very limited segments
of the entire coast. Given the vast distances
involved, few of the FWS vessels passing
through the area have had the time to
thoroughly examine any cormorant colonies
or roosts bird by bird. Even for those who
pause, the ever present swells and the constant
chop of the summer westerlies make
positive identification difficult.</p>

<p>It is possible that Brandt's cormorant has
been in the area in small numbers for a long
time, either regularly or intermittently. It
could have escaped observation because of the
conditions described above. This species may
be there as a relict, as a pioneer, or only because
surplus birds are being pushed into
marginal habitat by population pressures on
their main range to the south.</p>


<h6>Pelagic Cormorant <em>(Phalacrocorax pelagicus)</em></h6>

<p>The pelagic cormorant is the most abundant
of the four cormorants residing in the
Gulf of Alaska. It is found throughout coastal
Alaska south of the Bering Strait and even in
some colonies in the southern Chukchi Sea.</p>

<p>Cormorants have a certain invisibility
which is brought about by their universal
presence. This blindness appears to have affected
everyone, even the earliest observers.</p>

<p>The earliest accounts provide a composite
picture of the distribution and abundance of
the pelagic cormorant which is very similar to
that encountered today. In southeastern
Alaska, beginning at the eastern edge of the
area under discussion, the pelagic cormorant
was pictured as the sole resident cormorant.
However, we know from Willet's collection of
a Brandt's cormorant at Forrester Island that
this might not be quite true. From Yakutat
Bay westward into the Aleutians this species
coexisted with the double-crested cormorant.
In the Western Aleutians there is some disagreement,
but in general it appears to have
been accepted that the red-faced cormorant
occurred there along with pelagic and possibly
double-crested cormorants. In the
Bering Sea this species coexisted with the red-faced
cormorant.</p>

<p>A number of recent authors (Gabrielson<span class="pagenum"><a name="Page_56" id="Page_56">[Pg 56]</a></span>
1940, 1944; Murie 1959; and others) have considered
the pelagic cormorant to be the most
widely distributed and abundant of the four
species found in Alaska. Since the modern picture
fits, in a general way at least, it would be
easy to conclude that the species enjoys an
unchanged status. There is just a faint suggestion
that this may not be true.</p>

<p>Dement'ev and Gladkov (1966) refer to a
great die-off of pelagic cormorants referred to
earlier, in the Commander Islands. Stejneger
(1885) enlarges on this disaster. It is true that
Stejneger visited these islands a relatively
short time after the die-off, but he reported
that even though the pelagic cormorants were
increasing, "people having seen their former
multitude think that there is no comparison
between the past and the present." Murie
(1959) thought that the pelagic cormorant,
while numerous, was outnumbered by the red-faced
cormorant in the Aleutians. More recently
there has been the rapid eastward expansion
of the red-faced cormorant. Although
it is not possible to determine what the real
status of the pelagic cormorant is relative to
its past status, I conclude that during this
century its status relative to that of the red-faced
cormorant has declined.</p>


<h6>Red-faced Cormorant <em>(Phalacrocorax urile)</em></h6>

<p>The red-faced cormorant, in spite of superficial
similarities to the pelagic cormorant,
just does not look the same to an experienced
observer. However, it would have been possible
for inexperienced observers in the days
before modern optics to overlook the differences.
The problem was further compounded
by the "invisibility" of the ubiquitous cormorants
referred to earlier. Apparent absences
or blank spots in their range may not
have been real.</p>

<p>Dement'ev and Gladkov (1966), reporting
on the Russian record, stated that the red-faced
cormorant was common in the Commander
Islands during the last part of the
19th century and into the early part of the
20th. Older authors had also reported it from
Kamchatka and the Kurile Islands. Now, according
to Dement'ev and Gladkov, it is an
uncommon breeder on Mednyi Island in the
Commander Islands and occurs only as an
autumn visitor to some of the southern Kurile
Islands.</p>

<p>Turner (1885) reported that the double-crested
cormorant was abundant in the Near
Islands and that the pelagic cormorant was
common, but makes no reference to the red-faced
cormorant. One specimen of the latter in
the Leningrad Academy of Science was taken
at Attu on 16 September 1844 (Gabrielson
and Lincoln 1959), which indicates that they
were probably present during the period reported
on by Turner and, therefore, relatively
uncommon. Clark (1911) identified red-faced
cormorants only a few times and in the Aleutians
only once, near Agattu. Dall (1874)
noted two red-faced cormorants collected at
Amchitka but he (Dall 1873) apparently did
not see any east of Unalaska.</p>

<p>Nelson (1887) apparently found red-faced
cormorants breeding on the Siberian and
Alaskan mainlands at either side of Bering
Strait, but Bailey (1948) searched for some
sign of their presence and found none. Nelson
(1887) also reported the red-faced cormorant
from St. Matthew and St. Lawrence islands in
the northern Bering Sea and from St. Michael
and Nelson Island on the Alaskan coast.
Gabrielson and Lincoln (1959) pointed out
that it has not been found breeding north of
the Pribilofs since then. Friedmann (1934) provides
support for Nelson by reporting red-faced
cormorant bones from archeological
sites on St. Lawrence. Gabrielson and Lincoln
(1959) cited two red-faced cormorants in the
Leningrad Academy of Science which were
collected in the Pribilofs in 1843. Dall and
Bannister (1869) reported them to be plentiful
on St. George Island. Baird (1869) also noted
their presence in the Pribilofs.</p>

<p>Bent (1964) makes no mention of seeing the
red-faced cormorant in the Aleutians. He
gives their breeding range as the Bering Sea
region, the Pribilof Islands, and perhaps the
western Aleutians, the Commander Islands,
and the coast of Siberia north of North Cape.
The American Ornithologists' Union (1931)
gave their breeding range as the Pribilof Islands,
the Commander Islands, and Siberia
north to North Cape.</p>

<p>Murie (1959) found a colony of between
4,000 and 5,000 red-faced cormorants nesting
on Amak Island in 1925. In 1936 he was surprised
to find that the red-faced cormorant
was the most abundant breeding cormorant in
the Aleutian Islands. Pelagic cormorants still<span class="pagenum"><a name="Page_57" id="Page_57">[Pg 57]</a></span>
appeared to be most numerous, but there were
large numbers of nonbreeding birds. In 1936
he located "a good sized colony" of red-faced
cormorants at Unga in the Shumagin Islands.
He found about 300 birds starting their nests
on 16 May.</p>

<p>In August 1946 Gabrielson (Gabrielson and
Lincoln 1959) visited the colony at Delarof
Harbor, Unga, where several thousand cormorants
were observed. From a number of
small samples he estimated that the red-faced
cormorants outnumbered pelagic cormorants
five to two. In 1973 I observed about 2,000
cormorants, mostly red-faced, in this colony.
Gabrielson also located them at two other
sites in the Shumagins and at Aghiyuk Island
in the Semidi Islands.</p>

<p>Howell (1948) noted only double-crested cormorants
at Double Island, Kodiak. Shortly
after that the leaflet, <cite>Birds of the Kodiak Island
National Wildlife Refuge</cite> (first issued in
1955), listed red-faced cormorants as common
summer residents. The red-faced cormorant
was next found at Katchemak Bay about
1963. Isleib (Isleib and Kessel 1973) first
noticed red-faced cormorants wintering in
Prince William Sound in 1969. In July 1972
Isleib and Sowl had found a colony containing
75 nests at Point Elrington at the western approach
to Prince William Sound. By 1974
Isleib and Haddock (unpublished data, FWS,
Anchorage, Alaska) found them east of the
Copper River Delta at Wingham Island.</p>

<p>The relatively rapid expansion of the range
and apparent population size of the red-faced
cormorant is remarkable. But has this been a
real expansion into vast stretches of new territory?
The record in the literature which I
have summarized shows, I think, something
else. We can demonstrate a historical range
for the red-faced cormorant that extends on
the Asiatic Coast from North Cape, Siberia,
south to the Kurile Islands, the entire Aleutian
Arc including the Commander Islands,
all the Bering Sea islands north to Bering
Strait, Norton Sound, Nelson Island, and the
islands south of the Alaska Peninsula at least
as far east as Kodiak Island. The recently
occupied coast from Cook Inlet to the Copper
River may represent a real range extension.
The breeding range of this species at the
present time does not include parts of its historical
range west of the Commander Islands
or north of the Pribilof Islands.</p>

<p>The fragmentary record appears to show a
long-term perturbation in the range and populations
of the red-faced cormorant that covers
at least 100 years. I believe that we are probably
seeing a recovery of lost range and a return
to something resembling a former distribution
and abundance.</p>

<p>What caused the perturbation? I am not
prepared to answer this question, but there
are two occurrences which I find suggestive.</p>

<p>It is interesting to note (Dement'ev and
Gladkov 1966) that on the Commander Islands
the red-faced cormorant was most abundant
during the first 50-odd years after the
pelagic cormorants had been wiped out in the
winter of 1876-77. Perhaps some clues are to
be found in the interactions between these
similar species.</p>

<p>It does not appear that the introduction of
fox could have been a causative factor. The
first observations of population expansion
were noted almost concurrently with the heyday
of the fox-farming industry. Because of
its choice of nesting habitat (very steep cliffs),
this cormorant would not have been affected
by predators except for the one that went into
a very rapid population decline at a time that
would fit&mdash;the Aleut.</p>

<p>Jochelson (1968) and Hrdlicka (1945) summarized
references to Aleut clothing in the
diaries and reports of early Russian visitors
to the Aleutian Islands. Evidently Aleut
women sometimes wore a long, robe-like
parka made of harbor seal <em>(Phoca vitulina)</em>
skins or, for women of high rank, parkas made
of sea otter <em>(Enhydra lutra)</em>. The men in almost
all reports were said to have worn bird-skin
parkas; puffins and guillemots appear to
have been preferred, but cormorants were
sometimes used. It took about 40 puffin skins
to fabricate a parka and a man evidently
needed from one to three of these garments
each year.</p>

<p>Sea otter populations were drastically reduced
by Russian hunters. Rats were introduced
to the Aleutians very early during the
Russian period and must have had a substantial
impact on populations of tufted puffins
and guillemots. The introduction of fox would
have had a further impact on burrow-nesting
birds. Turner (1885) noted that Aleuts in the
Near Islands kept the fox confined to Attu so<span class="pagenum"><a name="Page_58" id="Page_58">[Pg 58]</a></span>
that they could keep the fox away from the
birds on Agattu. This is evidence of an Aleut
recognition of serious competition. Could cormorants,
particularly red-faced cormorants,
have been preferred sources of fiber? Were
Aleuts forced to rely more heavily on cormorant
skins as puffin and guillemot numbers
were reduced by rats and fox and sea otters
by men?</p>

<p>Whatever the cause and effect, the status of
red-faced cormorants now appears to be
better in the Gulf of Alaska than for at least
the last 100 years.</p>


<h6>Glaucous-winged Gull <em>(Larus glaucescens)</em></h6>

<p>The glaucous-winged gull is apparently one
of the more successful seabirds breeding in
the Gulf of Alaska. While it is outnumbered
(both locally and in total abundance) by the
black-legged kittiwake, it is generally the
most commonly seen and most uniformly distributed
gull in the Gulf of Alaska. Murie
(1959) called it the common breeding gull
about the Alaska Peninsula. Cahalane (1943,
1944) considered it to be numerous to abundant
around Kodiak and in the Shelikoff
Strait area. Gabrielson (1944) reported that it
could be seen in small numbers everywhere.
Most recently, Isleib and Kessel (1973) reported
it to be an abundant resident in the
north Gulf Coast area. My own experience
would confirm these observations.</p>

<p>This gull appears to use a wider variety of
nesting sites than some others (Gabrielson
and Lincoln 1959). Except where man's activities
have created new food sources, there appears
to be a close link between the location of
glaucous-winged gull colonies and those of
murres, kittiwakes, and cormorants. Swartz
(1966) found that during the breeding season
glaucous-winged gulls at Cape Thompson derived
almost all of their food from murre eggs
and chicks. I have noted small numbers of
these gulls nesting, usually on turf near the
tops of cliffs, in most colonies of favored prey
species.</p>

<p>The glaucous-winged gull is the principal
scavenger throughout much of coastal south-central
Alaska. This has sometimes resulted
in the development of large concentrations
near canneries and, more recently, near
dumps.</p>

<p>Two glaucous-winged gull concentrations
stand out in the northern Gulf of Alaska. One
of these is on Egg Island at the western end of
the Copper River Delta. Patten (1976) estimated
that this colony contained 10,000-12,000
gulls in 1975. At times it appears to
spread onto nearby Hinchinbrook Island.
M. E. Isleib (personal communication) has estimated
its size as high as 25,000 gulls. The
other large concentration is on the Susitna
Flats across Cook Inlet from Anchorage. This
colony, or colony cluster, may be larger than
the one at Egg Island. There are no other
known colonies even approaching these in
size. Most colonies range between a few pairs
and 2,000-3,000.</p>

<p>Glaucous-winged gulls do not appear to
have had any great changes in population
that can be detected from the literature.
There have almost certainly been local fluctuations
in the number of breeding birds as
food supplies, such as canneries and dumps,
have appeared or disappeared in an area.
Long-term changes in salmon runs have undoubtedly
had an impact as well. One other
change, the reduced level of egging, has undoubtedly
had an effect also. Along the
Alaska Peninsula and in the Shumagin Islands,
cannery workers of Filipino heritage
and fishermen who have a strong Aleut heritage
still harvest gull eggs for food. However,
this activity is much reduced from what it
must have been.</p>


<h6>Herring Gull <em>(Larus argentatus)</em></h6>

<p>The herring gull is a resident of Upper Cook
Inlet and is found up and down the coast from
Prince William Sound to the Alaska Peninsula.
Not too much was learned about it during
the recent FWS reconnaissance. Williamson
and Peyton (1963) reported the interbreeding
of herring gulls and glaucous-winged
gulls in this area. This interbreeding has resulted
in a situation in which assignment of
these gulls to one group or another in the field
can be rather arbitrary. The result has most
often been that field observers tend to lump
them with glaucous-winged gulls unless their
herring gull characteristics are obvious.
Specimens collected by Williamson and Peyton
(1963) indicate that herring gulls have the
edge in numbers in Upper Cook Inlet.</p>

<p><span class="pagenum"><a name="Page_59" id="Page_59">[Pg 59]</a></span></p>


<h6>Black-legged Kittiwake <em>(Rissa tridactyla)</em></h6>

<p>The black-legged kittiwake is the most
abundant gull in the northern and western
Gulf of Alaska. Colonies of this species can be
found throughout the entire area, and range
in size from a few pairs (Class I) to more than
100,000 birds (Class V). They may be found in
essentially pure colonies, but are often found
sharing colonies with murres.</p>

<p>The center of abundance for breeding black-legged
kittiwakes in the Gulf of Alaska is in
the Semidi Islands, where Palmer Sekora
(U.S. Bureau of Sport Fisheries and Wildlife
1973) estimated that there were 426,000
breeding kittiwakes in 1972. He located kittiwake
colonies at eight sites, ranging in size
from 1,000 to 109,000 nesting birds. The size
of the average colonial site was 27,000 birds.
Ten sites were Class IV in size and one was a
solid Class V.</p>

<p>The easternmost known colony in the northern
Gulf of Alaska is at Wingham Island. Up
to 1973, 22 colonies had been located in Prince
William Sound. The largest of these contained
only 5,636 nests in 1972 (Isleib and Kessel
1973). Class IV or larger colonies are found at
Cape Resurrection, the Barren Islands, Chisik
Island, Boulder Bay and Cape Chiniak on
Kodiak Island, and at Delarof Harbor and the
Haystacks in the Shumagin Islands. It is interesting
to note that Gabrielson (1940) considered
Whale Island to be one of the largest
known kittiwake colonies in Alaska. He stated
that there were many thousands of pairs extending
over a mile or more of cliff. He saw a
second site which he did not visit but looked
equally large. A photograph in an article by
East (1943) also indicated the presence of a
large colony. C. J. Lensink (personal communication)
estimated that there were about
100,000 kittiwakes in the colony in 1956.
When last visited by Vernon Berns (personal
communication), this colony contained only
3,000 birds. It is also of interest that Gabrielson
(1940, 1944) did not notice either the kittiwakes
or the murres now breeding on Nord Island
in the Barren Islands or the kittiwakes
on East Amatuli Island.</p>

<p>Whale Island and possibly the colonies in
the Barren Islands give evidence of local
population fluctuations, but for the most part
I have not found an indication of a major perturbation
over the past 40 years. Before 1936,
the record is too fragmentary to allow an
assessment.</p>

<p>One of the interesting aspects of kittiwake
ecology in the Gulf of Alaska is the common
occurrence of breeding failure. David Snarski
(December 1943 Quarterly Progress Report,
Alaska Cooperative Wildlife Research Unit,
University of Alaska) observed breeding failure
on colonies in the Tuxedni National Wildlife
Refuge in 1970 and 1971 and obtained circumstantial
evidence of another failure in
1972. In 1973 all of the breeding cliffs were
occupied and nesting was successful. Whatever
the cause of these periodic failures, they
do not yet appear to have had a permanent
impact that we are able to measure.</p>


<h6>Red-legged Kittiwake <em>(Rissa brevirostris)</em></h6>

<p>Red-legged kittiwakes are not now known
to breed in the western Gulf of Alaska. Turner
(1886) stated that he saw a few at Sanak in
1878. We also have Stejneger's (1885) statement,
that "red-legged" kittiwakes nest from
Bering Island to Sanak. Friedmann (1937) reported
two humeri from Kodiak Island
middens. During the summer of 1976, two
birds were observed off Kodiak Island by
Irving M. Warner (personal communication),
and one at 158&deg;W and 54&deg;30'-54&deg;20'N south
and east of the Shumagin Islands (Patrick J.
Gould, personal communication).</p>

<p>Turner (1885) listed the red-legged kittiwake
as abundant and breeding in the Near
Islands. Turner (1886) also stated that he had
seen quite a number about a cliff back of the
village on Akutan Island in 1878. He added
that to the westward this kittiwake was more
abundant than the black-legged kittiwake.
Murie (1959) expressed the opinion that
Turner had confused the short-billed gull with
the "short-billed" kittiwake. Clark (1911) also
reported that he had seen the red-legged kittiwake
in small numbers near Unalaska and
that they became progressively more common
west to the Near Islands. Nelson (1887) reported
seeing large numbers of red-legged
kittiwakes at Unalaska. Murie (1959) and
Gabrielson (1940, 1944) did not see any red-legged
kittiwakes in the Aleutian Islands. The
species has recently been discovered breeding
at Buldir and Bogoslof islands (G. Vernon<span class="pagenum"><a name="Page_60" id="Page_60">[Pg 60]</a></span>
Byrd, personal communication).</p>

<p>Is it possible that we have here another
species which is exhibiting a response to some
unknown long-term perturbation? The suggestion
that such an event has occurred is
faint, but it is there. Do we have in the red-legged
and black-legged kittiwakes an
example of yet another congener pair that has
been affected by some perturbation in which
one was affected positively and the other
negatively? Clark (1911) reported small numbers
of black-legged kittiwakes to go with
large numbers of red-legged kittiwakes in the
Near Islands, which is the reverse of the current
situation.</p>


<h6>Arctic Tern <em>(Sterna paradisaea)</em></h6>

<p>Gabrielson and Lincoln (1959) attribute to
the Arctic tern the most extensive range of
any Alaskan water bird. It is found in suitable
habitat everywhere north of Tracy Arm in
Southeastern Alaska. Murie (1959) stated
that he found it nesting at suitable sites
everywhere he went. Isleib and Kessel (1973)
considered it to be an abundant breeder in
Prince William Sound and along the northern
Gulf Coast.</p>

<p>The Arctic tern was observed in FWS aerial
surveys in Prince William Sound, and surveys
in July and August 1972 provided an estimate
of 45,000 terns in the Sound (Isleib and Kessel
1973). On the other hand, tern colonies were
located only rarely in the FWS colony surveys
before 1975. This is, however, a reflection of
the equipment and methods used and not of
the abundance of terns.</p>

<p>From the fragmentary data available, it is
not possible to detect changes in Arctic tern
status at the present time. We have to assume
that the widespread introduction of fox had at
least local impact. Although this tern uses a
wide variety of nesting sites, it tends to nest
on flat sites where access by mammalian
predators is easy.</p>


<h6>Aleutian Tern <em>(Sterna aleutica)</em></h6>

<p>No Aleutian tern colonies were discovered
in the Gulf of Alaska area during FWS colony
surveys in the early 1970's. This is again a reflection
of the fact that surveys were not designed
to locate tern colonies. Aleutian terns
were encountered at least twice, once during
late March 1972 in Hawkins Cutoff, Prince
William Sound, and again when two birds
were noted offshore from the Katmai National
Monument on 30 May 1973 (L. W. Sowl, personal
observations).</p>

<p>The type specimen of the Aleutian tern and
a single egg were collected at Kodiak Island
on 12 June 1868 by Bischoff (Coues 1874).
Fisher (Gabrielson and Lincoln 1959) collected
four more eggs in 1882. The bird was not
found breeding there until Howell (1948)
found a colony of 50 pairs at Bell's Flats in
1944. Walker (1923) found them nesting on
the Situk River, Yakutat, in 1917 and shortly
thereafter saw them at the Alsek River Flats.
He also reported that D. H. Stevenson of the
Bureau of Biological Survey had told him that
they nested on the Isanotski Islands at the
end of the Alaska Peninsula. This latter report
was the only one from the Aleutian Island
chain for many years. Isleib and Kessel
(1973) considered it an uncommon local
breeder in the northern Gulf of Alaska. Isleib
estimated its population at a few hundred pairs
on the Copper River Delta in May 1973 and
300-500 birds in June 1970. He also reported
that they appeared more or less regularly near
Controller Bay and off the Situk River.</p>

<p>In recent years Aleutian terns have been
seen with increasing frequency in many places
in western Alaska and the Aleutian Islands.
This is probably partly due to the increasing
level of field work. At Amchitka Island the
several colonies that have been found in recent
years are almost certainly exhibiting a
response to the removal of fox from the
island.</p>

<p>Although there is no way of determining
what the past status of the Aleutian tern has
been in the Gulf of Alaska area, it has been
there in small numbers since it was first discovered
on Kodiak. It has probably not been
abundant at any time and may have suffered
a long-term decline brought about by the introduction
of fox.</p>


<h6>Common Murre <em>(Uria aalge)</em></h6>

<p>The common murre is resident in the northern
and western Gulf of Alaska from Pinnacle
Rock, Kayak Island, westward. East of Cook
Inlet colonies are located at Wingham Island,
the Martin Islands, Middleton Island, Por<span class="pagenum"><a name="Page_61" id="Page_61">[Pg 61]</a></span>poise
Rock in Hinchinbrook Entrance, Barwell
Island/Cape Resurrection, the Chiswell
Islands, the Barren Islands, and Chisik
Island.</p>

<p>For some reason, the islands of the Kodiak-Afognak
Archipelago do not host any known
major murre colonies. There is also a rather
large gap between the Chisik Island colony
and the next major colony at Oil Creek west of
Puale Bay. Directly west of Oil Creek is another
colony at Cape Unalishagvak. Both of
these latter colonies are Class V and they are
the first colonies of this size to be encountered
in the Gulf of Alaska. West of these colonies
the next large colony is at Atkulik Island. To
the south, midway between the last-named
colonies, lies the major composite murre
colony in the Semidi Islands. These sites
make up the only Class VI colony in the Gulf
of Alaska. Westward, the next major colony,
a Class V, is at Spitz Island south of Mitrofania
Island. In the Shumagin Islands one
Class V colony is at Karpa Island, and lesser
colonies with large murre components are
found at the Haystacks, Castle Rock, and
Bird Island. Only minor murre colonies are
found between the Shumagin Islands and the
end of the Alaska Peninsula.</p>

<p>Gabrielson and Lincoln (1959) were aware
only of the colonies at Cape Resurrection
(which Gabrielson considered to be large), at
the Chiswell Islands, and at Chisik Island for
the area from Cook Inlet east. Gabrielson
visited the Barren Islands on 13 June 1940
and apparently did not notice the present
murre colonies, both Class IV, at East Amutuli
(an island which he visited) and Nord
Island.</p>

<p>Gabrielson (Gabrielson and Lincoln 1959)
found a few small colonies at Kodiak, mostly
on small offshore islands. Gabrielson found
common murres to be abundant in the Semidi
Islands and stated that there were no notable
colonies in the Shumagins, although on his return
to the Shumagins in 1949 he did find a
fairly large colony at the Haystacks. That size
description would fit the colony that is there
now. He obviously did not see the other colonies.
Rausch (1958) reported murres from
Middleton Island.</p>

<p>There is quite a difference between the distribution
of murres as we know it today and
the way Gabrielson and Lincoln pictured it.
Why does this difference exist? There are two
possible answers: either the number of colonies
has increased, or the coverage of colony
locations has improved. The latter case, at
least, is established. I must confess to being
puzzled by the way Gabrielson was able to
move about close to what are now known to be
sizeable colonies without seeing them, those
in the Barren Islands and the Shumagin Islands
in particular. Perhaps this represents
the vague outlines of yet another population
change.</p>

<p>The center of abundance for murre distribution
in the Gulf of Alaska today is from Paule
Bay west to eastern Shumagin Islands. The
Semidi Islands are the heartland of this area
of maximum abundance. We have no definitive
data on species composition of these colonies.
Common murres undoubtedly dominate
in most of the colonies; the only ones where we
know of a sizeable thick-billed murre component
are in the Shumagin Islands.</p>


<h6>Thick-billed Murre <em>(Uria lomvia)</em></h6>

<p>Thick-billed murre population information
cannot be separated from that of the common
murre on the basis of existing data. A direct
assessment of present-day status is not possible.
After reviewing what we know about
their distribution, I suggest a way to examine
the question indirectly.</p>

<p>The thick-billed murre is found in colonies
with the common murre from Middleton Island
westward; Rausch (1958) noted about
400 murres at Middleton Island and observed
that the thick-billed murre outnumbered the
common murre by several times. Isleib and
Sowl (FWS, unpublished data) saw a thick-billed
murre mixed with common murres at
Porpoise Rock in July 1972. Isleib and Kessel
(1973) expressed the opinion that small numbers
of thick-billed murres will be found in
most common murre colonies in the northern
Gulf of Alaska when it is possible to survey
these colonies in detail. Karpa Island had a
significant component of thick-billed murres
in June 1973, and they constituted 40% of the
colony at the Haystacks (L. W. Sowl, unpublished
data).</p>

<p>Bent (1963) reported that many thick-billed
murre eggs have been taken by collectors at
Round Island in the Shumagin Islands. Dall
and Bannister (1869) reported a thick-billed<span class="pagenum"><a name="Page_62" id="Page_62">[Pg 62]</a></span>
murre that was taken at Kodiak in 1867.</p>

<p>The Gulf of Alaska is at the periphery of the
breeding range of the thick-billed murre.
While it probably occurs in mixed colonies
with the common murre throughout this area,
the thick-billed murre is much less abundant.
Occasionally in the Gulf of Alaska, a colony
will be occupied predominantly by the thick-billed
murre. Gabrielson and Lincoln (1959)
noted that the thick-billed murre outnumbered
the common murre in many colonies in
the Aleutians and that it became progressively
more common at higher latitudes.</p>

<p>We have almost no data relative to the
species composition of murre colonies in the
Gulf of Alaska. Until we do it will not be possible
to fully understand the population
status of the thick-billed murre. It appears
that changes in the species composition of
murre colonies in the Bering Sea may be an indicator
of perturbation. The data for the Gulf
of Alaska are still too fragmentary to provide
any indication of whether or not the same indicator
would work there. Close monitoring of
the Shumagin Islands colonies over a number
of years might produce the answer.</p>

<p>Earlier in this paper I noted the dramatic
changes in species composition of murre colonies
on Walrus Island. Gabrielson and Lincoln
(1959) also commented on this well-documented
and anything but static situation. Investigators
who visited this island during
1976 reported seeing no murres on the island
and only small numbers on offshore rocks.
James Bartonek (personal communication)
said that this situation has prevailed for
several years.</p>

<p>There is an indication that a similar population
fluctuation and change in species composition
of murre colonies have also occurred
on St. Matthew Island. Bent (1963) found
mostly common murres and few thick-billed
murres at St. Matthew. Hanna (1916) saw
only thick-billed murres. Later, Gabrielson
(1941) found this to be true in 1940.</p>

<p>Dramatic fluctuation in murre populations
may be common and, at least in some cases,
the two species may be affected differently.
Perhaps this phenomenon has potential for
providing us with an indicator of some natural
perturbations.</p>

<p>Peterson and Fisher (1955) expressed the
opinion that thick-billed murres arrived at the
nesting ledges later than the common murre
and had to take the sites that were left. Tuck
(1960) reported data from the western Atlantic
showing that thick-billed murres arrive
later than common murres. On the other
hand, Belopol'skii (1961) reported data showing
that the two species arrive on breeding
colonies in East Murman simultaneously. At
Cape Thompson, Swartz (1966) found that
thick-billed murres arrived about a week before
common murres. The date of arrival,
while perhaps a contributing factor, is probably
not decisive. Interspecific competition of
another sort is indicated.</p>

<p>In mixed murre colonies where there are
large numbers of common murres, this species
occupies the choice nesting sites. Thick-billed
murres are usually left with the narrower
ledges while the common murres occupy the
longer, broader ledges (Belopol'skii 1961). The
broader ledges have lower chick and egg mortality
(Spring 1971). Spring also noted that
thick-billed murres are excluded from the centers
of mixed colonies. Johnson (1938) found
that this contributes to higher losses of eggs
to predators and to the loss of other social
benefits of occupying the colony center (Johnson
1941).</p>

<p>Kozlova (1961) said that during the occupation
of a colony there is a sharp competitive
struggle between the two species. In the end
thick-billed murres are pushed out to the periphery
of the colonies or left with narrow
ledges or other equally unfavorable sites.
Spring (1971) studied the functional anatomy
of both species and concluded that the common
murre is more successful in these encounters
because it has a more upright gait and
greater agility than the thick-billed murre.</p>

<p>It follows that in a portion of their respective
ranges, where the two species overlap and
where there is an equal chance that either
common murres or thick-billed murres will
dominate a given colony, the common murre
dominates. I conclude from this that where
there are dramatic changes in species composition
of murre colonies, such as at Walrus
Island, it is probably because the common
murre has been greatly reduced in numbers at
the colony.</p>

<p>Spring (1971) concluded that the common
murre is well adapted to pursuit and capture
of pelagic fishes and that the thick-billed<span class="pagenum"><a name="Page_63" id="Page_63">[Pg 63]</a></span>
murre is better adapted for deep diving and
the capture of benthic fishes and pelagic and
benthic invertebrates. Having greater latitude
for food selection, the thick-billed murre
would have a greater tolerance for ecological
perturbations affecting the available food
supply. The common murre has an advantage
when pelagic fishes are available but cannot
switch to the other foods as readily as can the
thick-billed murre. The low density of pelagic
fishes in high arctic areas probably also accounts
for the greater success of the thick-billed
murre at higher latitudes relative to
common murres.</p>

<p>Belopol'skii (1961) presented data from
East Murman which indicates that the common
murre restricts its diet almost entirely to
a small number of fish species. Swartz (1966)
found strong indications that there were significant
differences in the food preferences of
the two species of murres. Thick-billed murres
made much greater use of invertebrates.
B&eacute;dard (1976) asserted that it is well known
that the common murre is quite partial to zooplankton.
So again the issue is not clear-cut.</p>

<p>The situation is, of course, much more complex
than I have portrayed it. Nonetheless, I
think that it offers potential for use as a tool
in assessing population change and perturbations
in the food supply which should be
studied quite closely.</p>


<h6>Pigeon Guillemot <em>(Cepphus columba)</em></h6>

<p>Gabrielson and Lincoln (1959) noted that
the pigeon guillemot was one of the most
regularly observed birds in Alaskan waters. It
is found everywhere throughout the northern
and western Gulf of Alaska area, with only a
few understandable and relatively small
blanks, such as in the silty waters of Upper
Cook Inlet. Because it obviously lacks the
breeding murres' need for close contact with
its nearest neighbors, it is able to exploit the
available nesting habitat to the fullest. It
seems that literally every bit of suitable nesting
habitat is normally occupied.</p>

<p>Because of the dispersed way in which it
breeds and because it does much of its feeding
in the onshore zone (which is hazardous for
boats) the pigeon guillemot is an almost impossible
species to inventory by standard
methods.</p>

<p>There is no evidence that the pigeon guillemot
has been greatly affected by any major
perturbation. Because of its choice of nesting
habitat, it is probably subject to the attack of
only one egg predator, the rat. Because of its
loose social structure and the way it selects
nesting sites, eggs and young do not sustain
loss from panic flights. Its dispersed distribution
should insure that man-made impacts
such as oil spills will have limited impact.</p>

<p>The population levels of the pigeon guillemot
are probably relatively very stable. The
widespread introduction of the rat to most of
its nesting range undoubtedly had impact,
but this impact has gone undocumented. It
would be interesting to follow the response of
guillemot populations on islands where rats
had been totally removed, if that ever becomes
more than a dream.</p>


<h6>Marbeled Murrelet <em>(Brachyramphus
marmoratus)</em></h6>

<p>The marbled murrelet apparently breeds
throughout most of the northern and western
Gulf of Alaska. This apparently is a necessary
condition because to date, at least in this part
of Alaska, we can only guess where and under
what conditions this murrelet breeds.</p>

<p>In some relatively sheltered waters like
Prince William Sound, where marbled murrelets
were estimated to number about 250,000
in 1972 (Isleib and Kessel 1973), they are the
most abundant seabirds. We know from
Dixon (1908) and Grinnell (1910) that this has
been so in Prince William Sound since the beginning
of the century. We know also that the
type specimens came from there as well
(Stresemann 1949), which is not necessarily
an indication of abundance but is suggestive
of their abundance relative to species not
collected.</p>

<p>Gabrielson (Gabrielson and Lincoln 1959)
found marbled murrelets common near Yakutat,
in Prince William Sound, in Resurrection
Bay, and at Kodiak, and reported seeing them
at the Chiswell Islands and at Chignik and
Pavlof Bay on the Alaska Peninsula. Cahalane
(1943, 1944) found them to be common in
Kupreanof Strait, and along the Alaska
Peninsula north of Katmai Bay. Murie (1959)
found them all along the Alaska Peninsula.
My own field notes from 1973 indicate that
the only place where they were common along
the Alaska Peninsula was at Wide Bay.</p>

<p><span class="pagenum"><a name="Page_64" id="Page_64">[Pg 64]</a></span></p>

<p>We can sample marbled murrelet numbers
by using standard transect methodology;
however, I have some very serious reservations
about our ability to convert these data
into a population estimate. This is not an unusual
assessment for Alaskan seabirds in
general, but I think it is particularly apropos
to this species.</p>

<p>We are still able only to guess at where the
marbled murrelet nests and we have not a clue
as to what sort of nesting strategy they pursue.
I am not prepared to accept, on the basis
of one North American record (Binford et al.
1975), that tree nesting is its habit throughout
its range. What has been proved is that
the marbled murrelet nests in trees and not,
as these authors would have us believe, that it
does not nest on the ground. It has become
rather fashionable to ignore the Chichagof Island
record (a ground nest), but it has not
been discredited. The color of the Chichagof
egg differs from that of the Big Basin egg, but
does agree with the one taken from an oviduct
by Cantwell (Gabrielson and Lincoln 1959).
My own experience leads me to believe that
tree nesting, if it occurs, is not the common
habit of marbled murrelets nesting in the
Prince William Sound region.</p>

<p>After many hours of observing marbled
murrelets over a period of several years, I am
intrigued by a number of things. These birds,
as often as not, appear to be clustered in
"pairs" as they feed. This occurs even at what
should be the height of the breeding season.
On several occasions I have noted a very pronounced
evening flight of these birds from
gathering areas on the water up into the surrounding
mountains at sunset. This has
moved me to wonder if their nesting strategy
includes incubating at night but less than full-time
attendance on days when the eggs can be
warmed by the sun. We know that periodic
egg-neglect is an aspect of storm-petrel behavior
(Pefaur 1974). Is this behavior also
possible on a more regular basis in an alcid? If
so, it would certainly help explain why nests
are hard to find.</p>

<p>It is apparent that more needs to be known
about the population dynamics and life history
of the marbled murrelet before we can
make a proper estimate of its abundance. In
spite of the fragmentary record, I conclude
that the marbled murrelet probably enjoys
the same relative abundance and distribution
that it did at the beginning of the century.</p>


<h6>Kittlitz's Murrelet <em>(Brachyramphus
brevirostris)</em></h6>

<p>The Kittlitz's murrelet is not as abundant
as the marbled murrelet, but locally it is sometimes
found in large numbers. FWS surveys
conducted during July-August 1972 provide
an estimate of 57,000 murrelets of this species
in Prince William Sound. Almost a fifth of
these were concentrated in Unakwik Inlet
above Unakwik Reef. Even more interesting,
about 2,500 of these birds were concentrated
in one loose flock.</p>

<p>In addition to Unakwik Inlet, Kittlitz's
murrelets concentrate in College Fjord in
Prince William Sound and in the waters fronting
the Bering-Malaspina ice-fields (Isleib and
Kessel 1973). Common as they are in these
waters, this species is supposed to be even
more abundant at Glacier Bay. The common
feature of these waters is the amount of ice
that can be found below their tributary
glaciers.</p>

<p>The Kittlitz's murrelet is apparently distributed
from LeConte Bay, east of Petersburg,
Alaska, north to Point Barrow and west
across the Aleutians to Attu, where Murie collected
a pair (Gabrielson and Lincoln 1959). I
once flushed a murrelet from an area of tread
and riser topography near the top of the
highest point on Kiska Island in heavy cloud
cover, and although I could not see this bird
well, I thought it to be of this species. From
the range description in Gabrielson and
Lincoln (1959) and Udvardy's (1963) range
map, it is apparent that the distribution of
this species is rather patchy, but I suspect
that for the more mountainous part of its
range this is more apparent than accurate.
The record is too fragmentary to allow an assessment
of any change in status during the
historical period.</p>


<h6>Ancient Murrelet <em>(Synthliboramphus
antiquus)</em></h6>

<p>Chase Littlejohn (Bendire 1895) spent the
spring and summer of 1894 collecting eggs on
islands south of the Alaska Peninsula. He has
left us a detailed record of what he saw but
not where he saw it. Bent (1963) stated flatly<span class="pagenum"><a name="Page_65" id="Page_65">[Pg 65]</a></span>
that the site of his collecting was Sanak Island
and this has common acceptance. Several
things in his account point to a site which was
a small island with several peers close by, but
this could not have been Sanak. It could have
been an island in the Sanak Island group or it
could equally well have been somewhere in the
Sandman Reefs. Unfortunately, because of
this the record is clouded. There has never
been anything approaching a survey of the
southern half of the Sandman Reefs. We do
not know what breeding colonies are there.</p>

<p>At any rate, Littlejohn told of the large
numbers of Leach's storm-petrels, fork-tailed
storm-petrels, auklets (of which only Cassin's
is specifically identified), and ancient murrelets
which occupied a large number of small islands.
He could not calculate the number of
breeding murrelets on his small island, the
size of which I interpret to have been of the
same order of magnitude as two others which
he estimated were about 2 acres. He does say
that the murrelets must have numbered
several thousand and could, if left alone by
the Aleuts, have quickly grown too numerous
for the island to accommodate.</p>

<p>Murie (1959) made a brief visit to Sanak in
1937 and learned that there were no longer
any large colonies of seabirds. He attributes
this to exploitation of the fisheries and to the
fox-farming industry. Littlejohn told of the
repeated visits of Aleuts to his small islands,
where they took hundreds of birds each time
and all of the eggs they could find. This kind
of activity could not help but disrupt the
breeding on these islands.</p>

<p>Littlejohn's description of the ancient
murrelet's nest leaves little doubt that the
birds could be reached by fox or rats with
ease. The birds showed no particular care in
selecting a nest site and often worked their
way back no more than about a meter into the
dead vegetative cover from preceding years,
where they scratched out a shallow nest.</p>

<p>There are few records of the ancient murrelet
from the northern and western Gulf of
Alaska. Friedmann (1935) reported the collection
of a series of eggs in 1884 on Kodiak Island.
Chase Littlejohn (Bendire 1895) collected
eggs from somewhere in the Sanak
Group in 1894. In 1908 Dixon (Grinnell 1910)
saw a bird in Port Nellie Juan. Several were
seen by Jaques (1930) near Belkofski in May
1928. Gabrielson collected one bird at Cordova
in September 1941 and another at the
Chiswell Islands in July 1945 (Gabrielson and
Lincoln 1959). He saw numerous flocks in the
Gulf of Alaska on 30 July of an unnamed year.
In 1943, he would have been near Cape
Spencer on that date. In 1945 he would have
been near the Chiswell Islands. In either case,
he was probably somewhere in Blying Sound.</p>

<p>The ancient murrelet is relatively uncommon
but regularly observed in the inshore
waters along the outer coasts of the islands
fronting Prince William Sound. FWS surveys
in July-August 1972 provided an estimate of
almost 1,000 birds, mostly in nonbreeding
plumage, along the outer coast of Prince
William Sound (Isleib and Kessel 1973). Small
numbers were found feeding close to the
Wooded Islands on 24 July (my personal observation).
Rausch (1958) saw a few off
Middleton Island in 1956. Isleib (Isleib and
Kessel 1973) saw 400-500 widely distributed
at the mouth of Yakutat Bay in July and August
1968. The only large numbers of ancient
murrelets encountered on the FWS survey of
the Alaskan Peninsula in 1973 were in the
Shumagin Islands. They were very common in
East Nagai Strait on 9 June and more than
half of the 1,300 seabirds per square nautical
mile encountered between Little Koniuji and
Chernabura Islands on 11 June were ancient
murrelets. At Nagai Island an estimated
5,000 ancient murrelets were observed in the
west bay at Pirate Shake, and later (on
19 June) several were observed in the vicinity
of Midun Island (FWS, Anchorage, Alaska,
unpublished data).</p>

<p>On the basis of the observations recounted
above, I have to conclude that ancient murrelets
are fairly regularly, if patchily, distributed
throughout the northern and western
Gulf of Alaska. I do not believe that the void
in their range shown for the northern Gulf of
Alaska by Udvardy (1963) is correct. Several
colonies are there, awaiting discovery.</p>

<p>Ancient murrelets are not abundant in the
Gulf of Alaska but they are certainly more
numerous than we have been able to prove. It
is not possible to tell from the existing data
whether they were once more abundant than
they are now. I suspect, on the basis of the
Sanak Island experience, that we can conclude
that this species has been reduced in
number by various of man's activities.</p>

<p><span class="pagenum"><a name="Page_66" id="Page_66">[Pg 66]</a></span></p>


<h6>Cassin's Auklet <em>(Ptychoramphus aleuticus)</em></h6>

<p>Cassin's auklet is a very uncommon bird in
the northern Gulf of Alaska. In the western
Gulf it is more common, particularly from the
Shumagins west.</p>

<p>This auklet apparently once bred in great
numbers on islands in or near the Sanak
Group where Chase Littlejohn (Bendire 1895)
found them to be twice as numerous as the ancient
murrelets. Murie (1959) did not find
them there.</p>

<p>Littlejohn began encountering Cassin's
auklets at sea some 290 km southeast of
Unga, Shumagin Islands. Murie (1959) encountered
them near the Shumagins in May
1937. During the FWS 1973 reconnaissance
survey of the Alaska Peninsula, these auklets
were not encountered (or at least not identified)
until we reached the vicinity of Unga
Strait where we saw a few in mixed flocks
with other murrelets and auklets. They were
most numerous in East Nagai Strait. We encountered
them only twice in a situation
which indicated they might be breeding&mdash;on
Hall and Herendeen islands on the north end
of Little Koniuji Island.</p>

<p>Murie (1959) considered Cassin's auklet to
be no longer common west of Kodiak. In
Gabrielson's many voyages through the
northern and western Gulf of Alaska he encountered
them only twice, once off Cape
Spencer and once in the Chiswell Islands.</p>

<p>Thoresen (1964) commented that throughout
the northern part of its range the Cassin's
auklet has become gradually less frequent. Although
there are no data to dispute this, I believe,
as do Isleib and Kessel (1973), that they
are more numerous than observations would
indicate, and I would apply this to the entire
area. There are certainly colonies remaining in
the Shumagin Islands, and quite probably
along the south coast of the Kenai Peninsula.
When it is possible to fully explore the Sandman
Reefs there is a good probability that
they will be found there.</p>

<p>We can only guess at the reasons for their
decline. Bendire (1895) and Murie (1959) have
described some contributing factors.</p>


<h6>Parakeet Auklet <em>(Cyclorrhyncus psittacula)</em></h6>

<p>Gabrielson and Lincoln (1959) described the
parakeet auklet as the least colonial of any of
the Alaskan auklets. They also considered the
Aleutian Islands to be its principal nesting
grounds. There are old records of breeding
parakeet auklets from Kodiak (Friedmann
1935) and Little Koniuji (Bean 1882). Grinnell
(1910) reported two that were seen on Green
Island, Prince William Sound, and several
more that were seen near Knight Island.</p>

<p>Murie (1959) did not see any parakeet auklets
near Kodiak and Afognak islands which
he considered to be the eastern part of their
range. He did not think they were abundant
anywhere along the Alaska Peninsula. He
found a few near Sutwik Island in May 1936
and then noted that they were fairly common
near the Shumagins in May 1937.</p>

<p>Gabrielson found this species to be quite
numerous on the north side of Chowiet Island
in the Semidi Islands in 1945 (Gabrielson and
Lincoln 1959). He also saw numerous individuals
in Marmot Strait and saw one in the
Chiswell Islands during the same year. David
Roseneau (Isleib and Kessel 1973) found hundreds
close to East Amatuli Island in the Barren
Islands in 1965.</p>

<p>During FWS colony surveys, parakeet auklets
have been found in close proximity to six
seabird colonies in Prince William Sound.
During the July-August 1972 surveys, they
were estimated to number about 3,000 in the
Sound. They have also been found closely associated
with Chisik Island (David Snarski,
personal communication), the Chiswell Islands,
Nord and Sud islands in the Barrens,
Sea Otter Island, and Central and Long islands
along the Alaska Peninsula. They were
most numerous in the Shumagin Islands,
where they were found near Castle Rock, Hall
(9,000), Herendeen (3,000), Atkins (more than
5,000), and Little Koniuju islands. They were
again encountered south and west of Cold Bay
at High, Fawn, Let, Amagat, Umga, and
Patton islands. Many of these islands are in
the north half of the Sandman Reefs, the only
portion where any attempt has been made to
survey seabird colonies.</p>

<p>The parakeet auklet may not be abundant
anywhere in the Gulf of Alaska but, based on
the numbers of places it has been seen in recent
years, its population appears to be well
dispersed and probably doing very well. This
auklet is most abundant from the Shumagin
Islands westward. It is almost certainly more<span class="pagenum"><a name="Page_67" id="Page_67">[Pg 67]</a></span>
numerous than has been thought. Its habits
are secretive enough so that it could easily
escape notice.</p>

<p>Because the parakeet auklet nests predominantly
under boulders, it probably was not
much affected by fox. Rats would certainly
have reduced its numbers wherever these
were introduced into its breeding habitat. We
have no data to tell us whether there may
have been population fluctuations in the past,
but there undoubtedly were at least minor
ones locally after rats were introduced.</p>


<h6>Crested Auklet <em>(Aethia pygmaea)</em></h6>

<p>Udvardy (1963) shows the breeding range of
the crested auklet as extending from southern
Kodiak Island westward. Within the northern
and western Gulf of Alaska, it is certainly
most abundant in the eastern Shumagin
Islands.</p>

<p>Isleib saw this auklet in Prince William
Sound 3 times during the winter of 1972-73.
These are the only records he was aware of for
that area (Isleib and Kessel 1973). David
Roseneau (Isleib and Kessel 1973) saw several
in Amatuli Cove, Barren Islands, in June
1965. I observed one in the vicinity of Cape
Spencer in August 1973.</p>

<p>Friedmann (1935) listed the crested auklet
as a breeding bird at Kodiak, but considered it
to be much more abundant as a wintering
bird. Townsend (1913) has provided us with a
vivid description of the myriads of crested
auklets he encountered at Yukon Harbor,
Little Koniuji Island. Gabrielson and Lincoln
(1959) noted large numbers of crested auklet
around Simeonof and Bird islands in the Shumagin
Islands in 1946 and stated that the
Yukon Harbor colony was still thriving.</p>

<p>Crested auklets were not encountered on
the 1973 FWS reconnaissance survey until we
reached the Shumagin Islands. They were
abundant only at the southeastern end of
Little Koniuji, where we encountered perhaps
10,000 in Yukon Harbor and more than 50,000
in a small cove directly south of Yukon Harbor
on the opposite side of the island. As
numerous as they were, they did not match
Townsend's myriads or even come close to his
assessment that they "were here more
numerous than the 'choochkies' at St.
George." St. George Island in the Pribilofs is
famous for its least auklets which, in the past,
have been estimated to number as high as 36
million (Peterson and Fisher 1955). The numbers
there today do not even approach this
level and we have no way of knowing how
abundant they were when Townsend visited
the Pribilofs, but I think it is safe to say that
they probably numbered in the millions. There
are probably more crested auklets than we observed
on Little Koniuji, but there is certainly
no longer anything approaching millions of
birds. Properly pronounced, Koniuji is the
Aleut name for the crested auklet, so we can
assume that the original inhabitants were impressed
by its numbers.</p>

<p>During the 1973 FWS survey we did not see
crested auklets at either Simeonof or Bird islands.
On the overgrazed and cattle-trampled
Simeonof it does not seem possible that any
could still exist.</p>

<p>I suspect that a cattleman's greed has been
the undoing of any crested auklets that may
have nested on Simeonof Island. This would
not account for the loss of any colonies that
may have been on Bird Island, but the decaying
fox-trapper's cabin on that island undoubtedly
tells the story. Churnabura, with
its feral cattle, presents much the same problem
as Simeonof. As for Little Koniuji, have
horned puffins been partly responsible for the
decrease in crested auklets? The puffin colony
at the south end of Little Koniuji must be
exactly where Townsend's millions of crested
auklets once nested.</p>


<h6>Least Auklet <em>(Aethia pusilla)</em></h6>

<p>No least auklets were encountered in FWS
surveys in the Gulf of Alaska in the early
1970's. Udvardy (1963) shows their breeding
range as starting well west in the Aleutians.
Gabrielson and Lincoln (1959) give the eastern
limit of their breeding range as the Shumagin
Islands. Bent (1963) listed their breeding
range as extending east to Kodiak Island,
and Friedmann (1935) knew of only a few
specimens taken in the winter from Kodiak.
Perhaps least auklets nested somewhere in
the western Gulf of Alaska, and they may
still, but at the moment we have no evidence
to prove that they do.</p>


<h6>Rhinoceros Auklet <em>(Cerorhinca monocerata)</em></h6>

<p>Udvardy (1963) would have us believe that
the rhinoceros auklet did not nest between<span class="pagenum"><a name="Page_68" id="Page_68">[Pg 68]</a></span>
southeastern Alaska and the southern Kurile
Islands. Bent (1963<em>b</em>), on the other hand, lists
their breeding range as extending from Washington
to Agattu. Clark (1910) noted this
species in small numbers at Atka and Agattu.
Because of the lack of proof, Udvardy probably
had no options. I believe that Bent was
probably closer to describing their original
range. I base this assumption on recent observations
and on the additional fragments of information
reported by Gabrielson and Lincoln
(1959). Murie (1959) failed to find this species
anywhere in the Aleutians, but his primary
reason for being there, the fox-farming industry,
may have had a lot to do with his not
being able to find any.</p>

<p>The FWS surveys in Prince William Sound
in July-August 1972 located small numbers of
rhinoceros auklets in breeding plumage at the
Wooded Islands and at Stoney Island and
Channel Island in Montague Strait. These
birds gave every impression of being local
breeders. David Roseneau (Isleib and Kessel
1973) encountered two at the Barren Islands
in June 1965. Isleib and Kessel (1973) list a
few other records from this area.</p>

<p>My own experience leads me to believe that
there is a large colony somewhere on Afognak
Island, probably on or near Tonki Cape. On 30
May 1973 I noted a lone bird north of Afognak
Island. Later, on 8 and 9 August, I saw
several in the same area. On 13 August in
Marmot Strait I observed a number of rhinoceros
auklets, either singly or in groups of
up to 12. Some of these had small fish in their
beaks. As they flushed, they all flew off
toward Tonki Cape. This observation was
made just at last light, and I believe that
there were many others about that could not
be seen in the dying light. We did not encounter
this species along the Alaska Peninsula
during the FWS survey in 1973 until we
reached the end. There I had one quick
glimpse of what I was certain was a rhinoceros
auklet at Amagat Island.</p>


<h6>Horned Puffin <em>(Fratercula corniculata)</em></h6>

<p>The horned puffin is one of the most abundant
breeding birds in the Gulf of Alaska.
There are only a few really large colonies but
these birds breed just about anywhere there is
a cliff (even a low one) with suitable fractures
and crevices. During the Alaska Peninsula
surveys in 1973, I estimated that the frequency
with which these birds were seen on
the water was about half that of the tufted
puffin. They have been recorded in so many
places that there is nothing to be gained by a
reiteration of the record in the literature.</p>

<p>The horned puffins reach their greatest density
in the Gulf of Alaska west of Kodiak Island.
Murie (1959) estimated that the colony
at Amagat Island, Morzhovi Bay, contained
15,000 birds, one of the largest he had seen. It
contained at least 50,000 in 1973. Even at
that, it was no match for the colony on Little
Koniuji Island with its minimum 140,000
horned puffins. Other colonies with large
horned puffin components were at High Island
(40,000), Castle Rock (20,000), Mitrofani
Island (35,000), and Sosbee Bay (15,000).</p>

<p>Earlier in this paper, I commented at length
on the great and often rapid fluctuations in
populations of tufted puffins. The same phenomenon
affects horned puffins. In 1975 there
were relatively small numbers of horned
puffins at Little Koniuji where they had
flourished 2 years earlier (James Bartonek,
personal communication). Because they are
apparently subject to erratically oscillating
populations, it is hard to tell how they have
fared over the years.</p>


<h6>Tufted Puffin <em>(Lunda cirrhata)</em></h6>

<p>The tufted puffin, as previously indicated,
is also a bird with widely fluctuating populations.
Until we develop an understanding of
their population dynamics and can understand
the underlying cause of these fluctuations
it will not be possible to assess trends in
their populations or understand the implications
of such trends.</p>

<p>Tufted puffins are abundant throughout the
Gulf of Alaska. Small colonies can be located
almost anywhere. Along the Alaska Peninsula
there are a number of colonies with an estimated
breeding population in 1973 of more
than 15,000 birds. These are: Ashiiak Island
(20,000), Central Island (90,000), the Brother
Islands (45,000), The Haystacks (19,000),
Castle Rock (85,000), Bird Island (none, but
may contain 500,000-1,000,000 at times),
Peninsula Islands (35,000), the Twins
(18,000), Amagat Island (40,000), and Umga
Island (22,000). These colonies correspond to
the area where colonies were listed for the<span class="pagenum"><a name="Page_69" id="Page_69">[Pg 69]</a></span>
horned puffin.</p>

<p>Tufted puffin populations respond readily
to some undetermined short-term perturbations.
This is clearly demonstrated by their
rapid population fluctuations. Because of
their numbers and because of the apparent
rapidity with which their numbers rebound, it
is not so apparent that they have been affected
by long-term perturbations, as so many
other seabirds apparently have.</p>

<p>There is much unused or underused nesting
habitat suitable for this species. In some
cases there are very strong clues pointing to
why this habitat is vacant. On many islands
along the Alaska Peninsula, which have very
good-looking tufted puffin nesting habitat
and no puffins, there are visible signs of the
presence of fox&mdash;either fox trails or abandoned
trappers' cabins. I also suspect that the
brown bear <em>(Ursa arctos)</em> is another possible
contributing factor to population declines of
burrow nesters along this coast. I have seen
brown bears swimming from island to island
on foraging expeditions. George J. Divoky
(personal communication) has found brown
bears visiting Ugaiushak Island, which is
13 km from shore. There are other islands between
Ugaiushak and the mainland but the
shortest route from shore would require one
swim of 7 km. The motivation must be strong.</p>

<p>Tufted puffins may shift from colony to
colony. This could be an explanation for apparent
local population fluctuation, but if so,
I am puzzled by the apparent tenacity with
which puffins cling to some sites. Their constant
occupancy of sites where the vegetative
mat is breakaway tundra (Amundsen 1972) or
is underlain by sand results in the destruction
of these sites. Tufted puffins often cling to
them in spite of the fact that they have been
reduced to "slums."</p>

<p>My conclusion is that in spite of their large
numbers it appears that tufted puffin populations
in the Gulf of Alaska probably have
been reduced to a level below that of their undisturbed
state.</p>


<h4>Conclusions</h4>

<p>Seabird numbers in the Gulf of Alaska are
not static. Generally, they are probably much
less abundant than they were when Bering
made his voyage of discovery. There are,
nonetheless, considerable numbers of seabirds
breeding along the coasts of these waters.
Some species show signs of recovery from
past insults by man. With enlightened management
there is still time to preserve the
vast natural heritage that they represent and,
in many cases, to improve their status.</p>

<p>In attempting to address a complicated
subject in short space and a relatively narrow
frame of reference, I have certainly erred a
number of times. I would like to see the
wealth of new data that will be derived from
current work applied to this concept. An understanding
of past population fluctuations
and the underlying perturbations that they
reflect is essential for managers faced with
the problem of making good decisions on
measures to mitigate the potential adverse
impact of development.</p>


<h4>References</h4>

<div class="hangindent">

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<p>Cahalane, V. H. 1944. Birds of the Katmai region,
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<p>Clark, A. H. 1910. The birds collected and observed
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<p>Cody, M. L. 1973. Coexistence, coevolution and convergent
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<p>Coues, E. 1874. Birds of the Northwest. U.S. Geological
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<p>Dall, W. H. 1873. Notes on the avifauna of the Aleutian
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<p>Dall, W. H. 1874. Notes on the avifauna of the Aleutian
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<p>Dall, W. H., and H. M. Bannister. 1869. List of the
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<p>Dement'ev, G. P., and N. A. Gladkov, eds. 1966.
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553 pp.</p>

<p>Dixon, J. S. 1908. Field notes from Alaska. Condor
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<p>East, B. 1943. Seabird cities of the Aleutians. Nat.
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<p>Friedmann, H. 1934. Bird bones from Eskimo ruins
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<p>Friedmann, H. 1935. The birds of Kodiak Island,
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<p>Friedmann, H. 1937. Bird bones from archeological
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<p>Gabrielson, I. N., and F. C. Lincoln. 1959. The birds
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<p>Hanna, G. D. 1916. Records of birds new to the
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<p>Howell, J. C. 1948. Observations on certain birds of
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<p>Hrdlicka, A. 1945. The Aleutian and Commander
Islands and their inhabitants. Wistar Institute
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<p>Isleib, M. E., and B. Kessel. 1973. Birds of the
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<p>Jochelson, W. 1968. History, ethnology and anthropology
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<p>Johnson, R. A. 1938. Predation of gulls in murre
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<p>Johnson, R. A. 1941. Nesting behavior of the Atlantic
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<p>Kozlova, E. V. 1961. Charadriiformes, Suborder
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<p>Patten, S. M., Jr. 1976. Sympatry and interbreeding
of herring and glaucous-winged gulls in southern
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<p>Pefaur, J. E. 1974. Egg-neglect in the Wilson's
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<p>Rausch, R. L. 1958. The occurrence and distribution
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<p><span class="pagenum"><a name="Page_71" id="Page_71">[Pg 71]</a></span></p>

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<p>Thoresen, A. C. 1964. The breeding behavior of the
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<p>Townsend, C. H. 1913. The crested auklet. Bird-Lore
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<p>Willet, G. 1918. Bird notes from Forrester Island,
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<p>Williamson, F. S. L., and L. S. Peyton. 1963. Interbreeding
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<p><span class="pagenum"><a name="Page_72" id="Page_72">[Pg 72]</a><br /><a name="Page_73" id="Page_73">[Pg 73]</a></span></p>
<div class="chapter"></div>
<hr class="chap" />




<h3><a name="Status_and_Distribution_of_Breeding_Seabirds_of" id="Status_and_Distribution_of_Breeding_Seabirds_of">Status and Distribution of Breeding Seabirds of
Southeastern Alaska, British Columbia, and Washington</a></h3>


<p class="center">by</p>

<p class="center">David A. Manuwal</p>

<p class="center">College of Forest Resources<br />
University of Washington<br />
Seattle, Washington 98195</p>

<p class="center">and</p>

<p class="center">R. Wayne Campbell</p>

<p class="center">Provincial Museum<br />
Victoria, British Columbia, Canada
</p>


<h4>Abstract</h4>

<div class="blockquot">

<p>Current breeding seabird population estimates, nest-site preferences, and
population changes are reviewed for southeastern Alaska, British Columbia, and
Washington. There are 19 species of seabirds and a minimum of 216,566 pairs
breeding in British Columbia and Washington. There are limited data on breeding
populations for southeastern Alaska. Species diversity ranges from 17
species in Alaska to 15 species in British Columbia and 14 species in Washington.
Eighty percent of all British Columbia seabirds breed on the east coast of
Queen Charlotte Islands and the northwest coast of Vancouver Island. The three
most numerous species in British Columbia are the fork-tailed storm-petrel,
<em>Oceanodroma furcata</em> (31.3%); Cassin's auklet, <em>Ptychoramphus aleuticus</em> (24.6%);
and ancient murrelet, <em>Synthliboramphus antiquus</em> (12.5%). In Washington, 74%
(43,274 pairs) of the seabird population resides on the Olympic coast; the remaining
26% are in the San Juan Island area. About 54% of this population consists
of the common murre <em>(Uria aalge)</em> and rhinoceros auklet <em>(Cerorhinca monocerata)</em>.
The rhinoceros auklet and glaucous-winged gull <em>(Larus glaucescens)</em> make up 97%
of the total seabird population of the San Juan Islands. About 68% of all seabirds
on the northeastern Pacific coast are nocturnal, burrow or rock crevice-nesting
species. Currently available population data are inadequate to determine
significant changes in population density for most species. Suggested topics for
future research are presented.</p></div>

<p>The purpose of this paper is to discuss the
known distribution, habitat, abundance, and
status of breeding seabirds of the Alexander
Archipelago in southeastern Alaska, the
Province of British Columbia, Canada, and
the State of Washington.</p>

<p>Even though several studies of the breeding
biology of several seabird species in this area
have been published, there have been few published
surveys of known breeding colonies. In
British Columbia the most extensive work
has been done by the British Columbia Provincial
Museum and the University of British
Columbia (Drent and Guiguet 1961). Gabrielson
and Lincoln (1959) summarized the available
literature on Alaskan birds up to about
1958. Since then, no extensive surveys have
been conducted in southeastern Alaska. The
U.S. Department of the Interior (1972), in its
environmental impact statement for the
Trans-Alaska Pipeline, presented additional
information on the seabirds of other parts of
Alaska. In Washington, there are no published
comprehensive surveys except those of
Kenyon and Scheffer (1961) and unpublished
surveys by the U.S. Fish and Wildlife Service
and the University of Washington.</p>

<p><span class="pagenum"><a name="Page_74" id="Page_74">[Pg 74]</a></span></p>

<div class="center">
<table border="0" cellpadding="4" cellspacing="0" summary="Table 1. Taxonomic distribution of marine birds breeding along the Pacific Coast of Washington, British Columbia, and southeastern Alaska.">
    <caption>Table 1. <em>Taxonomic distribution of marine birds breeding along the Pacific Coast of Washington, British Columbia, and southeastern Alaska.</em></caption>
  <tr>
    <th rowspan="2">Family</th>
    <th rowspan="2">Common name</th>
    <th colspan="0">Regions</th>
  </tr>
  <tr>
    <th>British Columbia</th>
    <th>Washington</th>
    <th>Southeastern Alaska</th>
    <th>Total forms</th>
  </tr>
  <tr>
    <td>Hydrobatidae</td>
    <td>Storm-petrels</td>
    <td class="tdn">2</td>
    <td class="tdn">2</td>
    <td class="tdn">2</td>
    <td class="tdn">2</td>
  </tr>
  <tr>
    <td>Phalacrocoracidae</td>
    <td>Cormorants</td>
    <td class="tdn">3</td>
    <td class="tdn">3</td>
    <td class="tdn">1</td>
    <td class="tdn">3</td>
  </tr>
  <tr>
    <td>Haematopodidae</td>
    <td>Oystercatchers</td>
    <td class="tdn">1</td>
    <td class="tdn">1</td>
    <td class="tdn">1</td>
    <td class="tdn">1</td>
  </tr>
  <tr>
    <td>Laridae</td>
    <td>Gulls and terns</td>
    <td class="tdn">2</td>
    <td class="tdn">2</td>
    <td class="tdn">3</td>
    <td class="tdn">4</td>
  </tr>
  <tr>
    <td>Alcidae</td>
    <td>Auks, murres, puffins</td>
    <td class="tdn">6</td>
    <td class="tdn">7</td>
    <td class="tdn">9</td>
    <td class="tdn">9</td>
  </tr>
  <tr>
    <td colspan="2" class="tdp">Total</td>
    <td class="tdn">14</td>
    <td class="tdn">15</td>
    <td class="tdn">16</td>
    <td class="tdn">19</td>
  </tr>
</table></div>


<h4>Taxonomic Distribution
of Marine Birds</h4>

<p>There are 19 species of seabirds that breed
along the Pacific coast of southeastern
Alaska, British Columbia, and Washington
(Table 1). Southeastern Alaska has the
largest number (17) of species. Errors in
species identification are most likely with the
Larus gulls, particularly in southeastern
Alaska where the herring gull <em>(L. argentatus)</em>
and glaucous-winged gull <em>(L. glaucescens)</em>
breed in mixed colonies (Patten and Weisbrod
1974). A similar situation exists in Washington
where the western <em>(L. occidentalis)</em> and
glaucous-winged gulls intergrade (Scott
1971). Brandt's <em>(Phalacrocorax penicillatus)</em>
and double-crested cormorants <em>(P. auritus)</em>
are often difficult to identify from the air.
This would be a problem in Washington and
the southwest coast of Vancouver Island,
where the two species are locally sympatric.</p>


<h4>Southeastern Alaska</h4>

<p>The area under consideration is the 400-km-long
Alexander Archipelago (Fig. 1). This
complex pattern of islands, bays, and inlets is
characterized by extremely high precipitation
and typical cool marine temperatures. Average
annual precipitation in the Sitka area is
245.4 cm (1931-60), and the average annual
temperature is 6.3&deg;C (U.S. Weather Bureau
1974). As a consequence of this cool, humid
environment, most of the islands are densely
covered with conifers, chiefly Sitka spruce
<em>(Picea sitkensis)</em> and hemlock <em>(Tsuga heterophylla)</em>,
and an almost impenetrable shrub
cover composed of salmonberry <em>(Rubus
spectabilis)</em>, elderberry <em>(Sambucus callicaipa)</em>,
devil's club <em>(Echinopanax horridus)</em>, and three
species of <em>Vaccinium</em> (Heath 1915).</p>

<div class="figcenter" style="width: 700px;">
<img src="images/p_074.jpg" width="700" height="552" alt="" />
<div class="caption"><p>Fig. 1. Map of southeastern Alaska showing major
seabird breeding colonies: 1&mdash;North Marble
Island; 2&mdash;Forrester Island.</p></div>
</div>

<p>There are 16 species of marine birds breeding
in the Alexander Archipelago. The major
seabird breeding colonies are located at Glacier
Bay and at St. Lazaria, Hazy, and Forrester
islands (Fig. 1; Table 2). Published surveys
of these colonies are available only for
St. Lazaria (Willett 1912) and Forrester islands
(Heath 1915; Willett 1915). Several
authors have reported on seabirds from surrounding
areas (Grinnell 1897, 1898, 1909;
Swarth 1911, 1922, 1936; Patten 1974). There
have been no surveys of seabirds of southeastern
Alaska since before the 1940's (J. G. King,
Jr., personal communication). However, since
census data are available for only two colonies,
we discuss them in more detail.</p>

<p><span class="pagenum"><a name="Page_75" id="Page_75">[Pg 75]</a></span></p>

<div class="center">
<table border="0" cellpadding="4" cellspacing="0" summary="Table 2. Population estimates of seabirds breeding on St. Lazaria and Forrester islands, southeastern Alaska">
    <caption>Table 2. <em>Population estimates of seabirds breeding on St. Lazaria and Forrester islands, southeastern Alaska</em> (data from Willett 1912 and 1915).</caption>
  <tr>
    <th rowspan="2">Bird species</th>
    <th colspan="2" class="br">St. Lazaria Island</th>
    <th colspan="0">Forrester Island</th>
  </tr>
  <tr>
    <th>Number of pairs</th>
    <th>Percent of total</th>
    <th>Number of pairs</th>
    <th>Percent of total</th>
  </tr>
  <tr>
    <td>Fork-tailed storm-petrel</td>
    <td class="tdn">2,000</td>
    <td class="tdn">8.0</td>
    <td class="tdn">10,000</td>
    <td class="tdn">6.0</td>
  </tr>
  <tr>
    <td>Leach's storm-petrel</td>
    <td class="tdn">20,000</td>
    <td class="tdn">80.0</td>
    <td class="tdn">50,000</td>
    <td class="tdn">30.0</td>
  </tr>
  <tr>
    <td>Pelagic cormorant</td>
    <td class="tdn">150</td>
    <td class="tdn">0.6</td>
    <td class="tdn">150</td>
    <td class="tdn">0.0</td>
  </tr>
  <tr>
    <td>Black oystercatcher</td>
    <td class="tdn">4</td>
    <td class="tdn">0.0</td>
    <td class="tdn">50</td>
    <td class="tdn">0.0</td>
  </tr>
  <tr>
    <td>Glaucous-winged gull</td>
    <td class="tdn">300</td>
    <td class="tdn">1.2</td>
    <td class="tdn">8,000</td>
    <td class="tdn">4.8</td>
  </tr>
  <tr>
    <td>Herring gull</td>
    <td></td>
    <td></td>
    <td class="tdn">220</td>
    <td class="tdn">0.0</td>
  </tr>
  <tr>
    <td>Common murre</td>
    <td class="tdn">300</td>
    <td class="tdn">1.2</td>
    <td class="tdn">20,000</td>
    <td class="tdn">12.0</td>
  </tr>
  <tr>
    <td>Pigeon guillemot</td>
    <td class="tdn">150</td>
    <td class="tdn">0.6</td>
    <td class="tdn">300</td>
    <td class="tdn">0.0</td>
  </tr>
  <tr>
    <td>Ancient murrelet</td>
    <td></td>
    <td></td>
    <td class="tdn">20,000</td>
    <td class="tdn">12.0</td>
  </tr>
  <tr>
    <td>Cassin's auklet</td>
    <td></td>
    <td></td>
    <td class="tdn">2,000</td>
    <td class="tdn">1.2</td>
  </tr>
  <tr>
    <td>Rhinoceros auklet</td>
    <td class="tdn">75</td>
    <td class="tdn">0.0</td>
    <td class="tdn">20,000</td>
    <td class="tdn">12.0</td>
  </tr>
  <tr>
    <td>Horned puffin</td>
    <td class="tdn">12</td>
    <td class="tdn">0.0</td>
    <td class="tdn">1,100</td>
    <td class="tdn">0.7</td>
  </tr>
  <tr>
    <td>Tufted puffin</td>
    <td class="tdn">2,000</td>
    <td class="tdn">8.0</td>
    <td class="tdn">35,000</td>
    <td class="tdn">21.0</td>
  </tr>
  <tr>
    <td class="tdp">Total</td>
    <td class="tdn">24,991</td>
    <td></td>
    <td class="tdn">166,820</td>
    <td></td>
  </tr>
</table></div>

<p>The studies by Willett (1912, 1915) and
Heath (1915) provide some base-line information
on species composition and abundance
with which future studies on St. Lazaria and
Forrester islands can be compared (Table 2).
The somewhat greater species diversity on
Forrester Island is primarily due to its
greater size and more suitable soil type for ancient
murrelets <em>(Synthliboramphus antiquus)</em>
and Cassin's auklets <em>(Ptychoramphus aleutica)</em>,
species that are absent on St. Lazaria.
Storm-petrels (<em>Oceanodroma</em> spp.) are the
most numerous species on both islands, but
there are proportionately more storm-petrels
(88%) on St. Lazaria than on Forrester (36%).
On the other hand, there are many large, burrowing
alcids on Forrester Island. Nearly a
third of the birds on Forrester are rhinoceros
auklets <em>(Cerorhinca monocerata)</em>, tufted
puffins <em>(Lunda cirrhata)</em>, and horned puffins
<em>(Fratercula corniculata)</em>.</p>

<p>The species composition of seabirds breeding
on other islands is similar to that found on
Forrester and St. Lazaria islands but less
abundant. In Glacier Bay, for example, the
only population data available are those provided
by Patten (1974) for North Marble Island:
pelagic cormorants, <em>Phalacrocorax
pelagicus</em> (30 pairs); black oystercatchers,
<em>Haematopus bachmani</em> (8); herring gulls (7);
glaucous-winged gulls (500); common murres,
<em>Uria aalge</em> (18); pigeon guillemots, <em>Cepphus
columba</em> (60); horned puffins (4); and tufted
puffins (30).</p>

<p>At the present time, it is impossible to draw
any conclusions about changes in population
density and distribution for most of the seabirds
breeding in southeastern Alaska. Adequate
data are available only for St. Lazaria
and Forrester islands where Willet and Heath
provided the only early extensive census data
for this part of Alaska.</p>


<h4>British Columbia</h4>

<p>The rugged British Columbia coastline is
characterized by 930 km of islands and inlets
(Figs. 2, 3). With the exception of the inner
southern portion, this coast is mostly uninhabited.
The physical characteristics of the
offshore islands are similar to those found off
the Washington coast. Descriptions of some
of these islands and the 15 species of breeding
seabirds on them have been given by Drent
and Guiguet (1961), Guiguet (1971), and
Summers (1974).</p>

<p>A detailed analysis of British Columbia seabirds
is not presented here since a more
thorough analysis is in preparation by R. W.
Campbell and R. H. Drent (manuscript). Instead,
we present seabird population estimates
available for the Province up to the
summer of 1975; Tables 3 and 4 summarize
these estimates for the five major portions of
coastal British Columbia. The coast of British
Columbia contains a myriad of small islands
where there may be small numbers of breed<span class="pagenum"><a name="Page_76" id="Page_76">[Pg 76]</a></span>ing
seabirds. Many of these have not been censused
and are too numerous to include in
Tables 3 and 4.</p>

<div class="figcenter" style="width: 700px;">
<img src="images/p_076a.jpg" width="700" height="529" alt="" />
<div class="caption"><p>Fig. 2. Map of northern British Columbia showing
sites of major seabird breeding colonies: 1&mdash;Skedans
Island; 2&mdash;Limestone Island; 3&mdash;Agglomerate
Island; 4&mdash;Bischoff Island; 5&mdash;Ramsey
Island; 6&mdash;Alder Island; 7&mdash;Rankins Island.</p></div>
</div>

<div class="figcenter" style="width: 700px;">
<img src="images/p_076b.jpg" width="700" height="563" alt="" />
<div class="caption"><p>Fig. 3. Map of southern British Columbia showing
sites of major seabird breeding colonies: 1&mdash;Triangle
Island; 2&mdash;Cleland Island.</p></div>
</div>

<p>More than half of the breeding seabirds in
British Columbia are found on the east coast
of the Queen Charlotte Islands, and the fork-tailed
storm-petrel <em>(Oceanodroma furcata)</em>
comprises more than half of that total. However,
new unpublished data (K. Vermeer) for
Triangle Island and the northwest coast of
Vancouver Island indicate that the population
figures in Table 3 for this area are underestimates.
Nevertheless, these two regions
have nearly 80% of all the breeding seabirds
in the Province. This results from the very
large populations of the rhinoceros auklet and
tufted puffin on Triangle Island and the fork-tailed
storm-petrel, ancient murrelet, and Cassin's
auklet on various islands on the east
coast of the Queen Charlotte Islands
(Table 3).</p>

<p>Continuing surveys of breeding seabirds are
being conducted by personnel of the British
Columbia Provincial Museum and the Canadian
Wildlife Service.</p>


<h4>Washington State</h4>


<h5><em>General Environment</em></h5>

<p>For this report, we have distinguished two
major geographical areas in Washington
where breeding seabirds are found&mdash;the western
coast of the Olympic Peninsula and the
San Juan Islands, including the Strait of Juan
de Fuca.</p>

<p>On the Olympic Peninsula, seabirds breed
on the offshore rocks, islands, and precipitous
cliffs from Copalis Beach to Cape Flattery
(Fig. 4). The offshore rocks and islands
throughout this area (except Tatoosh Island)
are now included in the Washington Islands
National Wildlife Refuge. Most of the larger
rocks and islands have dense stands of salmonberry,
salal, and grasses, and a few support
stands of stunted conifers (Fig. 5); most
are inaccessible to man. The adjacent coast is
dominated by the Olympic rain forest where
the mean annual precipitation is about
337.1 cm (U.S. Weather Bureau 1956, 1965<em>a</em>,
1965<em>b</em>).</p>

<p>Because the San Juan Islands lie northeast
of the Olympic Peninsula and east of Vancouver
Island (Fig. 6) they are in a rain
shadow; however, because of highly variable
topography and aspect, most islands have a
diverse assemblage of plant communities
(Franklin and Dyrness 1973). Exposed south-facing
slopes are occupied by grassland vegetation
and frequently by scattered trees,
usually <em>Pseudotsuga menziesii</em> and <em>Arbutus
menziesii</em>. Most of the seabird colonies are located
on rather small exposed islands with<span class="pagenum"><a name="Page_77" id="Page_77">[Pg 77]</a></span>
short, grassy, shrubby vegetation. In general,
these islands are not suitable for burrowing
species.</p>

<div class="center">
<table border="0" cellpadding="4" cellspacing="0" summary="Table 3. Species composition, population estimates, and distribution of seabirds in British Columbia.">
    <caption>Table 3. <em>Species composition, population estimates, and distribution of seabirds in British Columbia.</em><a name="FNanchor_5_5" id="FNanchor_5_5"></a><a href="#Footnote_5_5" class="fnanchor">[5]</a><a name="FNanchor_6_6" id="FNanchor_6_6"></a><a href="#Footnote_6_6" class="fnanchor">[6]</a> (+ = Present.)</caption>
  <tr>
    <th>Bird species</th>
    <th>Straits of Georgia and Juan de Fuca</th>
    <th>Southwest coast of Vancouver Island</th>
    <th>Northwest coast of Vancouver Island</th>
    <th>West coast of Queen Charlotte Island</th>
    <th>East coast of Queen Charlotte Island</th>
    <th>Prince Rupert to Queen Charlotte Island</th>
    <th>Total birds</th>
    <th>Percent of total</th>
  </tr>
  <tr>
    <td>Fork-tailed storm-petrel</td>
    <td></td>
    <td class="tdc">+</td>
    <td class="tdc">+</td>
    <td class="tdn">1,050</td>
    <td class="tdn">97,100</td>
    <td class="tdc">+</td>
    <td class="tdn">98,160</td>
    <td class="tdn">31.3</td>
  </tr>
  <tr>
    <td>Leach's storm-petrel</td>
    <td></td>
    <td class="tdn">10,000</td>
    <td class="tdc">+</td>
    <td class="tdn">1,800</td>
    <td class="tdn">180</td>
    <td class="tdn">750</td>
    <td class="tdn">12,730</td>
    <td class="tdn">4.1</td>
  </tr>
  <tr>
    <td>Double-crested cormorant</td>
    <td class="tdn">1,058</td>
    <td class="tdn">0</td>
    <td class="tdn">0</td>
    <td class="tdn">0</td>
    <td class="tdn">0</td>
    <td class="tdn">0</td>
    <td class="tdn">2,116</td>
    <td class="tdn">&gt;1.0</td>
  </tr>
  <tr>
    <td>Brandt's cormorant</td>
    <td></td>
    <td class="tdn">370</td>
    <td class="tdn">0</td>
    <td class="tdn">0</td>
    <td class="tdn">0</td>
    <td class="tdn">0</td>
    <td class="tdn">370</td>
    <td class="tdn">&gt;1.0</td>
  </tr>
  <tr>
    <td>Pelagic cormorant</td>
    <td class="tdn">2,174</td>
    <td class="tdn">336</td>
    <td class="tdn">3,350</td>
    <td class="tdn">1,456</td>
    <td class="tdn">496</td>
    <td class="tdn">12</td>
    <td class="tdn">9,998</td>
    <td class="tdn">3.2</td>
  </tr>
  <tr>
    <td>Glaucous-winged gull</td>
    <td class="tdn">10,123</td>
    <td class="tdn">6,870</td>
    <td class="tdn">600</td>
    <td class="tdn">412</td>
    <td class="tdn">866</td>
    <td class="tdn">540</td>
    <td class="tdn">29,534</td>
    <td class="tdn">9.4</td>
  </tr>
  <tr>
    <td>Common murre</td>
    <td></td>
    <td class="tdn">16</td>
    <td class="tdn">3,000</td>
    <td class="tdn">0</td>
    <td class="tdn">0</td>
    <td class="tdn">0</td>
    <td class="tdn">3,016</td>
    <td class="tdn">1.0</td>
  </tr>
  <tr>
    <td>Pigeon guillemot</td>
    <td class="tdn">1,029</td>
    <td class="tdn">204</td>
    <td class="tdn">250</td>
    <td class="tdn">358</td>
    <td class="tdn">1,458</td>
    <td class="tdn">1,650</td>
    <td class="tdn">5,978</td>
    <td class="tdn">1.9</td>
  </tr>
  <tr>
    <td>Ancient murrelet</td>
    <td></td>
    <td class="tdn">0</td>
    <td class="tdn">0</td>
    <td class="tdn">200</td>
    <td class="tdn">42,150</td>
    <td class="tdn">4</td>
    <td class="tdn">42,354</td>
    <td class="tdn">13.5</td>
  </tr>
  <tr>
    <td>Cassin's auklet</td>
    <td></td>
    <td class="tdc">+</td>
    <td class="tdn">50,000</td>
    <td class="tdc">+</td>
    <td class="tdn">26,500</td>
    <td class="tdn">450</td>
    <td class="tdn">76,950</td>
    <td class="tdn">4.6</td>
  </tr>
  <tr>
    <td>Rhinoceros auklet</td>
    <td></td>
    <td class="tdn">1,200</td>
    <td class="tdc">+</td>
    <td class="tdn">10,000</td>
    <td class="tdn">300</td>
    <td class="tdn">200</td>
    <td class="tdn">11,700</td>
    <td class="tdn">3.7</td>
  </tr>
  <tr>
    <td>Tufted puffin</td>
    <td class="tdn">1</td>
    <td class="tdn">154</td>
    <td class="tdn">20,000</td>
    <td class="tdn">190</td>
    <td class="tdn">0</td>
    <td class="tdn">42</td>
    <td class="tdn">20,388</td>
    <td class="tdn">6.5</td>
  </tr>
  <tr>
    <td class="tdp">Total</td>
    <td class="tdn">14,385</td>
    <td class="tdn">19,150</td>
    <td class="tdn">77,200</td>
    <td class="tdn">15,466</td>
    <td class="tdn">169,050</td>
    <td class="tdn">3,648</td>
    <td class="tdn">313,294</td>
    <td class="tdn">81.2</td>
  </tr>
</table></div>


<div class="pagenum"><a name="Page_78" id="Page_78">[Pg 78]</a></div>

<div class="center">
<table border="0" cellpadding="4" cellspacing="0" summary="Table 4. Breeding seabird population estimates for British Columbia">
    <caption>Table 4. <em>Breeding seabird population estimates for British Columbia</em><a name="FNanchor_7_7" id="FNanchor_7_7"></a><a href="#Footnote_7_7" class="fnanchor">[7]</a><a name="FNanchor_8_8" id="FNanchor_8_8"></a><a href="#Footnote_8_8" class="fnanchor">[8]</a></caption>
  <tr>
    <th>Geographic location</th>
    <th>Population estimate</th>
    <th>Percent of total</th>
  </tr>
  <tr>
    <td>Straits of Georgia and Juan de Fuca</td>
    <td class="tdr">14,385</td>
    <td class="tdr">9.2</td>
  </tr>
  <tr>
    <td>Southwest Coast of Vancouver Island</td>
    <td class="tdr">9,575</td>
    <td class="tdr">6.1</td>
  </tr>
  <tr>
    <td>Northwest Coast of Vancouver Island</td>
    <td class="tdr">38,600</td>
    <td class="tdr">24.6</td>
  </tr>
  <tr>
    <td>West Coast of Queen Charlotte Island</td>
    <td class="tdr">7,733</td>
    <td class="tdr">4.9</td>
  </tr>
  <tr>
    <td>East Coast of Queen Charlotte Island</td>
    <td class="tdr">84,530</td>
    <td class="tdr">54.0</td>
  </tr>
  <tr>
    <td>Prince Rupert to Queen Charlotte Strait</td>
    <td class="tdr">1,824</td>
    <td class="tdr">1.2</td>
  </tr>
  <tr>
    <td class="tdp">Total</td>
    <td class="tdr">156,647</td>
    <td class="tdr">100.0</td>
  </tr>
</table></div>

<div class="figcenter" style="width: 490px;">
<img src="images/p_078.jpg" width="490" height="700" alt="" />
<div class="caption"><p>Fig. 4. Map of the Olympic Peninsula of
Washington State showing sites of major seabird
breeding colonies: 1&mdash;Protection Island; 2&mdash;Carroll
Island; 3&mdash;Destruction Island.</p></div>
</div>

<p>In the Strait of Juan de Fuca, the two most
important sites are Smith and Protection islands.
Both are composed of glacial deposits
and heavy sod that has developed under dense
grassy vegetation (Fig. 7). Consequently,
these two islands support most of the burrowing
seabirds in the region. Unfortunately,
both islands have historically been subjected
to much human disturbance (Richardson
1961; Manuwal 1974).</p>

<p>The existing information on seabird colonies
in both the coastal and San Juan Island
areas has been largely derived from aerial surveys
by the U.S. Fish and Wildlife Service.
These surveys are inherently biased toward
surface-nesting species such as gulls and cormorants.
Population estimates for guillemots,
auklets, storm-petrels, and puffins are less
accurate. Some additional information obtained
by direct island visitation has been provided
by Kenyon and Scheffer (1961),
Richardson (1961), Thoresen and Galusha
(1971), G. Eddy (unpublished data), and D. A.
Manuwal (unpublished data). Although other
accounts of Washington seabirds are available,
the references listed above are specifically
oriented toward population assessment.</p>


<h5><em>Olympic Peninsula</em></h5>

<p>Despite the large number of offshore rocks,
islets, and islands along the Pacific coast of
Washington, significant seabird colonies are
present only on about 30 islands. Since
Table 5 summarizes the population estimates
for 12 species of seabirds breeding on 24
major sites, it represents only the majority
and not the total number of breeding seabirds
on the Pacific coast of Washington. About
74% of the entire Washington seabird population
resides on the coastal rocks and islands.</p>

<p>Major colony sites with more than 2,500
breeding pairs are Grenville Arch,<span class="pagenum"><a name="Page_79" id="Page_79">[Pg 79]</a></span>
Willoughby Rock, Destruction Island, Cake
Rock, Carroll Island, and Bodelteh Island.
More intensive censusing, especially of nocturnal
burrowing species will undoubtedly
raise the population estimates for these and
other islands off the coast. About 54% of the
total coastal population is composed of the
common murre and rhinoceros auklet.</p>

<div class="figcenter" style="width: 700px;">
<img src="images/p_079a.jpg" width="700" height="451" alt="" />
<div class="caption"><p>Fig. 5. Photograph of Destruction Island off the coast of Washington.</p></div>
</div>

<div class="figcenter" style="width: 700px;">
<img src="images/p_079b.jpg" width="700" height="595" alt="" />
<div class="caption"><p>Fig. 6. Map of the San Juan Archipelago showing
sites of major seabird breeding colonies: 1&mdash;Viti
Rocks; 2&mdash;Colville Island; 3&mdash;Smith Island.</p></div>
</div>


<h5><em>San Juan Islands</em></h5>

<p>There are about 86 actual or potential seabird
colony sites in this area; 25 (30%) are now
considered important. Eleven islands are
under Federal protection as National Wildlife
Refuges. Part of Protection Island is owned
by the Washington State Game Department
to protect the largest rhinoceros auklet colony
in the State. Most colony sites are on small islands
with poorly developed soil which prevents
burrowing species from using them.
Consequently, the dominant species are surface
nesters (such as gulls and cormorants)
and rock-crevice nesters (like the pigeon
guillemot). In all, about 31,000 seabirds of 7
species breed in the San Juan Island area.
Breeding seabird population estimates for 49
of the 86 nesting sites are given in Table 6.
Even though this does not represent all the
colonies, it covers the most important islands
and those islands where there appears to be
potential for seabird breeding.</p>

<p><span class="pagenum"><a name="Page_80" id="Page_80">[Pg 80]</a></span></p>

<div class="figcenter" style="width: 500px;">
<img src="images/p_080.jpg" width="500" height="700" alt="" />
<div class="caption"><p>Fig. 7. Photograph of Smith Island in the Strait of Juan de Fuca, Washington. The glacial deposits are evident
from the composition of the cliff faces.</p></div>
</div>

<p><span class="pagenum"><a name="Page_81" id="Page_81">[Pg 81]</a></span></p>

<div class="center">
<table border="0" cellpadding="4" cellspacing="0" summary="Table 5. Estimated breeding seabird populations of the outer coast of Washington">
    <caption>Table 5. <em>Estimated breeding seabird populations of the outer coast of Washington</em>.<a name="FNanchor_9_9" id="FNanchor_9_9"></a><a href="#Footnote_9_9" class="fnanchor">[9]</a> (Unpublished data from U.S. Fish and Wildlife Service and University of Washington)</caption>
  <tr>
    <th rowspan="3">Breed&shy;ing site</th>
    <th colspan="0">Species</th>
  </tr>
  <tr>
    <th colspan="2" class="br">Storm-pet&shy;rels</th>
    <th colspan="4">Cor&shy;mo&shy;rants</th>
    <th rowspan="2">Black oys&shy;ter-catch&shy;er</th>
    <th colspan="2">Gulls</th>
    <th rowspan="2">Com&shy;mon mur&shy;re</th>
    <th rowspan="2">Pi&shy;geon guil&shy;le&shy;mot</th>
    <th colspan="2">Auk&shy;lets</th>
    <th rowspan="2">Tuft&shy;ed puf&shy;fin</th>
    <th rowspan="2">To&shy;tal</th>
  </tr>
  <tr>
    <th>Fork-tail&shy;ed</th>
    <th>Le&shy;ach's</th>
    <th>Un&shy;i&shy;den&shy;ti&shy;fied</th>
    <th>Dou&shy;ble-crest&shy;ed</th>
    <th>Bran&shy;dt's</th>
    <th>Pe&shy;lag&shy;ic</th>
    <th>West&shy;ern</th>
    <th>Glau&shy;cous-wing&shy;ed west&shy;ern</th>
    <th>Cass&shy;in's</th>
    <th>Rhi&shy;noc&shy;er&shy;os</th>
  </tr>
  <tr>
    <td>Copalis Rock</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdn">15</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdn">30</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdn">45</td>
  </tr>
  <tr>
    <td>Point Grenville</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdn">60</td>
    <td class="tdn">30</td>
    <td class="tdn">80</td>
    <td class="tdc">&mdash;</td>
    <td class="tdn">165</td>
    <td class="tdn">40</td>
    <td class="tdn">1,100</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdn">1,475</td>
  </tr>
  <tr>
    <td>Grenville Arch</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdn">30</td>
    <td class="tdn">20</td>
    <td class="tdc">&mdash;</td>
    <td class="tdn">1</td>
    <td class="tdn">60</td>
    <td class="tdc">&mdash;</td>
    <td class="tdn">3,000</td>
    <td class="tdn">4</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdn">3</td>
    <td class="tdn">3,118</td>
  </tr>
  <tr>
    <td>Flat Rock</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdn">30</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdn">60</td>
    <td class="tdn">300</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdn">390</td>
  </tr>
  <tr>
    <td>Split Rock</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdn">100</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdn">1</td>
    <td class="tdn">150</td>
    <td class="tdc">&mdash;</td>
    <td class="tdn">2,100</td>
    <td class="tdn">4</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdn">2,355</td>
  </tr>
  <tr>
    <td>Willoughby Rock</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdn">80</td>
    <td class="tdn">40</td>
    <td class="tdn">15</td>
    <td class="tdc">&mdash;</td>
    <td class="tdn">150</td>
    <td class="tdc">&mdash;</td>
    <td class="tdn">3,000</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdn">25</td>
    <td class="tdn">3,310</td>
  </tr>
  <tr>
    <td>South Rock</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdn">40</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdn">50</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdn">90</td>
  </tr>
  <tr>
    <td>Abbey Islet</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdn">30</td>
    <td class="tdn">3</td>
    <td class="tdc">&mdash;</td>
    <td class="tdn">50</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdn">10</td>
    <td class="tdn">93</td>
  </tr>
  <tr>
    <td>Destruction Island</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdn">12</td>
    <td class="tdn">350</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdn">25</td>
    <td class="tdc">&mdash;</td>
    <td class="tdn">10,940</td>
    <td class="tdn">350</td>
    <td class="tdn">11,677</td>
  </tr>
  <tr>
    <td>Middle Rock</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdn">25</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdn">25</td>
    <td class="tdc">&mdash;</td>
    <td class="tdn">50</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdn">100</td>
  </tr>
  <tr>
    <td>North Rock</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdn">25</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdn">25</td>
  </tr>
  <tr>
    <td>Alexander Island</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdn">50</td>
    <td class="tdn">5</td>
    <td class="tdc">&mdash;</td>
    <td class="tdn">225</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdn">1,550</td>
    <td class="tdn">1,830</td>
  </tr>
  <tr>
    <td>Rounded Island</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdn">25</td>
    <td class="tdc">&mdash;</td>
    <td class="tdn">25</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdn">1</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdn">51</td>
  </tr>
  <tr>
    <td>Giant's Graveyard</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdn">10</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdn">150</td>
    <td class="tdc">&mdash;</td>
    <td class="tdn">50</td>
    <td class="tdn">150</td>
    <td class="tdc">&mdash;</td>
    <td class="tdn">360</td>
  </tr>
  <tr>
    <td>Quillayute Needles</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdn">50</td>
    <td class="tdn">50</td>
    <td class="tdn">50</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdn">150</td>
    <td class="tdn">900</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdn">350</td>
    <td class="tdn">1,550</td>
  </tr>
  <tr>
    <td>James Island</td>
    <td class="tdc">&mdash;</td>
    <td class="tdn">30</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdn">40</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdn">150</td>
    <td class="tdn">750</td>
    <td class="tdn">40</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdn">20</td>
    <td class="tdn">1,030</td>
  </tr>
  <tr>
    <td>Cake Rock</td>
    <td class="tdc">&mdash;</td>
    <td class="tdn">500</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdn">150</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdn">600</td>
    <td class="tdn">300</td>
    <td class="tdn">12</td>
    <td class="tdc">&mdash;</td>
    <td class="tdn">50</td>
    <td class="tdn">1,000</td>
    <td class="tdn">2,612</td>
  </tr>
  <tr>
    <td>Sealion Rock</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdn">70</td>
    <td class="tdc">&mdash;</td>
    <td class="tdn">30</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdn">250</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdn">5</td>
    <td class="tdn">355</td>
  </tr>
  <tr>
    <td>Carroll Island</td>
    <td class="tdc">&mdash;</td>
    <td class="tdn">3,100</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdn">100</td>
    <td class="tdn">3</td>
    <td class="tdc">&mdash;</td>
    <td class="tdn">550</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdn">25</td>
    <td class="tdn">250</td>
    <td class="tdn">2,400</td>
    <td class="tdn">6,428</td>
  </tr>
  <tr>
    <td>Ball Rock</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdn">50</td>
    <td class="tdn">7</td>
    <td class="tdc">&mdash;</td>
    <td class="tdn">150</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdn">750</td>
    <td class="tdn">957</td>
  </tr>
  <tr>
    <td>White Rock</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdn">100</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdn">75</td>
    <td class="tdn">250</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdn">100</td>
    <td class="tdn">525</td>
  </tr>
  <tr>
    <td>Ozette Island</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdn">1</td>
    <td class="tdc">&mdash;</td>
    <td class="tdn">15</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdn">16</td>
  </tr>
  <tr>
    <td>Bodelteh Island</td>
    <td class="tdn">1,900</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdn">100</td>
    <td class="tdn">2</td>
    <td class="tdc">&mdash;</td>
    <td class="tdn">300</td>
    <td class="tdc">&mdash;</td>
    <td class="tdn">5</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdn">750</td>
    <td class="tdn">3,057</td>
  </tr>
  <tr>
    <td>Tatoosh Island</td>
    <td class="tdc">&mdash;</td>
    <td class="tdn">25</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdn">100</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdn">1,500+</td>
    <td class="tdn">100</td>
    <td class="tdn">20</td>
    <td class="tdn">25?</td>
    <td class="tdn">25?</td>
    <td class="tdn">30</td>
    <td class="tdn">1,825</td>
  </tr>
  <tr>
    <td class="tdp">Total</td>
    <td class="tdn">1,900</td>
    <td class="tdn">3,655</td>
    <td class="tdn">45</td>
    <td class="tdn">390</td>
    <td class="tdn">140</td>
    <td class="tdn">995</td>
    <td class="tdn">35</td>
    <td class="tdn">930</td>
    <td class="tdn">4,215</td>
    <td class="tdn">11,950</td>
    <td class="tdn">161</td>
    <td class="tdn">100</td>
    <td class="tdn">11,415</td>
    <td class="tdn">7,343</td>
    <td class="tdn">43,274</td>
  </tr>
</table>
<span class="pagenum"><a name="Page_82" id="Page_82">[Pg 82]</a></span></div>

<div class="center">
<table border="0" cellpadding="4" cellspacing="0" summary="Table 6. Breeding seabird population estimates for the San Juan Islands and Strait of Juan de Fuca, Washington, 1973-75">
    <caption>Table 6. <em>Breeding seabird population estimates for the San Juan Islands and Strait of Juan de Fuca, Washington, 1973-75</em>.<a name="FNanchor_10_10" id="FNanchor_10_10"></a><a href="#Footnote_10_10" class="fnanchor">[10]</a></caption>
  <tr>
    <th rowspan="3">Breeding site</th>
    <th colspan="0">Species</th>
  </tr>
  <tr>
    <th colspan="2">Cor&shy;mo&shy;rants</th>
    <th rowspan="2">Black oys&shy;ter-catch&shy;er</th>
    <th rowspan="2">Glau&shy;cous-wing&shy;ed gull</th>
    <th rowspan="2">Pi&shy;geon guil&shy;le&shy;mot</th>
    <th rowspan="2">Tuft&shy;ed puf&shy;fin</th>
    <th rowspan="2">Rhi&shy;noc&shy;er&shy;os Auk&shy;let</th>
    <th rowspan="2">To&shy;tal</th>
  </tr>
  <tr>
    <th>Dou&shy;ble-crest&shy;ed</th>
    <th>Pe&shy;lag&shy;ic</th>
  </tr>
  <tr>
    <td>Bare Island</td>
    <td class="tdc">&mdash;</td>
    <td class="tdn">50</td>
    <td class="tdn">1</td>
    <td class="tdn">120</td>
    <td class="tdc">+</td>
    <td class="tdc">&mdash;</td>
    <td class="tdn">2</td>
    <td class="tdn">173</td>
  </tr>
  <tr>
    <td>Barren Island</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdn">0</td>
  </tr>
  <tr>
    <td>Battleship Island</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdn">0</td>
  </tr>
  <tr>
    <td>Bird Rocks</td>
    <td class="tdn">30</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">+</td>
    <td class="tdn">320</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdn">350</td>
  </tr>
  <tr>
    <td>Cactus Island</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdn">1</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdn">1</td>
  </tr>
  <tr>
    <td>Castle Island</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdn">30</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdn">30</td>
  </tr>
  <tr>
    <td>Colville Island</td>
    <td class="tdc">&mdash;</td>
    <td class="tdn">40</td>
    <td class="tdn">1</td>
    <td class="tdn">1,000</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdn">1,041</td>
  </tr>
  <tr>
    <td>Danger Island</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdn">125</td>
    <td class="tdn">7</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdn">132</td>
  </tr>
  <tr>
    <td>Decatur Island</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdn">0</td>
  </tr>
  <tr>
    <td>Eliza Island</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdn">3</td>
    <td class="tdn">1</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdn">4</td>
  </tr>
  <tr>
    <td>Eliza Rock</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdn">1</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdn">1</td>
  </tr>
  <tr>
    <td>Flat Top Island</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">+</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">+</td>
  </tr>
  <tr>
    <td>Flower Island</td>
    <td class="tdc">&mdash;</td>
    <td class="tdn">17</td>
    <td class="tdc">&mdash;</td>
    <td class="tdn">90</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdn">107</td>
  </tr>
  <tr>
    <td>Goose Island</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdn">60</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdn">60</td>
  </tr>
  <tr>
    <td>Gull Rock</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">+</td>
    <td class="tdn">125</td>
    <td class="tdn">7</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdn">132</td>
  </tr>
  <tr>
    <td>Gull Reef</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdn">0</td>
  </tr>
  <tr>
    <td>Hall Island</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdn">1</td>
    <td class="tdn">275</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdn">276</td>
  </tr>
  <tr>
    <td>Harbor Rock</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdn">0</td>
  </tr>
  <tr>
    <td>Iceberg Island</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdn">0</td>
  </tr>
  <tr>
    <td>Johns Island</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdn">1</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdn">1</td>
  </tr>
  <tr>
    <td>Long Island</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdn">8</td>
    <td class="tdn">80</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdn">88</td>
  </tr>
  <tr>
    <td>Low Island</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdn">1</td>
    <td class="tdn">75</td>
    <td class="tdn">17</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdn">93</td>
  </tr>
  <tr>
    <td>Lummi Rocks</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdn">4</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdn">4</td>
  </tr>
  <tr>
    <td>Matia Island</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">+</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">+</td>
  </tr>
  <tr>
    <td>Mummy Rocks</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdn">55</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdn">55</td>
  </tr>
  <tr>
    <td>Minor Island</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdn">100</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdn">100</td>
  </tr>
  <tr>
    <td>O'Neal Island</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdn">0</td>
  </tr>
  <tr>
    <td>Patos Island</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdn">20</td>
    <td class="tdc">+</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdn">20</td>
  </tr>
  <tr>
    <td>North Peapod Island</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdn">1</td>
    <td class="tdn">220</td>
    <td class="tdn">2</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdn">223</td>
  </tr>
  <tr>
    <td>South Peapod Island</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdn">1</td>
    <td class="tdn">75</td>
    <td class="tdn">2</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdn">78</td>
  </tr>
  <tr>
    <td>Pearl Island</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdn">0</td>
  </tr>
  <tr>
    <td>Pointer Island</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdn">58</td>
    <td class="tdn">2</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdn">60</td>
  </tr>
  <tr>
    <td>Protection Island</td>
    <td class="tdn">3</td>
    <td class="tdn">110</td>
    <td class="tdn">3</td>
    <td class="tdn">1,500</td>
    <td class="tdn">30</td>
    <td class="tdn">9,200</td>
    <td class="tdn">35</td>
    <td class="tdn">10,881</td>
  </tr>
  <tr>
    <td>Puffin Island</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdn">1</td>
    <td class="tdn">350</td>
    <td class="tdn">15</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdn">366</td>
  </tr>
  <tr>
    <td>Ripple Island</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdn">0</td>
  </tr>
  <tr>
    <td>Sentinel Island</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdn">10</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdn">10</td>
  </tr>
  <tr>
    <td>Sentinel Rock</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdn">1</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdn">1</td>
  </tr>
  <tr>
    <td>Skip Jack Island</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdn">75</td>
    <td class="tdn">20</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdn">95</td>
  </tr>
  <tr>
    <td>Smith Island</td>
    <td class="tdc">&mdash;</td>
    <td class="tdn">20</td>
    <td class="tdn">6</td>
    <td class="tdn">10</td>
    <td class="tdn">30</td>
    <td class="tdn">600</td>
    <td class="tdc">&mdash;</td>
    <td class="tdn">666</td>
  </tr>
  <tr>
    <td>Speiden Island</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdn">0</td>
  </tr>
  <tr>
    <td>South Sister Island</td>
    <td class="tdn">2</td>
    <td class="tdn">11</td>
    <td class="tdn">1</td>
    <td class="tdn">131</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdn">145</td>
  </tr>
  <tr>
    <td>Middle Sister Island</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdn">1</td>
    <td class="tdn">22</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdn">23</td>
  </tr>
  <tr>
    <td>North Sister Island</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdn">2</td>
    <td class="tdn">412</td>
    <td class="tdn">3</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdn">417</td>
  </tr>
  <tr>
    <td>Viti Rocks</td>
    <td class="tdn">29</td>
    <td class="tdn">80</td>
    <td class="tdn">1</td>
    <td class="tdn">387</td>
    <td class="tdn">1</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdn">498</td>
  </tr>
  <tr>
    <td>Waldron Island</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdn">2</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdn">2</td>
  </tr>
  <tr>
    <td>Williamson Rocks</td>
    <td class="tdc">&mdash;</td>
    <td class="tdn">67</td>
    <td class="tdn">1</td>
    <td class="tdn">346</td>
    <td class="tdn">2</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdn">416</td>
  </tr>
  <tr>
    <td>Whale Island</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdn">1</td>
    <td class="tdn">70</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdn">71</td>
  </tr>
  <tr>
    <td>White Rock</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">+</td>
    <td class="tdn">125</td>
    <td class="tdn">13</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdn">138</td>
  </tr>
  <tr>
    <td>Yellow Island</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdn">0</td>
  </tr>
  <tr>
    <td class="tdp">Total per species</td>
    <td class="tdn">64</td>
    <td class="tdn">395</td>
    <td class="tdn">34</td>
    <td class="tdn">6,234</td>
    <td class="tdn">194</td>
    <td class="tdn">9,800</td>
    <td class="tdn">37</td>
    <td class="tdn">16,758</td>
  </tr>
  <tr>
    <td class="tdp">Percent of total population</td>
    <td class="tdn">0.4</td>
    <td class="tdn">2.3</td>
    <td class="tdn">0.2</td>
    <td class="tdn">37.2</td>
    <td class="tdn">1.2</td>
    <td class="tdn">58.5</td>
    <td class="tdn">0.2</td>
    <td class="tdn">100.0</td>
  </tr>
</table><span class="pagenum"><a name="Page_83" id="Page_83">[Pg 83]</a></span></div>

<p>The major colony sites with more than 250
breeding pairs are located at Protection and
Smith islands, Bird Rocks, Colville Island,
Hall Island, North and South Peapod rocks,
Puffin Island, North Sisters, Viti Rocks, and
Williamson Rocks (Fig. 6). Glaucous-winged
gulls are the predominant species on all these
islands except Protection and Smith islands,
where there are large colonies of rhinoceros
auklets. Rhinoceros auklets (65%) and glaucous-winged
gulls (32%) make up 97% of the
total San Juan Islands seabird population.</p>


<h4>Nest-site Preferences</h4>

<p>Food supply and availability of nest sites
are two critically important factors influencing
the distribution and abundance of seabirds.
Whereas information on general diet
composition is known for most seabird
species, we know little about the availability
of favored seabird prey. The dynamics of seabird
food chains is reviewed elsewhere in these
proceedings.</p>

<p>The nest-site preferences for seabirds of the
northeast Pacific Ocean are given in Table 7,
and Table 8 indicates the proportion of seabirds
that belong to specific nest-site categories.
These preferences, in conjunction with
knowledge of the physical characteristics of
seabird habitat, permit a partial explanation
of the present distribution and abundance of
seabirds. For example, if we compare the San
Juan Island habitats with those of the Washington
coast, it is apparent that there are
more cliff-nesting species on the coast. This
reflects the physical characteristics of the two
habitats. There are few cliffs in the San Juan
Islands, and those that exist are very unstable.
Colony sites in the San Juan Islands
are typically on low, flat islands. Glaucous-winged
gulls are the most abundant nesting
species there. Coastal islands, on the other
hand, are either covered by dense vegetation
or are large monolithic chunks of rock with
few available flat areas. Population estimates
for the Washington coast are heavily biased
toward surface nesters, since most of the data
have been gathered by aerial surveys. Consequently,
the burrow and rock crevice categories
are underestimated. The aerial survey
is appropriate for only about 43% of the birds
nesting on the Washington coast.</p>

<div class="center">
<table border="0" cellpadding="4" cellspacing="0" summary="Table 7. Nest-site preference for seabirds breeding from Cape Fairweather, Alaska, to the Columbia River, Washington.">
    <caption>Table 7. <em>Nest-site preference for seabirds breeding from Cape Fairweather, Alaska, to the Columbia River, Washington.</em></caption>
  <tr>
    <th>Nest-site type</th>
    <th>Bird species</th>
  </tr>
  <tr>
    <td colspan="0">Burrow-rock crevice</td>
  </tr>
  <tr>
    <td rowspan="3" class="tdp">Diurnal</td>
    <td>Pigeon guillemot</td>
  </tr>
  <tr>
    <td>Horned puffin</td>
  </tr>
  <tr>
    <td>Tufted puffin</td>
  </tr>
  <tr>
    <td rowspan="6" class="tdp">Nocturnal</td>
    <td>Fork-tailed storm-petrel</td>
  </tr>
  <tr>
    <td>Leach's storm-petrel</td>
  </tr>
  <tr>
    <td>Kittlitz's murrelet</td>
  </tr>
  <tr>
    <td>Ancient murrelet</td>
  </tr>
  <tr>
    <td>Cassin's auklet</td>
  </tr>
  <tr>
    <td>Rhinoceros auklet</td>
  </tr>
  <tr>
    <td colspan="0">Open nests</td>
  </tr>
  <tr>
    <td rowspan="6" class="tdp">Flat or slope</td>
    <td>Double-crested cormorant</td>
  </tr>
  <tr>
    <td>Brandt's cormorant</td>
  </tr>
  <tr>
    <td>Glaucous-winged gull</td>
  </tr>
  <tr>
    <td>Herring gull</td>
  </tr>
  <tr>
    <td>Western gull</td>
  </tr>
  <tr>
    <td>Black oystercatcher</td>
  </tr>
  <tr>
    <td rowspan="3" class="tdp">Cliff face</td>
    <td>Pelagic cormorant</td>
  </tr>
  <tr>
    <td>Common murre</td>
  </tr>
  <tr>
    <td>Black-legged kittiwake</td>
  </tr>
  <tr>
    <td rowspan="0" class="tdp">Tree branch</td>
    <td>Marbled murrelet</td>
  </tr>
</table></div>

<p>Northern and southern British Columbia
provide another good example of habitat
availability as revealed through seabird population
estimates. The population data are
more comprehensive and have largely been
gathered by island visitations. The islands in
the northern portion are heavily vegetated
and many have well-developed soil into which
storm-petrels, auklets, and murrelets can burrow.
Indeed, 96% of the seabird population
consists of nocturnal, burrow-nesting species.
In southern British Columbia, however, there
are more open-nest species, particularly glaucous-winged
gulls and cormorants.</p>

<p>Overall, 68% of the breeding seabirds found
along the northeastern Pacific coast are nocturnal
and nest in burrows or rock crevices
(Table 8). The most conspicuous nesting birds
such as gulls, cormorants, and murres, comprise
only 22% of the total population. Consequently,
our current estimates of breeding
seabirds still underestimate the more secretive,
nocturnal, burrow-nesting species.</p>

<p><span class="pagenum"><a name="Page_84" id="Page_84">[Pg 84]</a></span></p>

<div class="center">
<table border="0" cellpadding="4" cellspacing="0" summary="Table 8. Proportional nest-site preferences of Pacific coast seabirds.">
    <caption>Table 8. <em>Proportional nest-site preferences of Pacific coast seabirds.<a name="FNanchor_11_11" id="FNanchor_11_11"></a><a href="#Footnote_11_11" class="fnanchor">[11]</a></em></caption>
  <tr>
    <th rowspan="3">Site</th>
    <th colspan="4" class="br">Estimated number of pairs</th>
    <th colspan="4" class="br">Percent of population</th>
    <th colspan="0" rowspan="2">Total</th>
  </tr>
  <tr>
    <th colspan="2">British Columbia</th>
    <th rowspan="2">San Juan Islands</th>
    <th rowspan="2">Washington coast</th>
    <th colspan="2">British Columbia</th>
    <th rowspan="2">San Juan Islands</th>
    <th rowspan="2">Washington coast</th>
  </tr>
  <tr>
    <th>Northern</th>
    <th>Southern</th>
    <th>Northern</th>
    <th>Southern</th>
    <th>Population</th>
    <th>Percent</th>
  </tr>
  <tr>
    <td colspan="0">Burrow-rock crevice</td>
  </tr>
  <tr>
    <td class="tdp">Diurnal</td>
    <td class="tdn">1,849</td>
    <td class="tdn">11,334</td>
    <td class="tdn">231</td>
    <td class="tdn">7,504</td>
    <td class="tdn">2.0</td>
    <td class="tdn">18.1</td>
    <td class="tdn">1.4</td>
    <td class="tdn">17.3</td>
    <td class="tdn">20,918</td>
    <td class="tdn">9.7</td>
  </tr>
  <tr>
    <td class="tdp">Nocturnal</td>
    <td class="tdn">90,347</td>
    <td class="tdn">30,600</td>
    <td class="tdn">9,800</td>
    <td class="tdn">17,070</td>
    <td class="tdn">96.0</td>
    <td class="tdn">48.9</td>
    <td class="tdn">58.6</td>
    <td class="tdn">39.4</td>
    <td class="tdn">147,817</td>
    <td class="tdn">68.1</td>
  </tr>
  <tr>
    <td colspan="0">Open nests</td>
  </tr>
  <tr>
    <td class="tdp">Flat or slope</td>
    <td class="tdn">909</td>
    <td class="tdn">15,101</td>
    <td class="tdn">6,298</td>
    <td class="tdn">5,755</td>
    <td class="tdn">1.0</td>
    <td class="tdn">24.2</td>
    <td class="tdn">37.6</td>
    <td class="tdn">13.3</td>
    <td class="tdn">28,063</td>
    <td class="tdn">13.0</td>
  </tr>
  <tr>
    <td class="tdp">Cliff face</td>
    <td class="tdn">982</td>
    <td class="tdn">5,525</td>
    <td class="tdn">395</td>
    <td class="tdn">12,945</td>
    <td class="tdn">1.0</td>
    <td class="tdn">8.8</td>
    <td class="tdn">2.4</td>
    <td class="tdn">30.0</td>
    <td class="tdn">19,847</td>
    <td class="tdn">9.2</td>
  </tr>
  <tr>
    <td class="tdc">Total</td>
    <td class="tdn">94,087</td>
    <td class="tdn">62,560</td>
    <td class="tdn">16,724</td>
    <td class="tdn">43,274</td>
    <td colspan="4"></td>
    <td class="tdn">216,645</td>
  </tr>
</table></div>


<h4>Population Changes</h4>

<p>The available data are inadequate to detect
changes in population distribution and density
for most species (Table 9). In Washington,
for instance, limited unsubstantiated information
suggests an overall decline of the
double-crested cormorant and tufted puffin in
the San Juan Island area. Likewise, there
seems to be an increase in glaucous-winged
gulls there. In British Columbia, Drent and
Guiguet (1961) were able to detect changes in
some species. For example, they noted increases
in the double-crested cormorant, pelagic
cormorant, and glaucous-winged gull. No
change was observed in the tufted puffin.
Since then, the Brandt's cormorant has established
a colony in Barkley Sound (Guiguet
1971). The data in southeastern Alaska are inadequate
for all species except, perhaps, the
Cassin's auklet which Gabrielson and Lincoln
(1959) reported to be declining throughout
Alaska. In short, no definitive statements can
now be made concerning changes in seabird
population numbers.</p>


<h5>Species Accounts</h5>


<h6>Fork-tailed Storm-petrel
<em>(Oceanodroma furcata)</em></h6>

<p>Storm-petrels are especially difficult to census
because they are nocturnal, and the burrows
and rock crevices where they breed are
often difficult to locate, especially in mixed-species
colonies. The census data are inadequate
to determine whether there have been
changes in population density and distribution.
Indeed, the biology of this species is perhaps
the least known of the North Pacific
colonial seabirds. In southeastern Alaska,
this species is outnumbered by at least 5 to 1
by the Leach's storm-petrel <em>(Oceanodroma
leucorhoa)</em>. The reasons for this are poorly
understood. There is some evidence that the
numbers of breeding fork-tailed storm-petrels
on Forrester Island may fluctuate drastically
from one year to the next (Gabrielson and
Lincoln 1959).</p>


<h6>Leach's Storm-petrel <em>(Oceanodroma
leucorhoa)</em></h6>

<p>Of the two subspecies of this petrel (<em>O. l. leucorhoa</em>
and <em>O. l. beali</em>), only <em>O. l. beali</em> is found
in southeastern Alaska. The <em>leucorhoa</em> subspecies
is more northerly in distribution.
Where both fork-tailed and Leach's storm-petrels
are sympatric, Leach's predominates;
however, this relationship becomes more unpredictable
in British Columbia and Washington.
This species is undoubtedly widespread
in the forested islands of the Alexander
Archipelago.</p>


<h6>Double-crested Cormorant
<em>(Phalacrocorax auritus)</em></h6>

<p>The double-crested cormorant apparently
does not breed in southeastern Alaska since
Willett (1912), Gabrielson and Lincoln (1959),
and S. Patten (personal communication) do
not report breeding colonies for the area. The
largest populations occur in southern British
Columbia principally in the Gulf Islands,
where 71% of all breeding double-crested cormorants
are found (Table 10). According to
Jewett et al. (1953), this species was less common
in Puget Sound than was Brandt's cormorant,
but is certainly not the case today
(D. A. Manuwal, unpublished data). The only
common cormorants in the San Juan Islands
are the pelagic and double-crested species.
The double-crested cormorant seems to have
declined in numbers on both coastal and inland
waters. On the basis of his observations,
R. W. Campbell believes that this species is
increasing in British Columbia.</p>

<p><span class="pagenum"><a name="Page_85" id="Page_85">[Pg 85]</a></span></p>

<div class="center">
<table border="0" class="tdc" cellpadding="4" cellspacing="0" summary="Table 9. Distribution and status of marine birds breeding
  along the Pacific coast of Washington, British Columbia, and
  southeastern Alaska.">
  <caption>Table 9. <em>Distribution and status of marine birds breeding
  along the Pacific coast of Washington, British Columbia, and
  southeastern Alaska.</em> (X = known to breed in the region;? =
  data insufficient; + = evidence indicates an overall increase in
  size of population; - = evidence indicates an overall decrease in
  size of population; 0 = no population change.)</caption>
  <tr>
    <th rowspan="2">Family and species</th>
    <th rowspan="2">Common name</th>
    <th colspan="2" class="br">Washington</th>
    <th colspan="2" class="br">British Columbia</th>
    <th colspan="0">Southeastern Alaska</th>
  </tr>
  <tr>
    <th>Presence</th>
    <th>Status</th>
    <th>Presence</th>
    <th>Status</th>
    <th>Presence</th>
    <th>Status</th>
  </tr>
  <tr>
    <td colspan="0" class="tdl">Hydrobatidae</td>
  </tr>
  <tr>
    <td class="tdp"><em>Oceanodroma furcata</em></td>
    <td class="tdl">Fork-tailed storm-petrel</td>
    <td>X</td>
    <td>?</td>
    <td>X</td>
    <td>?</td>
    <td>X</td>
    <td>?</td>
  </tr>
  <tr>
    <td class="tdp"><em>O. leucorhoa</em></td>
    <td class="tdl">Leach's storm-petrel</td>
    <td>X</td>
    <td>?</td>
    <td>X</td>
    <td>-</td>
    <td>X</td>
    <td>?</td>
  </tr>
  <tr>
    <td colspan="0" class="tdl">Phalacrocoracidae</td>
  </tr>
  <tr>
    <td class="tdp"><em>Phalacrocorax auritus</em></td>
    <td class="tdl">Double-crested cormorant</td>
    <td>X</td>
    <td>-</td>
    <td>X</td>
    <td>-</td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td class="tdp"><em>P. penicillatus</em></td>
    <td class="tdl">Brandt's cormorant</td>
    <td>X</td>
    <td>?</td>
    <td>X</td>
    <td>0</td>
    <td>?</td>
    <td></td>
  </tr>
  <tr>
    <td class="tdp"><em>P. pelagicus</em></td>
    <td class="tdl">Pelagic cormorant</td>
    <td>X</td>
    <td>?</td>
    <td>X</td>
    <td>+</td>
    <td>X</td>
    <td>?</td>
  </tr>
  <tr>
    <td colspan="0" class="tdl">Haematopodidae</td>
  </tr>
  <tr>
    <td class="tdp"><em>Haematopus bachmani</em></td>
    <td class="tdl">Black oystercatcher</td>
    <td>X</td>
    <td>?</td>
    <td>X</td>
    <td>+</td>
    <td>X</td>
    <td>?</td>
  </tr>
  <tr>
    <td colspan="0" class="tdl">Laridae</td>
  </tr>
  <tr>
    <td class="tdp"><em>Larus glaucescens</em></td>
    <td class="tdl">Glaucous-winged gull</td>
    <td>X</td>
    <td>+</td>
    <td>X</td>
    <td>+</td>
    <td>X</td>
    <td>?</td>
  </tr>
  <tr>
    <td class="tdp"><em>L. occidentalis</em></td>
    <td class="tdl">Western gull</td>
    <td>X</td>
    <td>?</td>
    <td>X</td>
    <td>?</td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td class="tdp"><em>L. argentatus</em></td>
    <td class="tdl">Herring gull</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td>X</td>
    <td>?</td>
  </tr>
  <tr>
    <td class="tdp"><em>Rissa tridactyla</em></td>
    <td class="tdl">Black-legged kittiwake</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td>X</td>
    <td>?</td>
  </tr>
  <tr>
    <td colspan="0" class="tdl">Alcidae</td>
  </tr>
  <tr>
    <td class="tdp"><em>Uria aalge</em></td>
    <td class="tdl">Common murre</td>
    <td>X</td>
    <td>?</td>
    <td>X</td>
    <td>-</td>
    <td>X</td>
    <td>?</td>
  </tr>
  <tr>
    <td class="tdp"><em>Cepphus columba</em></td>
    <td class="tdl">Pigeon guillemot</td>
    <td>X</td>
    <td>?</td>
    <td>X</td>
    <td>+</td>
    <td>X</td>
    <td>?</td>
  </tr>
  <tr>
    <td class="tdp"><em>Brachyramphus marmoratus</em></td>
    <td class="tdl">Marbled murrelet</td>
    <td>X</td>
    <td>?</td>
    <td>X</td>
    <td>?</td>
    <td>X</td>
    <td>?</td>
  </tr>
  <tr>
    <td class="tdp"><em>B. brevirostris</em></td>
    <td class="tdl">Kittlitz's murrelet</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td>X</td>
    <td>?</td>
  </tr>
  <tr>
    <td class="tdp"><em>Synthliboramphus antiquus</em></td>
    <td class="tdl">Ancient murrelet</td>
    <td>X</td>
    <td>?</td>
    <td>X</td>
    <td>?</td>
  </tr>
  <tr>
    <td class="tdp"><em>Ptychoramphus aleuticus</em></td>
    <td class="tdl">Cassin's auklet</td>
    <td>X</td>
    <td>?</td>
    <td>X</td>
    <td>?</td>
    <td>X</td>
    <td>-</td>
  </tr>
  <tr>
    <td class="tdp"><em>Cerorhinca monocerata</em></td>
    <td class="tdl">Rhinoceros auklet</td>
    <td>X</td>
    <td>?</td>
    <td>X</td>
    <td>+</td>
    <td>X</td>
    <td>?</td>
  </tr>
  <tr>
    <td class="tdp"><em>Fratercula corniculata</em></td>
    <td class="tdl">Horned puffin</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td>X</td>
    <td>?</td>
  </tr>
  <tr>
    <td class="tdp"><em>Lunda cirrhata</em></td>
    <td class="tdl">Tufted puffin</td>
    <td>X</td>
    <td>-</td>
    <td>X</td>
    <td>0</td>
    <td>X</td>
    <td>?</td>
  </tr>
  <tr>
    <td class="tdc">Total species</td>
    <td></td>
    <td>14</td>
    <td></td>
    <td>15</td>
    <td></td>
    <td>16</td>
    <td></td>
  </tr>
</table></div>


<h6>Brandt's Cormorant <em>(Phalacrocorax
penicillatus)</em></h6>

<p>Brandt's cormorant is the least abundant of
the three cormorant species that nest in the
study area. Washington is at the northernmost
edge of the breeding distribution of this
species. Only one more northerly colony
exists, on Sartine Island off Vancouver Island
(Vermeer et al. 1976). Brandt's cormorant
comprises about 85% of the cormorant population
in Oregon (U.S. Fish and Wildlife Service,
unpublished data). However, in Washington
it is only about 9% and in British
Columbia 3% of the total cormorant
population.</p>

<p><span class="pagenum"><a name="Page_86" id="Page_86">[Pg 86]</a></span></p>

<div class="center">
<table border="0" cellpadding="4" cellspacing="0" summary="Table 10. Estimated seabird populations breeding from Cape Fairweather, Alaska, to the Columbia River, Washington.">
  <caption>Table 10. <em>Estimated seabird populations breeding
  from Cape Fairweather, Alaska, to the Columbia River,
  Washington.</em><a name="FNanchor_12_12" id="FNanchor_12_12"></a><a
  href="#Footnote_12_12" class="fnanchor">[12]</a><a
  name="FNanchor_13_13" id="FNanchor_13_13"></a><a
  href="#Footnote_13_13" class="fnanchor">[13]</a><a
  name="FNanchor_14_14" id="FNanchor_14_14"></a><a
  href="#Footnote_14_14" class="fnanchor">[14]</a> (? = present in
  unknown numbers; - = inadequate data.)</caption>
  <tr>
    <th rowspan="2">Bird species</th>
    <th colspan="2" class="br">Northern British Columbia</th>
    <th colspan="2" class="br">Southern British Columbia</th>
    <th colspan="2" class="br">San Juan Islands</th>
    <th colspan="2" class="br">Washington coast</th>
    <th colspan="0">Total all regions</th>
  </tr>
  <tr>
    <th>Pop&shy;u&shy;la&shy;tion</th>
    <th>Per&shy;cent</th>
    <th>Pop&shy;u&shy;la&shy;tion</th>
    <th>Per&shy;cent</th>
    <th>Pop&shy;u&shy;la&shy;tion</th>
    <th>Per&shy;cent</th>
    <th>Pop&shy;u&shy;la&shy;tion</th>
    <th>Per&shy;cent</th>
    <th>Pop&shy;u&shy;la&shy;tion</th>
    <th>Per&shy;cent</th>
  </tr>
  <tr>
    <td>Fork-tailed storm-petrel</td>
    <td class="tdn">49,080</td>
    <td class="tdn">52.2</td>
    <td class="tdc">?</td>
    <td class="tdc">-</td>
    <td class="tdn">0</td>
    <td class="tdc">-</td>
    <td class="tdn">1,900</td>
    <td class="tdn">4.4</td>
    <td class="tdn">50,980</td>
    <td class="tdn">23.5</td>
  </tr>
  <tr>
    <td>Leach's storm-petrel</td>
    <td class="tdn">1,365</td>
    <td class="tdn">1.5</td>
    <td class="tdn">5,000</td>
    <td class="tdn">8.0</td>
    <td class="tdn">0</td>
    <td class="tdc">-</td>
    <td class="tdn">3,655</td>
    <td class="tdn">8.5</td>
    <td class="tdn">10,020</td>
    <td class="tdn">4.6</td>
  </tr>
  <tr>
    <td>Double-crested cormorant</td>
    <td class="tdn">0</td>
    <td class="tdc">-</td>
    <td class="tdn">1,058</td>
    <td class="tdn">1.7</td>
    <td class="tdn">64</td>
    <td class="tdn">&gt;0.1</td>
    <td class="tdn">390</td>
    <td class="tdn">&gt;0.1</td>
    <td class="tdn">1,512</td>
    <td class="tdn">&gt;0.1</td>
  </tr>
  <tr>
    <td>Brandt's cormorant</td>
    <td class="tdn">0</td>
    <td class="tdc">-</td>
    <td class="tdn">185</td>
    <td class="tdn">&gt;0.1</td>
    <td class="tdn">0</td>
    <td class="tdc">-</td>
    <td class="tdn">140</td>
    <td class="tdn">&gt;0.1</td>
    <td class="tdn">325</td>
    <td class="tdn">&gt;0.1</td>
  </tr>
  <tr>
    <td>Pelagic cormorant</td>
    <td class="tdn">982</td>
    <td class="tdn">1.0</td>
    <td class="tdn">4,017</td>
    <td class="tdn">6.4</td>
    <td class="tdn">395</td>
    <td class="tdn">2.4</td>
    <td class="tdn">995</td>
    <td class="tdn">2.3</td>
    <td class="tdn">6,389</td>
    <td class="tdn">3.0</td>
  </tr>
  <tr>
    <td>Glaucous-winged gull</td>
    <td class="tdn">909</td>
    <td class="tdn">1.0</td>
    <td class="tdn">13,858</td>
    <td class="tdn">22.2</td>
    <td class="tdn">6,234</td>
    <td class="tdn">37.3</td>
    <td class="tdn">4,215</td>
    <td class="tdn">9.8</td>
    <td class="tdn">25,216</td>
    <td class="tdn">11.6</td>
  </tr>
  <tr>
    <td>Western gull</td>
    <td class="tdn">0</td>
    <td class="tdc">-</td>
    <td class="tdc">?</td>
    <td class="tdc">-</td>
    <td class="tdn">0</td>
    <td class="tdc">-</td>
    <td class="tdn">930</td>
    <td class="tdn">2.2</td>
    <td class="tdn">930</td>
    <td class="tdn">&gt;0.1</td>
  </tr>
  <tr>
    <td>Common murre</td>
    <td class="tdn">0</td>
    <td class="tdc">-</td>
    <td class="tdn">1,508</td>
    <td class="tdn">2.4</td>
    <td class="tdn">0</td>
    <td class="tdc">-</td>
    <td class="tdn">11,950</td>
    <td class="tdn">27.7</td>
    <td class="tdn">13,458</td>
    <td class="tdn">6.2</td>
  </tr>
  <tr>
    <td>Pigeon guillemot</td>
    <td class="tdn">1,733</td>
    <td class="tdn">1.8</td>
    <td class="tdn">1,256</td>
    <td class="tdn">2.0</td>
    <td class="tdn">194</td>
    <td class="tdn">1.2</td>
    <td class="tdn">161</td>
    <td class="tdn">&gt;0.1</td>
    <td class="tdn">3,345</td>
    <td class="tdn">1.5</td>
  </tr>
  <tr>
    <td>Ancient murrelet</td>
    <td class="tdn">21,177</td>
    <td class="tdn">22.5</td>
    <td class="tdn">0</td>
    <td class="tdc">-</td>
    <td class="tdn">0</td>
    <td class="tdc">-</td>
    <td class="tdn">0</td>
    <td class="tdc">-</td>
    <td class="tdn">21,177</td>
    <td class="tdn">9.8</td>
  </tr>
  <tr>
    <td>Cassin's auklet</td>
    <td class="tdn">13,475</td>
    <td class="tdn">14.3</td>
    <td class="tdn">25,000</td>
    <td class="tdn">40.0</td>
    <td class="tdn">0</td>
    <td class="tdc">-</td>
    <td class="tdn">100</td>
    <td class="tdn">&gt;0.1</td>
    <td class="tdn">38,575</td>
    <td class="tdn">17.8</td>
  </tr>
  <tr>
    <td>Rhinoceros auklet</td>
    <td class="tdn">5,250</td>
    <td class="tdn">5.6</td>
    <td class="tdn">6,000</td>
    <td class="tdn">&gt;0.1</td>
    <td class="tdn">9,800</td>
    <td class="tdn">58.6</td>
    <td class="tdn">11,415</td>
    <td class="tdn">26.4</td>
    <td class="tdn">27,065</td>
    <td class="tdn">12.5</td>
  </tr>
  <tr>
    <td>Horned puffin</td>
    <td class="tdn">0</td>
    <td class="tdc">-</td>
    <td class="tdn">0</td>
    <td class="tdc">-</td>
    <td class="tdn">0</td>
    <td class="tdc">-</td>
    <td class="tdn">0</td>
    <td class="tdc">-</td>
    <td class="tdn">0</td>
    <td class="tdc">-</td>
  </tr>
  <tr>
    <td>Tufted puffin</td>
    <td class="tdn">116</td>
    <td class="tdn">&gt;0.1</td>
    <td class="tdn">10,078</td>
    <td class="tdn">16.1</td>
    <td class="tdn">37</td>
    <td class="tdn">&gt;0.1</td>
    <td class="tdn">7,343</td>
    <td class="tdn">17.0</td>
    <td class="tdn">17,574</td>
    <td class="tdn">8.1</td>
  </tr>
  <tr>
    <td class="tdp">Total</td>
    <td class="tdn">94,087</td>
    <td></td>
    <td class="tdn">67,960</td>
    <td></td>
    <td class="tdn">16,724</td>
    <td></td>
    <td class="tdn">43,194</td>
    <td></td>
    <td class="tdn">216,566</td>
    <td></td>
  </tr>
</table><span class="pagenum"><a name="Page_87" id="Page_87">[Pg 87]</a></span></div>

<p>Comparing information in Jewett et al.
(1953) with the current situation, it is apparent
that there has been a drastic change in
the distribution and probably in the numbers
of this species in Washington. Today, there
are no Brandt's cormorant colonies in the San
Juan Islands or Strait of Juan de Fuca. Yet
Jewett et al. (1953) reported colonies at Bellingham
Bay and on Lopez and Matia islands.
We have observed juvenile Brandt's cormorants
in the San Juan Islands during the
summer. This species may be particularly susceptible
to human disturbance, since all three
areas listed above are heavily used in the summer
for recreation.</p>


<h6>Pelagic Cormorant <em>(Phalacrocorax
pelagicus)</em></h6>

<p>The distribution of breeding colonies of the
pelagic cormorant is strongly determined by
the availability of the steep cliffs on which it
constructs its nest. This is the only common
cormorant in southeastern Alaska. Throughout
its extensive range, this species is generally
found breeding in small numbers. Nothing
is known about fluctuations in its numbers
in Alaska.</p>

<p>This species is common in both British
Columbia and Washington; nesting sites are
of the same type as those in Alaska except in
the San Juan Islands, where 200-300 birds
nest on cliff faces composed of glacial deposits.
Here, there is frequent nest loss due to
slippage off the cliff face; this loss is especially
severe on Smith and Protection islands.
There do not appear to be any changes in the
distribution of pelagic cormorants, but an
accurate assessment of abundance is impossible
from the data currently available.</p>


<h6>Glaucous-winged Gull <em>(Larus glaucescens)</em></h6>

<p>The glaucous-winged gull is the characteristic
gull of southeastern Alaska and British
Columbia. In Washington, it is the dominant
gull in the San Juan Island area but interbreeds
with the western gull on the Washington
outer coast from Tatoosh to Copalis
Beach (Scott 1971). In Alaska, it is widely distributed
and locally abundant on Forrester Island,
St. Lazaria, and throughout Glacier Bay
(S. Patten, personal communication). The
biology of this species has been extensively
studied in the southern part of its range, especially
by Vermeer (1963) and James-Veitch
and Booth (1954). The only study of the breeding
biology of this species in southeastern
Alaska is by Patten (1974) for Glacier Bay.
Glaucous-winged gulls are apparently increasing
in British Columbia (R. W. Campbell, unpublished
data) and in Washington (T. R.
Wahl, personal communication). This increase
is undoubtedly a result of the proximity of
breeding colonies to garbage dumps and commercial
fishing fleets in both Canada and the
United States. Little is known about changes
in populations of gulls in southeastern
Alaska.</p>


<h6>Western Gull <em>(Larus occidentalis)</em></h6>

<p>The western gull is the common breeding
gull of the Washington outer coast; however,
there is increased interbreeding with glaucous-winged
gulls northward from Destruction
Island to Tatoosh Island. The percentage
of glaucous-winged gulls steadily increases
until Vancouver Island and the Strait of Juan
de Fuca, where western gulls are rare. Population
estimates of gulls on the outer coast of
Washington are derived primarily from aerial
flights. This makes identification of gulls difficult,
and in view of the amount of interbreeding,
it is probably impossible to classify many
of the breeding gulls as to species. Western
gulls appear to be increasing in the Grays
Harbor area (G. D. Alcorn, personal communication).</p>


<h6>Herring Gull <em>(Larus argentatus)</em></h6>

<p>The herring gull is typically found in inland
Alaska but can be found uncommonly along
the coast of southeastern Alaska, where it
often forms mixed colonies with glaucous-winged
gulls. These two species apparently
hybridize where they are sympatric (Williamson
and Peyton 1963; Patten and Weisbrod
1974; Patten 1974).</p>


<h6>Black-legged Kittiwake <em>(Rissa tridactyla)</em></h6>

<p>The black-legged kittiwake is found only in
the northern portions of southeastern Alaska.
It apparently is a common breeding bird in
Glacier Bay National Monument (S. M.
Patten, Jr., personal communication). No
population estimates are available for this
species other than that it is locally abundant.</p>

<p><span class="pagenum"><a name="Page_88" id="Page_88">[Pg 88]</a></span></p>


<h6>Common Murre <em>(Uria aalge)</em></h6>

<p>Common murres are common in southeastern
Alaska and the coast of Washington but
breed only in small numbers in British Columbia
and are absent in the San Juan Islands.
Since this species usually prefers cliffs or the
tops of inaccessible rocks, they are probably
limited by island topography in British
Columbia, and are most certainly so limited in
the San Juan and Gulf Island groups.</p>

<p>In Alaska, common murres breed in unknown
numbers in Glacier Bay and in large
numbers on St. Lazaria, Forrester, and the
Hazy islands. No data on population changes
are available for any of the three regions.</p>


<h6>Pigeon Guillemot <em>(Cepphus columba)</em></h6>

<p>The pigeon guillemot is common throughout
the region from Cape Fairweather to
Washington. Even though it is not truly colonial,
it may be locally abundant where there
are suitable nest sites. Since these nest sites
are usually difficult to find, population estimates
are seldom accurate, usually being conservative.
It is evident that guillemots appear
to be small in number when compared with
other seabirds nesting at major colony sites in
the north Pacific region (Table 10). This disparity
may be exaggerated by the difficulty of
censusing guillemots.</p>


<h6>Marbled Murrelet <em>(Brachyramphus
marmoratus)</em></h6>

<p>Since the marbled murrelet has been found
to nest in coniferous forests (Binford et al.
1975), traditional census techniques are unsuitable.
This species is common in southeastern
Alaska (Gabrielson and Lincoln 1959), in
British Columbia (Drent and Guiguet 1961),
and in Washington (Jewett et al. 1953).</p>


<h6>Kittlitz's Murrelet <em>(Brachyramphus
brevirostris)</em></h6>

<p>The difficulties in assessing breeding populations
of Kittlitz's murrelet are the same as
those for the marbled murrelet. This species
nests on the ground at high elevation near the
coast (Bailey 1973). The largest concentrations
are in the vicinity of Glacier Bay National
Monument (Gabrielson and Lincoln
1959). They are not found breeding in Washington
or British Columbia.</p>


<h6>Ancient Murrelet <em>(Synthliboramphus
antiquus)</em></h6>

<p>Ancient murrelets appear to be locally common
throughout southeastern Alaska. Their
presence is probably strongly dependent upon
a suitable soil in which to excavate burrows.
The only available population estimates are
those by Willett (1915) for Forrester Island
(Table 1). Censusing this species is especially
difficult because its burrows are easily confused
with those of Cassin's auklet. There are
no studies of this species in southeastern
Alaska; however, it has been well studied in
the Queen Charlotte Islands to the south by
Sealy (1975).</p>


<h6>Cassin's Auklet <em>(Ptychoramphus
aleuticus)</em></h6>

<p>A synthesis of literature and unpublished
observations led Gabrielson and Lincoln
(1959) to conclude that Cassin's auklet has
greatly decreased in numbers and is not abundant
anywhere in Alaska. They also concluded
that the colony on Forrester Island
(Table 1) was the only well-documented colony
in southeastern Alaska. Fishermen in the
southeastern Alaska area occasionally see
this species (M. E. Isleib, personal communication),
but it is apparently still uncommon
though more widespread than just Forrester
Island. The nocturnal habits and burrowing in
dense vegetation makes censusing this
species very difficult. Nothing is known about
the ecology of this species in Alaska.</p>


<h6>Rhinoceros Auklet <em>(Cerorhinca monocerata)</em></h6>

<p>Rhinoceros auklets seem to be found breeding
only on islands where there is a well-developed
soil in which to excavate their extensive
burrows. From the limited evidence available,
it appears that the largest rhinoceros
auklet populations probably are to be found in
southeastern Alaska. Willett (1912) found a
very large population on Forrester Island
(Table 2), and the species has been found in
the summer in the Barren Islands east of
Kodiak Island (E. P. Bailey, personal communication).
More intensive surveys of the
Alexander Archipelago will probably reveal
other populations of this species.</p>

<p><span class="pagenum"><a name="Page_89" id="Page_89">[Pg 89]</a></span></p>

<p>This species is less common in British
Columbia than either Alaska or Washington.
A possible reason for this is lack of suitable
nesting areas. In Washington, the two largest
colonies are at Protection Island in the Strait
of Juan de Fuca and Destruction Island on
the outer coast. Smaller numbers exist on
other coastal islands and on Smith Island in
the Strait of Juan de Fuca. The Smith Island
colony is an interesting one since it appears
that early human disturbance in the late 19th
or early 20th century eliminated the species
from the island. In their discussion of Smith
Island, Jewett et al. (1953) made no mention
of auklets, only of puffins and guillemots.
Couch (1929) did not record the species in
1925. The colony now numbers about 600
pairs.</p>


<h6>Horned Puffin <em>(Fratercula corniculata)</em></h6>

<p>Although the horned puffin is one of the
most abundant seabirds in other parts of
Alaska, it is much less abundant in the southeastern
portion. In addition to the information
discussed by Sealy (1973), it now appears
that this species may breed as far south as
Triangle Island, British Columbia (K. Vermeer,
personal communication; D. A. Manuwal,
personal observation). Here, as on Forrester
Island, it is greatly outnumbered by
the tufted puffin. No data are available on the
breeding or status of this species in the study
area.</p>


<h6>Tufted Puffin <em>(Lunda cirrhata)</em></h6>

<p>The tufted puffin is found breeding on
scattered islands throughout the region. The
largest known colonies are on Forrester Island,
Alaska, Triangle Island, British Columbia,
and Carroll Island, Washington. It is
notably absent from most of the gulf and San
Juan Islands. Even though puffins have apparently
never been numerous in the San
Juan Islands, their population has noticeably
declined during the past 35 years. For
example, Jewett et al. (1953) reported a
colony of 50 pairs on Bare Island in 1937, but
in 1973 only 2 pairs were counted (D. A.
Manuwal, unpublished data). Likewise, in
1915 there were more than 250 pairs on Smith
Island, but by 1916 there were only 75 pairs
(Jewett et al. 1953). The decline is attributed
to rapid erosion of the glacial-deposit cliffs.
There are no puffins on Smith Island today,
and the largest colony in the Puget Sound
area is the 35 pairs on Protection Island
(D. A. Manuwal, unpublished data).</p>


<h4>Discussion</h4>

<p>The total minimum estimate of the breeding
seabird populations of British Columbia and
Washington is 216,500 pairs (Table 10). No
comprehensive estimates are available for
breeding seabirds of southeastern Alaska. It
is likely, however, that the number of breeding
seabirds in the Alexander Archipelago
may be equal to (or exceed) the populations of
both British Columbia and Washington. Data
are desperately needed from that area. Of the
total seabird population in the study area
(Table 10) 43% reside in northern British
Columbia. The Washington State population
represents 28% of the total. Fork-tailed
storm-petrels comprise almost 25% of all the
breeding seabirds in the area under consideration.
The Cassin's auklet is the next most
numerous species (18% of the total).</p>

<p>It is apparent that current data are, for the
most part, inadequate for assessing anything
but catastrophic changes in seabird breeding
colonies. This inadequacy is due to inadequate
censusing because of excessive reliance upon
aerial surveys; in the past, this has often been
a result of insufficient funding.</p>

<p>Of the several threats facing seabird populations,
none may be as important as oil pollution.
A general review of this subject is presented
elsewhere by Vermeer and Vermeer
(1975). It is apparent from this review that
the most vulnerable species are those that
dive beneath the sea surface, including all the
alcids and cormorants breeding along the
coast that are discussed in this paper. This
group makes up almost 60% of all the breeding
seabirds in this area. Unfortunately, our
knowledge of several of these species is scanty
and our current census techniques are unsuitable
for most of these birds.</p>

<p>Studies of the changes in seabird numbers
have been made in other oceans. For example,
in Great Britain (Bourne 1972<em>a</em>, 1972<em>b</em>; Harris
1970), eastern Canada (Nettleship 1973), and
the Atlantic coast of the United States (Kadlec
and Drury 1968), two major trends seem
apparent. First, there is an overall decline in<span class="pagenum"><a name="Page_90" id="Page_90">[Pg 90]</a></span>
alcid and tern numbers. The decline in auks
may be due to their extreme vulnerability to
oil pollution (Bourne 1972<em>a</em>, 1972<em>b</em>; Vermeer
and Vermeer 1975). The Atlantic puffin, however,
may be suffering the additional effects
of gull cleptoparasitism (Nettleship 1972).
Secondly, there seems to be an increase in gull
populations on both sides of the Atlantic, particularly
the herring gull and black-legged
kittiwake.</p>

<p>Compared with the Atlantic coast of North
America and northern Europe, the data base
for seabird populations of the Pacific coast is
poor. The fragmentary evidence now available
indicates that there may be small population
increases in the western and glaucous-winged
gulls and range extensions of the Brandt's
and double-crested cormorants and of the
rhinoceros auklet (Scott et al. 1974). Whether
these changes represent actual population increases
or displacements remains unclear. The
remote locations of most of the large Pacific
seabird colonies may provide unofficial protection
from human interference. Intensive
surveys are needed to establish base-line inventories
in these areas.</p>

<p>As a consequence of this first comprehensive
review of the status of breeding marine
birds of the northeast Pacific coast of North
America, we recommend the following future
research topics as necessary for the conservation
of this great international resource.</p>

<div class="hangindent">
<p>&#8226; Seabird colony census techniques should
  be refined since almost 68% of the seabirds
  in this area are nocturnal and nest in burrows.
  The present reliance on aerial censusing,
  although economical, is inadequate to
  census most breeding seabird populations;
  more on-site surveys are needed. For surface-nesting
  species and diurnal, burrowing
  species, studies on species specific activity
  cycles are needed so that census data
  can be corrected for birds not observed at
  the colony. For nocturnal, burrowing
  species seasonal burrow occupancy rates
  must be determined so that burrow counts
  can be corrected for inactive burrows.</p>

<p>&#8226; Comprehensive surveys should be made
  every 3-5 years.</p>

<p>&#8226; In 1980 a coordinated breeding bird survey
  of the entire Pacific coasts of Mexico,
  Canada, and the United States should be
  conducted.</p>

<p>&#8226; Specific islands where key populations
  exist should be carefully monitored for
  subtle changes in population density or
  species composition.</p>

<p>&#8226; Increased study of the breeding biology of
  seabirds should be carried out so that base-line
  reproductive characteristics can be
  determined.</p>

<p>&#8226; Detailed studies of the effects of human
  disturbance should be made, especially for
  species that breed near large coastal cities
  or marine recreation areas.</p>
</div>


<h4>References</h4>

<div class="hangindent">

<p>Bailey, E. P. 1973. Discovery of a Kittlitz's murrelet
nest. Condor 75:457.</p>

<p>Binford, L., B. Elliott, and S. Singer. 1975. Discovery
of a nest and the downy young of the
marbled murrelet. Wilson Bull. 87:303-319.</p>

<p>Bourne, W. 1972<em>a</em>. The decline of auks in Great
Britain. Biol. Conserv. 4:144-146.</p>

<p>Bourne, W. 1972<em>b</em>. Threats to seabirds. Int. Counc.
Bird Preserv. Bull. XI:200-218.</p>

<p>Couch, L. 1929. Introduced European rabbits in the
San Juan Islands, Washington. J. Mammal.
10:334-336.</p>

<p>Drent, R. H., and C. Guiguet. 1961. A catalogue of
British Columbia seabird colonies. Occas. Pap.
B. C. Prov. Mus. 12. 173 pp.</p>

<p>Franklin, J., and C. Dyrness. 1973. Natural vegetation
of Oregon and Washington. U.S. For. Serv.
Gen. Tech. Rep. PNW-8. 417 pp.</p>

<p>Gabrielson, I., and F. Lincoln. 1959. The birds of
Alaska. The Stackpole Company, Harrisburg,
Penn., and Wildlife Management Institute,
Washington, D.C. 922 pp.</p>

<p>Grinnell, J. 1897. Petrels of Sitka, Alaska.
Nidologist 4:76-78.</p>

<p>Grinnell, J. 1898. Summer birds of Sitka, Alaska.
Auk 15:122-131.</p>

<p>Grinnell, J. 1909. Birds and mammals of the 1907
Alexander Expedition to southeastern Alaska.
Univ. Calif. Publ. Zool. 5:171-264.</p>

<p>Guiguet, C. 1971. A list of seabird nesting sites in
Barkley Sound, British Columbia. Syesis 4:253-259.</p>

<p>Harris, M. P. 1970. Rates and causes of increases of
some British gull populations. Bird Study 17:325-335.</p>

<p>Heath, H. 1915. Birds observed on Forrester Island,
Alaska during the summer of 1913. Condor
17:20-41.</p>

<p>James-Veitch, E., and E. Booth. 1954. Behavior and
life history of the glaucous-winged gull. Walla
Walla Coll., Publ. Biol. 12.</p>

<p>Jewett, S., W. Taylor, W. Shaw, and J. Aldrich.
1953. Birds of Washington State. Univ. of Washington
Press, Seattle. 767 pp.</p>

<p><span class="pagenum"><a name="Page_91" id="Page_91">[Pg 91]</a></span></p>

<p>Kadlec, J., and W. Drury. 1968. Structure of the
New England herring gull population. Ecology
49:644-676.</p>

<p>Kenyon, K., and V. Scheffer. 1961. Wildlife surveys
along the northwest coast of Washington. Murrelet
42:29-37.</p>

<p>Manuwal, D. A. 1974. Conservation notes (Protection
Island). Pac. Seabird Group Bull. 1.</p>

<p>Nettleship, D. 1972. Breeding success of the common
puffin (<em>Fratercula arctica</em> L.) on different
habitats at Great Island, Newfoundland. Ecol.
Monogr. 42:239-268.</p>

<p>Nettleship, D. 1973. Tenth census of seabirds in the
sanctuaries of the north shore of the Gulf of St.
Lawrence. Can. Field-Nat. 87:395-402.</p>

<p>Patten, S. 1974. Breeding ecology of the glaucous-winged
gull <em>(Larus glaucescens)</em> in Glacier Bay,
Alaska. M.S. Thesis. Univ. of Washington,
Seattle. 78 pp.</p>

<p>Patten, S., and A. R. Weisbrod. 1974. Sympatry
and interbreeding of herring and glaucous-winged
gulls in southeastern Alaska. Condor
76:343.</p>

<p>Richardson, F. 1961. Breeding biology of the
rhinoceros auklet on Protection Island, Washington.
Condor 63:456-473.</p>

<p>Scott, J. M. 1971. Interbreeding of the glaucous-winged
gull and western gull in the Pacific Northwest.
Calif. Birds 2:129-133.</p>

<p>Scott, M., W. Hoffman, D. Ainley, and C. Zeillemaker.
1974. Range expansion and activity patterns
in rhinoceros auklets. West. Birds 5:13-20.</p>

<p>Sealy, S. 1973. Breeding biology of the horned
puffin on St. Lawrence Island, Bering Sea, with
zoogeographical notes on the north Pacific
puffins Pac. Sci. 27:99-199.</p>

<p>Sealy, S. 1975. Feeding ecology of the ancient and
marbled murrelets near Langara Island, British
Columbia. Can. J. Zool. 53:418-433.</p>

<p>Summers, K. 1974. Seabirds breeding along the
east coast of Moresby Island, Queen Charlotte
Islands, British Columbia. Syesis 7:1-12.</p>

<p>Swarth, H. 1911. Birds and mammals of the 1909
Alexander Alaska Expedition. Univ. Calif. Publ.
Zool. 7:9-172.</p>

<p>Swarth, H. 1922. Birds and mammals of the Stikine
River Region of northern British Columbia and
southeastern Alaska. Univ. Calif. Publ. Zool.
24:194-314.</p>

<p>Swarth, H. 1936. Origins of the fauna of the Sitkan
District. Alaska. Proc. Calif. Acad. Sci. Ser. 4,
15:59-78.</p>

<p>Thoresen, A., and J. Galusha. 1971. A nesting population
study of some islands in the Puget Sound
area. Murrelet 52:20-23.</p>

<p>U.S. Department of the Interior, Interagency Task
Force. 1972. Environmental setting between Port
Valdez, Alaska, and west coast ports. Final Environmental
Impact Statement, Vol. 3.</p>

<p>U.S. Weather Bureau. 1956. Climatic summary of
the United States&mdash;supplement for 1931 through
1952, Washington. Climatography of the United
States 11-39. 79 pp.</p>

<p>U.S. Weather Bureau, Environmental Data Service.
1965<em>a</em>. Climatic summary of the United
States&mdash;supplement for 1951 through 1960, Oregon.
Climatography of the United States 86-31.
96 pp.</p>

<p>U.S. Weather Bureau, Environmental Data Service.
1965<em>b</em>. Climatic summary of the United
States&mdash;supplement for 1951 through 1960,
Washington. Climatography of the United States
86-39. 92 pp.</p>

<p>U.S. Weather Bureau, Environmental Data Service.
1974. Climatological Data, Alaska Annual
Summary 1974, 60:13.</p>

<p>Vermeer, K. 1963. The breeding ecology of the glaucous
winged gull <em>(Larus glaucescens)</em> on Mandarte
Island. B.C. Occas. Pap. B.C. Prov. Mus.
13. 104 pp.</p>

<p>Vermeer, K., D. A. Manuwal, and D. S. Bingham.
1976. Seabirds and pinnipeds of Sartine Island,
Scott Island group, British Columbia. Murrelet
57(1):14-16.</p>

<p>Vermeer, K., and R. Vermeer. 1975. Oil threat to
birds on the Canadian west coast. Can. Field-Nat.
89:278-298.</p>

<p>Willett, G. 1912. Report of George Willett, Agent
and Warden stationed on St. Lazaria Bird Reservation,
Alaska. Bird-lore 14:419-426.</p>

<p>Willett, G. 1915. Summer birds of Forrester Island,
Alaska. Condor 19:15-17.</p>

<p>Williamson, F. S. L., and L. Peyton. 1963. Interbreeding
of glaucous-winged and herring gulls in
the Cook Inlet region, Alaska. Condor 65:24-28.</p></div>

<p><span class="pagenum"><a name="Page_92" id="Page_92">[Pg 92]</a></span></p>


<div class="footnotes"><h4>FOOTNOTES:</h4>

<div class="footnote">

<p><a name="Footnote_5_5" id="Footnote_5_5"></a><a href="#FNanchor_5_5"><span class="label">[5]</span></a> Data are minimum estimates of pairs and do not include breeding sites with less than 100 birds.</p></div>

<div class="footnote">

<p><a name="Footnote_6_6" id="Footnote_6_6"></a><a href="#FNanchor_6_6"><span class="label">[6]</span></a> Does not include the black oystercatcher, marbled murrelet, and western gull.</p></div>

<div class="footnote">

<p><a name="Footnote_7_7" id="Footnote_7_7"></a><a href="#FNanchor_7_7"><span class="label">[7]</span></a> Estimates only for colonies of 100 or more birds.</p></div>

<div class="footnote">

<p><a name="Footnote_8_8" id="Footnote_8_8"></a><a href="#FNanchor_8_8"><span class="label">[8]</span></a> Estimates are in number of pairs.</p></div>

<div class="footnote">

<p><a name="Footnote_9_9" id="Footnote_9_9"></a><a href="#FNanchor_9_9"><span class="label">[9]</span></a> Estimates are number of pairs.</p></div>

<div class="footnote">

<p><a name="Footnote_10_10" id="Footnote_10_10"></a><a href="#FNanchor_10_10"><span class="label">[10]</span></a> Estimates are numbers of pairs.</p></div>

<div class="footnote">

<p><a name="Footnote_11_11" id="Footnote_11_11"></a><a href="#FNanchor_11_11"><span class="label">[11]</span></a> Data for southeastern Alaska were inadequate to enable estimates of breeding pairs.</p></div>

<div class="footnote">

<p><a name="Footnote_12_12" id="Footnote_12_12"></a><a href="#FNanchor_12_12"><span class="label">[12]</span></a> Population estimates are minimum and represent numbers of pairs.</p></div>

<div class="footnote">

<p><a name="Footnote_13_13" id="Footnote_13_13"></a><a href="#FNanchor_13_13"><span class="label">[13]</span></a>  Does not include the following species for which population estimates are lacking: black oystercatcher, herring gull, black-legged kittiwake,
marbled murrelet, Kittlitz's murrelet.</p></div>

<div class="footnote">

<p><a name="Footnote_14_14" id="Footnote_14_14"></a><a href="#FNanchor_14_14"><span class="label">[14]</span></a> Data for southeastern Alaska were inadequate to enable estimates of breeding pairs.</p>

<p><span class="pagenum"><a name="Page_93" id="Page_93">[Pg 93]</a></span></p></div></div>

<p><span class="pagenum"><a name="Page_94" id="Page_94">[Pg 94]</a><br /><a name="Page_95" id="Page_95">[Pg 95]</a></span></p>
<div class="chapter"></div>
<hr class="chap" />




<h2><a name="THE_BIOLOGY_AND_ECOLOGY_OF" id="THE_BIOLOGY_AND_ECOLOGY_OF">THE BIOLOGY AND ECOLOGY OF
MARINE BIRDS IN THE NORTH</a></h2>
<div class="chapter"></div>
<hr class="chap" />




<h3><a name="Trophic_Relations_of_Seabirds_in_the_Northeastern" id="Trophic_Relations_of_Seabirds_in_the_Northeastern">Trophic Relations of Seabirds in the Northeastern
Pacific Ocean and Bering Sea</a></h3>


<p class="center">by</p>

<p class="center">David G. Ainley</p>

<p class="center">Point Reyes Bird Observatory<br />
Stinson Beach, California 94970</p>

<p class="center">and</p>

<p class="center">Gerald A. Sanger<a name="FNanchor_15_15" id="FNanchor_15_15"></a><a href="#Footnote_15_15" class="fnanchor">[15]</a></p>

<p class="center">National Marine Fisheries Service<br />
Marine Mammal Division<br />
Seattle, Washington<br />
</p>


<h4>Abstract</h4>

<div class="blockquot">

<p>Literature on the diets of seabirds is reviewed for 70 species found in five subarctic
oceanographic regions of the northeastern North Pacific Ocean and Bering
Sea. Species inhabiting estuaries and sheltered bays are not included. The diets
of cormorants, marine ducks, alcids, and marine raptors are best known; less
information is available for loons, grebes, petrels, and gulls. Enough is known,
however, to broadly characterize the diet of each species. Less than 7% of all
species feed on one type of prey, about 60% feed on two or three types, and the
rest feed on four or more types. Only 12% of all species feed on eight or more
types of prey. Most seabirds (77%) feed as secondary and tertiary carnivores.
Where overlap in diet exists, seabirds partition resources through use of different
feeding methods, selection of different-sized prey, and zonation of habitat.
Species that have specialized diets are probably more susceptible than others to
local environmental catastrophes. Species whose feeding methods are highly
adapted for exploitation of resources in polar and subpolar habitats are not
adapted for coping with oil pollution. Competition between birds and man for
marine resources can sometimes benefit seabirds and at other times harm them.
More research is needed on seabird feeding relations so that the ecological roles
played by marine birds can be defined and placed in perspective. Such work
should be conducted at the community level, year-round, and should be so conducted
as to facilitate comparison with biological oceanographic data.</p></div>

<p>The ecology, morphology, and much of the
behavior of a seabird species are definable in
terms of the food resources it exploits year-round
and the spatial and temporal relations
between food and breeding sites. This general
point unifies such important reports as those
by Kuroda (1954), B&eacute;dard (1969<em>a</em>), Ashmole
and Ashmole (1967), Ashmole (1971), Spring
(1971), and Sealy (1972). More concretely, information
on trophic relations of seabirds is
useful in several ways. In conjunction with
biological oceanographic data, it can provide
insight into geographic location, marine habitat,
depth, time of day, and general method of
food capture by seabirds. Collected over
several years, it can provide a basis for understanding
annual differences in seabird breeding
phenology and success. Finally, supplemented
with data on how much seabirds eat
and excrete, it is necessary for an understanding
of the energetic and ecological roles
played by the birds in the functioning of
marine ecosystems.</p>
<p><span class="pagenum"><a name="Page_96" id="Page_96">[Pg 96]</a></span></p>
<p>Several studies that describe trophic relations
within seabird communities have helped
to define the principals of community organization
pertaining to the exploitation of available
food resources and have given clues to
food-chain pathways. Trophic relations have
been described for breeding communities in
the Barents Sea (Uspenski 1958; Belopol'skii
1961), in the tropical Pacific Ocean (Ashmole
and Ashmole 1967; Ashmole 1968), in the
North Sea (Pearson 1968), and in the Chukchi
Sea (Swartz 1966). The last-named study pertained
most directly to the geographic region
discussed in this paper, but several other
studies have provided sound information on
segments of communities in the northeastern
North Pacific and Bering Sea. These include
the work on three species of auklets (<em>Aethia</em>,
<em>Cyclorrhynchus</em>) in the Bering Sea (B&eacute;dard
1969<em>a</em>); investigations on cormorants and
other fish predators in British Columbia by
Munro (1941), Munro and Clemens (1931), and
Robertson (1974); studies of murres in Bristol
Bay by Ogi and Tsujita (1973); observations
on several species near the Pribilof Islands by
Preble and McAtee (1923); work on diving
species off Oregon by Scott (1973); and
studies of murrelets by Sealy (1975).</p>

<p>A review of available reports reveals three
obvious gaps in the emphasis placed in seabird
food studies. First, few studies have ever
considered in detail the trophic relations of
seabird communities during the winter or nonbreeding
season. Partial exceptions are the
works by Cottam (1939) and others on marine
diving ducks, species that are seabirds only
during the winter, and by several researchers
(Munro and Clemens 1931; Munro 1941;
Robertson 1974) on seabirds in British Columbia.
Divoky (1976) studied diets of pack-ice
gulls during the nonbreeding season, but
those species are not included in the present
analysis because they rarely are found south
of the Bering Strait. Second, no study has
considered the trophic relationships of an entire
seabird community, i.e., not just breeding
species but also nonbreeding species. In the
rather broad communities considered here,
50-70% or more of the birds breed in another
part of the world. To say that these nonbreeding
species have no significant impact on resource
exploitation or on organization and
evolution among breeding members would be
naive. Finally, few investigators have attempted
to fit birds into an entire ecosystem,
including lower trophic level origins as well as
fish, marine mammals, and man.</p>

<p>The reasons for these gaps in study emphasis
are readily apparent: the inconvenience of
marine research during the winter when
weather is stormy, the need for costly study
platforms (boats), and the difficulties in organizing
the specialized community of biologists
required for such tasks. A less obvious
but important reason is that oceanographers
and fishery biologists have overlooked seabirds
as important members of marine
ecosystems.</p>


<h4>Diets of Seabirds in
Western North America</h4>

<p>Relatively good information exists for most
pelecaniformes of the region. A notable exception
is the brown pelican <em>(Pelecanus occidentalis)</em>,
an endangered species. This is unfortunate
because dietary information is important
for understanding the species' ecology. Observations
in eastern North America (Palmer
1962) and Peru (Murphy 1936) indicated that
their diet consisted of fish that occur at the
surface. The larger cormorants are piscivorous,
particularly on schooling fishes that
occur at moderate to great depths (Table 1).
The smaller cormorants feed more heavily on
benthic fish and decapod crustaceans. Cormorants
apparently feed only during daylight
and then only for short periods because their
wettable plumage loses its buoyancy. Thus
they remain relatively close (50 km) to nesting
and loafing areas.</p>

<p><span class="pagenum"><a name="Page_97" id="Page_97">[Pg 97]</a></span></p>

<div class="center">
<table border="0" cellpadding="4" cellspacing="0" summary="Table 1. Food of cormorants in different localities">
<caption>Table 1. <em>Food of cormorants in different localities</em> (x = major prey, o = minor prey and * = incidental prey species)</caption>
<col></col>
<col span="25" width="3%"></col>
  <tr>
    <th rowspan="3">Lo&shy;ca&shy;tion</th>
    <th colspan="0">Di&shy;et</th>
  </tr>
  <tr>
    <th colspan="3" class="br">CRUS&shy;TA&shy;CEAN</th>
    <th colspan="0">FISH</th>
  </tr>
  <tr>
    <th>A<br />m<br />p<br />h<br />i<br />p<br />o<br />d<br /></th>
    <th>I<br />s<br />o<br />p<br />o<br />d<br /></th>
    <th>D<br />e<br />c<br />a<br />p<br />o<br />d<br /></th>
    <th><em>C<br />l<br />u<br />p<br />e<br />a<br /></em></th>
    <th><em>E<br />n<br />g<br />r<br />a<br />u<br />l<br />i<br />s<br /></em></th>
    <th>S<br />a<br />l<br />m<br />o<br />n<br />i<br />d<br /></th>
    <th>A<br />r<br />g<br />e<br />n<br />t<br />i<br />n<br />i<br />d<br /></th>
    <th><em>P<br />o<br />r<br />i<br />c<br />h<br />t<br />h<br />y<br />s<br /></em></th>
    <th><em>O<br />t<br />o<br />p<br />h<br />i<br />d<br />i<br />u<br />m<br /></em></th>
    <th><em>B<br />o<br />r<br />e<br />o<br />g<br />a<br />d<br />u<br />s<br /></em></th>
    <th><em>M<br />i<br />c<br />r<br />o<br />g<br />a<br />d<br />u<br />s<br /></em></th>
    <th><em>G<br />a<br />s<br />t<br />e<br />r<br />o<br />s<br />t<br />e<br />u<br />s<br /></em></th>
    <th><em>S<br />e<br />b<br />a<br />s<br />t<br />e<br />s<br /></em></th>
    <th>H<br />e<br />x<br />a<br />g<br />r<br />a<br />m<br />m<br />i<br />d<br /></th>
    <th>C<br />o<br />t<br />t<br />i<br />d<br /></th>
    <th>A<br />g<br />o<br />n<br />i<br />d<br /></th>
    <th>E<br />m<br />b<br />i<br />o<br />t<br />o<br />c<br />i<br />d<br /></th>
    <th><em>C<br />h<br />r<br />o<br />m<br />i<br />s<br /></em></th>
    <th><em>O<br />x<br />y<br />j<br />u<br />l<br />i<br />s<br /></em></th>
    <th>S<br />t<br />i<br />c<br />h<br />a<br />c<br />i<br />d<br /></th>
    <th>P<br />h<br />o<br />l<br />i<br />d<br /></th>
    <th>G<br />o<br />b<br />i<br />i<br />d<br /></th>
    <th><em>A<br />m<br />m<br />o<br />d<br />y<br />t<br />e<br />s<br /></em></th>
    <th>P<br />l<br />e<br />u<br />r<br />o<br />n<br />e<br />c<br />t<br />i<br />d<br /></th>
    <th>B<br />o<br />t<br />h<br />i<br />d<br /></th>
  </tr>
  <tr>
    <td colspan="0">Double-crested cormorant <em>(Phalacrocorax auritus)</em><a name="FNanchor_16_16" id="FNanchor_16_16"></a><a href="#Footnote_16_16" class="fnanchor">[16]</a></td>
  </tr>
  <tr>
    <td class="tdp">Alaska Peninsula (Palmer 1962)</td>
    <td class="tdc">o</td>
    <td class="tdc">o</td>
    <td class="tdc">o</td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">o</td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
  </tr>
  <tr>
    <td class="tdp">SE Alaska (Bailey 1927)</td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td class="tdp">Mandarte Island (Robertson 1974)</td>
    <td></td>
    <td></td>
    <td class="tdc">*</td>
    <td class="tdc">*</td>
    <td class="tdc">*</td>
    <td class="tdc">*</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">*</td>
    <td></td>
    <td></td>
    <td class="tdc">*</td>
    <td></td>
    <td class="tdc">o</td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td class="tdp">Vancouver Island (Munro and Clemens 1931)</td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td class="tdp">Oregon (Palmer 1962)</td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td class="tdc">*</td>
    <td></td>
    <td></td>
    <td class="tdc">*</td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td class="tdc">*</td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">*</td>
    <td></td>
    <td class="tdc">*</td>
    <td></td>
  </tr>
  <tr>
    <td class="tdp">Farallon Island (PRBO, unpublished data)</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">*</td>
    <td></td>
    <td></td>
    <td class="tdc">*</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">*</td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">*</td>
    <td></td>
    <td class="tdc">*</td>
    <td class="tdc">*</td>
  </tr>
  <tr>
    <td colspan="0">Brandt's cormorant <em>(P. penicillatus)</em><a name="FNanchor_17_17" id="FNanchor_17_17"></a><a href="#Footnote_17_17" class="fnanchor">[17]</a></td>
  </tr>
  <tr>
    <td class="tdp">Vancouver Island (Robertson, unpublished data)</td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">o</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td class="tdp">Vancouver Island (Munro and Clemens 1931)</td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td class="tdp">Washington (Jewett et al. 1953)</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
  </tr>
  <tr>
    <td class="tdp">Yaquina Head (Scott 1973)</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td class="tdc">o</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">o</td>
    <td class="tdc">*</td>
    <td class="tdc">o</td>
    <td></td>
    <td class="tdc">o</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">o</td>
    <td></td>
  </tr>
  <tr>
    <td class="tdp">Farallon Island (PRBO, unpublished data)</td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">*</td>
    <td class="tdc">o</td>
    <td class="tdc">*</td>
    <td></td>
    <td class="tdc">*</td>
    <td class="tdc">x</td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td class="tdc">x</td>
    <td class="tdc">*</td>
    <td class="tdc">o</td>
    <td></td>
    <td class="tdc">*</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">*</td>
    <td class="tdc">*</td>
  </tr>
  <tr>
    <td class="tdp">San Diego (Hubbs et al. 1970)</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">*</td>
    <td></td>
    <td class="tdc">*</td>
    <td></td>
    <td class="tdc">o</td>
    <td class="tdc">o</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">*</td>
    <td class="tdc">x</td>
  </tr>
  <tr>
    <td colspan="0">Pelagic cormorant <em>(P. pelagicus)</em><a name="FNanchor_18_18" id="FNanchor_18_18"></a><a href="#Footnote_18_18" class="fnanchor">[18]</a></td>
  </tr>
  <tr>
    <td class="tdp">Cape Thompson (Swartz 1966)</td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td class="tdp">Pribilof Island (Preble and McAtee 1923)</td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td class="tdp">Alaska (Palmer 1962)</td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td class="tdc">x</td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td class="tdc">x</td>
    <td class="tdc">x</td>
    <td></td>
  </tr>
  <tr>
    <td class="tdp">SE Alaska (Heath 1915)</td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td class="tdp">Mandarte Island (Robertson 1974)</td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">*</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">*</td>
    <td class="tdc">x</td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td class="tdp">Vancouver Island (Munro and Clemens 1921)</td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td class="tdp">Washington (Jewett et al. 1953)</td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
  </tr>
  <tr>
    <td class="tdp">Netarts, Oregon (Gabrielson and Jewett 1940)</td>
    <td class="tdc">*</td>
    <td class="tdc">*</td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td class="tdp">Yaquina Head (Scott 1973)</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">o</td>
    <td class="tdc">o</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td class="tdp">Farallon Island (PRBO, unpublished data)</td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td colspan="0">Red-faced cormorant <em>(P. urile)</em></td>
  </tr>
  <tr>
    <td class="tdp">Pribilof Islands (Preble and McAtee 1923)</td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td class="tdc">x</td>
    <td class="tdc">x</td>
    <td></td>
  </tr>
</table></div>

<div class="pagenum"><a name="Page_98" id="Page_98">[Pg 98]</a></div>

<div class="center">
<table border="0" cellpadding="4" cellspacing="0" summary="Table 2. Food of marine ducks and geese">
    <caption>Table 2. <em>Food of marine ducks and geese (x = major prey, o = minor prey, and * = incidental prey species</em>).<a name="FNanchor_19_19" id="FNanchor_19_19"></a><a href="#Footnote_19_19" class="fnanchor">[19]</a></caption>
  <tr>
    <th rowspan="3">Location</th>
    <th colspan="0">Diet<a href="#Footnote_19_19" class="fnanchor">[19]</a></th>
  </tr>
  <tr>
    <th rowspan="2" class="tdt">PLANTS</th>
    <th colspan="3" class="br">CRUS&shy;TA&shy;CEANS</th>
    <th colspan="6">MOL&shy;LUSCS</th>
    <th rowspan="2" class="tdt">ECHI&shy;NO&shy;DERMS</th>
    <th rowspan="2" class="tdt">FISH</th>
    <th rowspan="2" class="tdt">FISH EGGS</th>
  </tr>
  <tr>
    <th>Am&shy;phi&shy;pods</th>
    <th>Dec&shy;a&shy;pods</th>
    <th>Bar&shy;na&shy;cles</th>
    <th>Mus&shy;sels</th>
    <th>Rock clams</th>
    <th>Ra&shy;zor clams</th>
    <th>Oys&shy;ters, Scal&shy;lops</th>
    <th>Lit&shy;tor&shy;i&shy;nids</th>
    <th>Chi&shy;tons</th>
  </tr>
  <tr>
    <td><strong>Geese</strong></td>
    <td class="tdp">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td colspan="0" class="tdp">(<em>Branta</em> spp.)</td>
  </tr>
  <tr>
    <td><strong>Emperor goose</strong></td>
    <td class="tdp">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td colspan="0" class="tdp"><em>(Philacte canagica)</em></td>
  </tr>
  <tr>
    <td><strong>Oldsquaw</strong></td>
    <td class="tdp">o</td>
    <td></td>
    <td></td>
    <td class="tdc">*</td>
    <td class="tdc">*</td>
    <td class="tdc">*</td>
    <td class="tdc">*</td>
    <td class="tdc">*</td>
    <td class="tdc">*</td>
    <td></td>
    <td class="tdp">*</td>
    <td class="tdp">o</td>
  </tr>
  <tr>
    <td colspan="0" class="tdp"><em>(Clangula hyemalis)</em></td>
  </tr>
  <tr>
    <td><strong>Harlequin duck</strong></td>
    <td class="tdp">*</td>
    <td class="tdc">x</td>
    <td class="tdc">x</td>
    <td class="tdc">o</td>
    <td class="tdc">*</td>
    <td></td>
    <td></td>
    <td class="tdc">*</td>
    <td class="tdc">o</td>
    <td class="tdc">o</td>
    <td class="tdp">*</td>
    <td class="tdp">*</td>
  </tr>
  <tr>
    <td colspan="0" class="tdp"><em>(Histrionicus histrionicus)</em></td>
  </tr>
  <tr>
    <td><strong>Steller's eider</strong></td>
    <td class="tdp">o</td>
    <td class="tdc">x</td>
    <td class="tdc">*</td>
    <td class="tdc">*</td>
    <td class="tdc">*</td>
    <td></td>
    <td class="tdc">*</td>
    <td class="tdc">o</td>
    <td class="tdc">*</td>
    <td class="tdc">*</td>
    <td class="tdp">*</td>
    <td class="tdp">*</td>
  </tr>
  <tr>
    <td colspan="0" class="tdp"><em>(Polysticta stelleri)</em></td>
  </tr>
  <tr>
    <td><strong>Common eider</strong></td>
    <td class="tdp">*</td>
    <td class="tdc">*</td>
    <td class="tdc">x</td>
    <td class="tdc">*</td>
    <td class="tdc">o</td>
    <td></td>
    <td class="tdc">*</td>
    <td class="tdc">o</td>
    <td class="tdc">o</td>
    <td class="tdc">*</td>
    <td class="tdp">o</td>
    <td class="tdp">*</td>
  </tr>
  <tr>
    <td colspan="0" class="tdp"><em>(Somateria mollissima)</em></td>
  </tr>
  <tr>
    <td><strong>King eider</strong></td>
    <td class="tdp">*</td>
    <td class="tdc">*</td>
    <td class="tdc">o</td>
    <td class="tdc">*</td>
    <td class="tdc">x</td>
    <td></td>
    <td class="tdc">*</td>
    <td class="tdc">o</td>
    <td class="tdc">o</td>
    <td class="tdc">*</td>
    <td class="tdp">x</td>
    <td class="tdp">*</td>
  </tr>
  <tr>
    <td colspan="0" class="tdp"><em>(S. spectabilis)</em></td>
  </tr>
  <tr>
    <td><strong>Spectacled eider</strong></td>
    <td class="tdp">x</td>
    <td class="tdc">*</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td class="tdc">o</td>
    <td class="tdc">*</td>
    <td class="tdc">*</td>
    <td class="tdp">*</td>
    <td class="tdp">*</td>
  </tr>
  <tr>
    <td colspan="0" class="tdp"><em>(S. fischeri)</em></td>
  </tr>
  <tr>
    <td><strong>White-winged scoter</strong></td>
    <td class="tdp">*</td>
    <td class="tdc">*</td>
    <td class="tdc">*</td>
    <td class="tdc">*</td>
    <td class="tdc">o</td>
    <td class="tdc">x</td>
    <td class="tdc">*</td>
    <td class="tdc">x</td>
    <td class="tdc">o</td>
    <td class="tdc">*</td>
    <td class="tdp">*</td>
    <td class="tdp">o</td>
    <td class="tdp">o</td>
  </tr>
  <tr>
    <td colspan="0" class="tdp"><em>(Melanitta deglandi)</em></td>
  </tr>
  <tr>
    <td><strong>Surf scoter</strong></td>
    <td class="tdp">o</td>
    <td class="tdc">*</td>
    <td class="tdc">*</td>
    <td class="tdc">*</td>
    <td class="tdc">x</td>
    <td class="tdc">*</td>
    <td class="tdc">*</td>
    <td class="tdc">x</td>
    <td class="tdc">o</td>
    <td class="tdc">*</td>
    <td class="tdp">*</td>
    <td class="tdp">*</td>
    <td class="tdp">o</td>
  </tr>
  <tr>
    <td colspan="0" class="tdp"><em>(M. perspicillata)</em></td>
  </tr>
  <tr>
    <td><strong>Black scoter</strong></td>
    <td class="tdp">o</td>
    <td class="tdc">*</td>
    <td class="tdc">*</td>
    <td class="tdc">o</td>
    <td class="tdc">x</td>
    <td class="tdc">*</td>
    <td class="tdc">*</td>
    <td class="tdc">x</td>
    <td class="tdc">*</td>
    <td class="tdc">*</td>
    <td class="tdp">*</td>
    <td class="tdp">*</td>
  </tr>
  <tr>
    <td colspan="0" class="tdp"><em>(M. nigra)</em></td>
  </tr>
  <tr>
    <td><strong>Red-breasted merganser</strong></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdp">x</td>
  </tr>
  <tr>
    <td colspan="0" class="tdp"><em>(Mergus serrator)</em></td>
  </tr>
</table></div>

<div class="pagenum"><a name="Page_99" id="Page_99">[Pg 99]</a></div>

<p>Information on diets of marine ducks
(Table 2) is more nearly complete than for
most other seabirds. These birds fall into four
groups with some overlap: species feeding on
plants (<em>Branta, Philacte</em>, <em>Anas</em>-type, and
<em>Somateria fischeri</em>); those feeding on benthic
crustaceans (<em>Clangula hyemalis</em>, <em>Histrionicus
histrionicus</em>, <em>Polysticta stelleri</em>, <em>S. mollissima</em>);
those feeding on benthic molluscs
(<em>Somateria</em> spp. and <em>Melanitta</em> spp.); and
those feeding on fish (<em>Mergus serrator</em>, <em>Clangula
hyemalis</em>, and <em>Melanitta deglandi</em>). A
study by Perthon (1968), one of the few on a
seabird's diet during most of a year, showed a
seasonal change in diet for <em>S. mollissima</em> in
Norway. In general, waterfowl seem to
specialize in their diets much more than other
seabirds and, for that reason, are perhaps
more restricted in their distributions. Some
marine ducks are known to dive to considerable
depths (reviewed by Kooyman 1974), but
usually they occur in shallow waters where
plants and sessile invertebrates are readily
available.</p>

<p>The summer diet of the pigeon guillemot
<em>(Cepphus columba)</em> is the best known among
seabirds in the region being considered here
(Table 3). Only in the extreme southern part
of its range (i.e., the California Channel Islands)
is there no information available on its
diet. The species feeds on organisms, mostly
fish, from rocky habitat and apparently can
dive to considerable depths (Follett and Ainley
1976). Because so much is known about
guillemot diets during summer, a study of the
winter diet would be valuable.</p>

<p>The diets of other alcids are known well
enough to at least characterize them broadly.
The larger species, murres, tufted and horned
puffins (<em>Lunda cirrhata</em>, <em>Fratercula corniculata</em>),
and the rhinoceros auklet <em>(Cerorhinca
monocerata)</em>, feed heavily on fish, mainly
species that school in midwater (Table 4). To a
great degree, these birds are opportunistic,
feeding rather heavily at times on cephalopods
and crustaceans, particularly nektonic
forms. Morphological differences between the
two murre species suggest that thick-billed
murres <em>(Uria lomvia)</em> feed on benthic organisms
much more than do common murres
<em>(U. aalge)</em>, and that the latter species is more
piscivorous (Spring 1971); however, field data
on diets are barely adequate to confirm this.
Ogi and Tsujita (1973) analyzed the stomach
contents of murres drowned in salmon gill
nets but did not separate the two species. For
the present paper we considered them to be
mostly <em>U. aalge</em>, since this species predominates
in the region of the food study (Bartonek
and Gibson 1972). Adult murres sometimes
eat different items than they feed to
their chicks (Spring 1971; Scott 1973). The
smaller alcids, ancient and marbled murrelets&mdash;<em>Synthliboramphus
antiquus</em> and
<em>Brachyramphus marmoratus</em>&mdash;(Table 5) and
auklets (Table 6), feed on macrozooplankton:
crustaceans, and fish and squid larvae. Little
is known about the food or feeding ecology of
Kittlitz's murrelet <em>(B. brevirostris)</em>. Its diet is
probably similar to that of the other murrelets,
especially the marbled murrelet, its allopatric
congener, but the diets of the other
murrelets differ somewhat (B&eacute;dard 1969<em>b</em>;
Sealy 1975). The Kittlitz's murrelet's shorter
bill suggests that it feeds more on invertebrates.
Alcids feed in deep or shallow water,
depending on food distribution. Some alcid
species can be found at great distances from
land, particularly in winter (Hamilton 1958;
Scott et al. 1971).</p>

<p>Information on the diets of other seabirds in
the region is fragmentary and sometimes
rather anecdotal. A little is known about the
feeding habits of loons (<em>Gavia</em> spp.) and
grebes (<em>Podiceps</em> spp. and <em>Aechmophorus
occidentalis</em>), especially off British Columbia
(Table 7). The larger of these birds feed mainly
on inshore fish, but as species become progressively
smaller, there is a tendency toward
eating crustaceans. Work by Madsen (1957) in
Denmark, indicated that loons and grebes
tend to take prey near or on the bottom. Much
more information is available on these birds'
diets at their freshwater breeding sites but
this provides only partial insight into what
they might eat in marine habitats.</p>

<p>Information is especially poor for albatrosses
and petrels (order Procellariiformes)
(Table 8). Yet, based on sheer numbers alone,
members of this diverse group are easily
among the most ecologically dominant of the
region (Sanger 1972; Ainley 1977). The
Laysan albatross <em>(Diomedea immutabilis)</em>
seems to be a squid specialist; the black-footed
albatross <em>(D. nigripes)</em>, northern fulmar
<em>(Fulmarus glacialis)</em>, scaled petrel <em>(Pterodroma
inexpectata)</em>, and the fork-tailed and
Leach's storm-petrels (<em>Oceanodroma furcata</em>
and <em>O. leucorhoa</em>) appear to be large,
medium, small, and tiny versions, respectively,
of surface-feeding generalists that eat
whatever they can find, including live and
dead fish, squid, coelenterates, crustaceans,
and other organisms. The shearwaters
(<em>Puffinus</em> spp.) feed to an unknown degree on
schooling fish, squid, and crustaceans that
occur near the surface. For these very abundant
shearwaters, that, unfortunately, is
close to the extent of our knowledge both for
the North Pacific, where they winter, and the
South Pacific, where they breed. Most petrels
remain in oceanic habitats, but shearwaters,
particularly the sooty shearwater <em>(Puffinus
griseus)</em>, and sometimes fulmars feed close to,
if not within, the inshore neritic habitat. A
much better understanding of the diets of this
group is sorely needed.</p>

<p><span class="pagenum"><a name="Page_100" id="Page_100">[Pg 100]</a></span></p>

<div class="center">
<table border="0" cellpadding="4" cellspacing="0" summary="Table 3. Food of the pigeon guillemot (Cepphus columba) in different localities">
    <caption>Table 3. <em>Food of the pigeon guillemot</em> (Cepphus columba) <em>in different localities (x = major prey, o = minor prey, and * = incidental prey species)</em>.</caption>
<col></col>
<col span="23" width="3%"></col>
  <tr>
    <th rowspan="3">Lo&shy;ca&shy;tion</th>
    <th colspan="0">Di&shy;et</th>
  </tr>
  <tr>
    <th colspan="3">CRUS&shy;TA&shy;CEAN</th>
    <th rowspan="2" class="tdt">O<br />C<br />T<br />O<br />P<br />U<br />S</th>
    <th colspan="0">FISH</th>
  </tr>
  <tr>
    <th>A<br />m<br />p<br />h<br />i<br />p<br />o<br />d</th>
    <th>I<br />s<br />o<br />p<br />o<br />d</th>
    <th>D<br />e<br />c<br />a<br />p<br />o<br />d</th>
    <th>P<br />e<br />t<br />r<br />o<br />m<br />y<br />z<br />o<br />n<br />t<br />i<br />d</th>
    <th>C<br />h<br />i<br />m<br />a<br />e<br />r<br />i<br />d</th>
    <th>C<br />l<br />u<br />p<br />e<br />i<br />d</th>
    <th>O<br />s<br />m<br />e<br />r<br />i<br />d</th>
    <th>G<br />a<br />d<br />i<br />d</th>
    <th>G<br />a<br />s<br />t<br />e<br />r<br />o<br />s<br />t<br />e<br />i<br />d</th>
    <th>S<br />c<br />o<br />r<br />p<br />a<br />e<br />n<br />i<br />d</th>
    <th>C<br />o<br />t<br />t<br />i<br />d</th>
    <th>A<br />g<br />o<br />n<br />i<br />d</th>
    <th>E<br />m<br />b<br />i<br />o<br />t<br />o<br />c<br />i<br />d</th>
    <th>B<br />a<br />t<br />h<br />y<br />m<br />a<br />s<br />t<br />e<br />r<br />i<br />d</th>
    <th>C<br />l<br />i<br />n<br />i<br />d</th>
    <th>C<br />r<br />y<br />p<br />t<br />a<br />c<br />a<br />n<br />t<br />h<br />o<br />d<br />i<br />d</th>
    <th>C<br />e<br />b<br />i<br />d<br />i<br />c<br />h<br />t<br />h<br />y<br />i<br />d</th>
    <th>S<br />t<br />i<br />c<br />h<br />a<br />e<br />i<br />d</th>
    <th>P<br />h<br />o<br />l<br />i<br />d</th>
    <th>A<br />m<br />m<br />o<br />n<br />d<br />y<br />t<br />i<br />d</th>
    <th>B<br />o<br />t<br />h<br />i<br />d</th>
    <th>P<br />l<br />e<br />u<br />r<br />o<br />n<br />e<br />c<br />t<br />i<br />d</th>
  </tr>
  <tr>
    <td>Cape Thompson (Swartz 1966)</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">o</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">o</td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td>Pribilof Island (Preble and McAttee 1923)</td>
    <td class="tdc">o</td>
    <td class="tdc">o</td>
    <td class="tdc">o</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td>Mandarte Island (Drent 1965; Koelink 1972)</td>
    <td></td>
    <td></td>
    <td class="tdc">o</td>
    <td></td>
    <td class="tdc">o</td>
    <td></td>
    <td class="tdc">*</td>
    <td></td>
    <td></td>
    <td class="tdc">o</td>
    <td></td>
    <td class="tdc">x</td>
    <td class="tdc">o</td>
    <td class="tdc">o</td>
    <td class="tdc">*</td>
    <td class="tdc">o</td>
    <td class="tdc">*</td>
    <td class="tdc">o</td>
    <td class="tdc">o</td>
    <td class="tdc">o</td>
    <td class="tdc">o</td>
    <td class="tdc">o</td>
    <td class="tdc">o</td>
  </tr>
  <tr>
    <td>Vancouver Island (Munro and Clemens 1931)</td>
    <td></td>
    <td></td>
    <td class="tdc">o</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">o</td>
    <td class="tdc">o</td>
  </tr>
  <tr>
    <td>Olympic Peninsula (Thoresen and Booth 1958)</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">o</td>
    <td></td>
    <td></td>
    <td class="tdc">o</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">o</td>
    <td class="tdc">o</td>
    <td></td>
  </tr>
  <tr>
    <td>Yaquina Head (Scott 1973)</td>
    <td></td>
    <td></td>
    <td class="tdc">o</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">o</td>
    <td class="tdc">o</td>
    <td></td>
    <td class="tdc">o</td>
    <td class="tdc">o</td>
  </tr>
  <tr>
    <td>Farallon Island (Follett and Ainley 1976)</td>
    <td></td>
    <td></td>
    <td class="tdc">o</td>
    <td class="tdc">o</td>
    <td></td>
    <td class="tdc">*</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td class="tdc">x</td>
    <td class="tdc">*</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">o</td>
    <td class="tdc">o</td>
    <td></td>
    <td class="tdc">o</td>
    <td class="tdc">o</td>
  </tr>
</table><span class="pagenum"><a name="Page_101" id="Page_101">[Pg 101]</a></span></div>

<p>Knowledge on the food of gulls, shorebirds,
and related species is surprisingly scanty in
view of all that is known about their breeding
biology and social behavior. Little is known
about the marine food of phalaropes, but by
inference from their association with storm-petrels,
plankton-feeding whales, and convergence
lines (Martin and Myers 1969), their
tiny size, and their method of feeding (picking
at minuscule items on the water surface), one
can guess that they feed on zooplankton and
detritus. Skuas <em>(Catharacta skua)</em> and jaegers
(<em>Stercorarius</em> spp.) apparently eat what they
can find at the surface, as well as whatever
they can steal from gulls and terns. Almost all
the literature on their feeding (Bent 1946)
dwells on accounts of their stealing from
other birds. That spectacular behavior would
seem to be so energetically costly, though,
that it is probably less important than we
have been led to believe. Rather surprisingly,
the question of what foods the gulls and terns
eat in the eastern North Pacific is difficult to
answer from the literature (Tables 9 and 10).
Some information exists for five of the larger
larids at isolated places, but little is known
about food elsewhere in their respective
ranges, and the diets of the seven smaller
gulls and the terns are practically unknown.
Studies on gull diets in the Atlantic region
(e.g., Spaans 1971; Harris 1965) provide information
on what to expect from the same
species in the Pacific, but that information
must be considered only in general terms because,
the birds being somewhat opportunistic,
their diets differ greatly from one
locality to another (Ingolfsson 1967). A few
observations are available for arctic terns
<em>(Sterna paradisaea)</em> in Alaska, but little information
exists for other terns (Table 10). Bent
(1921) noted that Aleutian terns <em>(S. aleutica)</em>
sometimes associate with arctic terns during
feeding.</p>

<p>Finally, we must include raptors, particularly
the peregrine <em>(Falco peregrinus)</em> and bald
eagle <em>(Haliaeetus leucocephalus)</em>, because
they are important predators on the smaller
seabirds (White et al. 1971, 1973). Peregrines
have, in fact, been observed feeding on storm-petrels
far at sea (Craddock and Carlson
1970).</p>


<h4>Trophic Relations Within
Seabird Communities</h4>

<p>We have compared and summarized in
general terms the food partitioning by species
in five rather broad oceanographic regions
and their subdivisions in the northeastern
North Pacific and Bering Sea, based on the
specific details on diets presented in Tables 1
through 10. The five broad regions, defined
oceanographically by Dodimead et al. (1963)
and Favorite et al. (1976) and modified by
Sanger (1972), are shown in Fig. 1. The five
oceanographic regions (domains) were divided
further into inshore neritic, offshore neritic,
and oceanic habitats (Sanger and King, this
volume). We did not include estuarine habitats
or sheltered bays in the analysis.</p>

<div class="figcenter" style="width: 700px;">
<img src="images/p_101.jpg" width="700" height="376" alt="" />
<div class="caption"><p>Fig. 1. Schematic oceanographic domains of the
subarctic Pacific regions (defined by Dodimead et
al. (1963) and Favorite et al. (1976) and modified
by Sanger (1972).)</p></div>
</div>

<p><span class="pagenum"><a name="Page_102" id="Page_102">[Pg 102]</a></span></p>

<div class="center">
<table border="1" cellpadding="4" cellspacing="0" summary="Table 4. Food of murres and puffins in different localities">
    <caption>Table 4. <em>Food of murres and puffins in different localities (x = major prey, o = minor prey, and * = incidental prey species).</em></caption>
<col></col>
<col span="30" width="3%"></col>
<thead>
  <tr>
    <th rowspan="3">LOCATION</th>
    <th colspan="0">Diet</th>
  </tr>
  <tr>
    <th colspan="4">CRUS&shy;TA&shy;CEAN</th>
    <th rowspan="2" class="tdt">P<br />O<br />L<br />Y<br />C<br />H<br />A<br />E<br />T<br />E</th>
    <th rowspan="2" class="tdt">C<br />E<br />P<br />H<br />A<br />L<br />O<br />P<br />O<br />D</th>
    <th colspan="0">FISH</th>
  </tr>
  <tr>
    <th>E<br />u<br />p<br />h<br />a<br />u<br />s<br />i<br />i<br />d</th>
    <th>A<br />m<br />p<br />h<br />i<br />p<br />o<br />d</th>
    <th>I<br />s<br />o<br />p<br />o<br />d</th>
    <th>D<br />e<br />c<br />a<br />p<br />o<br />d</th>
    <th>C<br />l<br />u<br />p<br />e<br />a</th>
    <th>S<br />a<br />r<br />d<br />i<br />n<br />o<br />p<br />s</th>
    <th>E<br />n<br />g<br />r<br />a<br />u<br />l<br />i<br />s</th>
    <th>S<br />a<br />l<br />m<br />o</th>
    <th>O<br />n<br />c<br />h<br />o<br />r<br />h<br />y<br />n<br />c<br />h<br />u<br />s</th>
    <th>H<br />y<br />p<br />o<br />m<br />e<br />s<br />u<br />s</th>
    <th>T<br />h<br />a<br />l<br />e<br />i<br />c<br />h<br />t<br />h<br />y<br />s</th>
    <th>M<br />a<br />l<br />l<br />o<br />t<br />u<br />s</th>
    <th>B<br />o<br />r<br />e<br />o<br />g<br />a<br />d<br />u<br />s</th>
    <th>M<br />i<br />c<br />r<br />o<br />g<br />a<br />d<br />u<br />s</th>
    <th>T<br />h<br />e<br />r<br />a<br />g<br />r<br />a</th>
    <th>L<br />y<br />c<br />o<br />d<br />e<br />s</th>
    <th>G<br />a<br />s<br />t<br />e<br />r<br />o<br />s<br />t<br />e<br />u<br />s</th>
    <th>S<br />e<br />b<br />a<br />s<br />t<br />e</th>
    <th>T<br />r<br />i<br />g<br />l<br />o<br />p<br />s</th>
    <th>M<br />y<br />o<br />x<br />o<br />c<br />e<br />p<br />h<br />a<br />l<br />u<br />s</th>
    <th>C<br />o<br />t<br />t<br />i<br />d</th>
    <th>C<br />y<br />m<br />a<br />t<br />o<br />g<br />a<br />s<br />t<br />e<br />r</th>
    <th>E<br />m<br />b<br />i<br />o<br />t<br />o<br />c<br />i<br />d</th>
    <th>C<br />h<br />i<br />r<br />o<br />l<br />o<br />p<br />h<br />i<br />s</th>
    <th>S<br />t<br />i<br />c<br />h<br />a<br />e<br />i<br />d</th>
    <th>A<br />m<br />m<br />o<br />d<br />y<br />t<br />e<br />s</th>
    <th>P<br />l<br />e<br />u<br />r<br />o<br />n<br />e<br />c<br />t<br />i<br />d</th>
    <th>L<br />i<br />p<br />a<br />r<br />i<br />d</th>
  </tr>
</thead>
<tbody>
  <tr>
    <td colspan="0"><strong>Common murre</strong><em>(Uria aalge)</em><a name="FNanchor_20_20" id="FNanchor_20_20"></a><a href="#Footnote_20_20" class="fnanchor">[20]</a></td>
  </tr>
  <tr>
    <td class="tdp">Cape Thompson (Swartz 1966)</td>
    <td class="tdc">o</td>
    <td class="tdc">o</td>
    <td class="tdc">o</td>
    <td></td>
    <td class="tdc">o</td>
    <td class="tdc">o</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">o</td>
    <td class="tdc">o</td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">o</td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td class="tdp">Pribilof Islands (Preble and McAtee 1923)</td>
    <td></td>
    <td class="tdc">x</td>
    <td class="tdc">o</td>
    <td></td>
    <td class="tdc">o</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">o</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td class="tdp">E. Bering Sea (Ogi and Tsujita 1973)</td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">o</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td class="tdp">Forrester Island (Heath 1915)</td>
    <td class="tdc">o</td>
    <td class="tdc">o</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td class="tdp">Vancouver Island</td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">o</td>
    <td></td>
    <td class="tdc">o</td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">o</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td class="tdp">(Robertson, unpublished data)</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">o</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">o</td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td class="tdp">Olympic Peninsula (Cody 1973)</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td class="tdc">o</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">o</td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">o</td>
    <td class="tdc">o</td>
    <td></td>
  </tr>
  <tr><td class="tdp">Yaquina Head</td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">o</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr><td class="tdp">(Scott 1973)</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td class="tdc">o</td>
    <td></td>
    <td class="tdc">o</td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">*</td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">o</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">o</td>
    <td></td>
  </tr>
  <tr><td class="tdp">Farallon Islands (PRBO, unpublished data)</td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">o</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td><strong>Thick-billed murre</strong><br /><em>(U. lomvia)</em><a name="FNanchor_21_21" id="FNanchor_21_21"></a><a href="#Footnote_21_21" class="fnanchor">[21]</a></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td class="tdc">o</td>
    <td></td>
    <td></td>
    <td class="tdc">o</td>
    <td class="tdc">o</td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">o</td>
    <td class="tdc">o</td>
    <td class="tdc">x</td>
    <td class="tdc">o</td>
    <td></td>
  </tr>
  <tr>
    <td class="tdp">Cape Thompson (Swartz 1966)</td>
    <td></td>
    <td class="tdc">o</td>
    <td></td>
    <td class="tdc">o</td>
    <td class="tdc">o</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td class="tdp">Pribilof Islands (Preble and McAtee 1923)</td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td class="tdp">Hooker Island</td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">o</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td class="tdp">(Demme 1934, <em>in</em> Dement'ev et al. 1968)</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">o</td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">*</td>
    <td class="tdc">x</td>
    <td class="tdc">*</td>
    <td class="tdc">o</td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td class="tdc">o</td>
    <td class="tdc">o</td>
    <td class="tdc">o</td>
  </tr>
  <tr>
    <td class="tdp">NE Canada (Tuck and Squires 1937)</td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td class="tdc">o</td>
    <td class="tdc">o</td>
    <td class="tdc">o</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td><b>Tufted puffin</b></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td class="tdp"><em>(Lunda cirrhata)</em><a name="FNanchor_22_22" id="FNanchor_22_22"></a><a href="#Footnote_22_22" class="fnanchor">[22]</a></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">*</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td class="tdp"> Cape Thompson</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">*</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td class="tdp"> (Swartz 1966)</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td class="tdp"> Forrester Island (Heath 1915)</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td class="tdp"> Langara Island</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td class="tdp"> (Sealy 1973<em>a</em>)</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td class="tdp">Washington</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">o</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td class="tdp"> (Jewett et al. 1953)</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td class="tdc">o</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td class="tdp"> Olympic Peninsula</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td class="tdp"> (Cody 1973)</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td class="tdc">o</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td class="tdp"> Farallon Island (PRBO, unpublished data)</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td><b>Horned puffin</b></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">o</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td class="tdp"><em>(Fratercula corniculata)</em></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">o</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td class="tdp"> Cape Thompson (Swartz 1966)</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">o</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td class="tdp"> Alaska (Bent 1946)</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td class="tdp"> Forrester Island (Heath 1915)</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td ><b>Rhinoceros auklet</b> <em>(Cerorhinca monocerata)</em></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">o</td>
    <td class="tdc">x</td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td class="tdp"> NW Pacific (Kozlova 1961;</td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td class="tdp"> Komaki 1967)</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td class="tdp"> Forrester Island (Heath 1915)</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td class="tdp"> Langara Island (Sealy 1973<em>a</em>)</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">o</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td class="tdp"> Destruction Island</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">o</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td class="tdp"> (Richardson 1961)</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">o</td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td class="tdp"> Olympic Peninsula (Cody 1973)</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td class="tdp"> So. California (Linton 1908; Grinnell 1899)</td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
</tbody>
</table><span class="pagenum"><a name="Page_104" id="Page_104">[Pg 104]</a></span></div>

<div class="center">
<table border="0" cellpadding="4" cellspacing="0" summary="Table 5. Food of ancient and marbled murrelets">
    <caption>Table 5. <em>Food of ancient and marbled murrelets (x = major prey, o = minor prey, and * = incidental prey species).</em></caption>
  <tr>
    <th rowspan="3">Location</th>
    <th colspan="0">DIET</th>
  </tr>
  <tr>
    <th colspan="10">CRUSTACEAN</th>
    <th class="bl br">SQU&shy;ID</th>
    <th colspan="0">FISH</th>
  </tr>
  <tr>
    <th>Eu&shy;phau&shy;si&shy;id</th>
    <th>Thy&shy;sa&shy;noes&shy;sa</th>
    <th>Eu&shy;phau&shy;sia</th>
    <th>My&shy;sid</th>
    <th>A&shy;can&shy;tho&shy;my&shy;sis</th>
    <th>Am&shy;phi&shy;pod</th>
    <th>Gam&shy;ma&shy;rid</th>
    <th>Ca&shy;rid shrimp</th>
    <th>Dec&shy;a&shy;pod</th>
    <th>Lar&shy;vae</th>
    <th>Lar&shy;vae</th>
    <th>En&shy;gra&shy;u&shy;lis</th>
    <th>Os&shy;mer&shy;id</th>
    <th>Scor&shy;paen&shy;id</th>
    <th>Cy&shy;ma&shy;to&shy;ga&shy;ster</th>
    <th>Sti&shy;chae&shy;id</th>
    <th>Am&shy;mo&shy;dy&shy;tes</th>
    <th>Lar&shy;vae</th>
  </tr>
  <tr>
    <td colspan="0"><strong>Ancient murrelet</strong> <em>(Synthliboramphus antiquus)</em><a name="FNanchor_23_23" id="FNanchor_23_23"></a><a href="#Footnote_23_23" class="fnanchor">[23]</a></td>
  </tr>
  <tr>
    <td class="tdp"> Commander Islands (Dement'ev et al. 1968)</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc"> x</td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td class="tdp"> Amchitka Island (White et al. 1971, 1973)</td>
    <td class="tdc">x</td>
    <td class="tdc"> x</td>
    <td></td>
    <td class="tdc"> x</td>
    <td class="tdc"> x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc"> x</td>
  </tr>
  <tr>
    <td class="tdp"> Langara Island (Sealy 1975)</td>
    <td class="tdc">x</td>
    <td class="tdc"> x</td>
    <td class="tdc"> x</td>
    <td></td>
    <td></td>
    <td class="tdc"> *</td>
    <td></td>
    <td></td>
    <td class="tdc"> *</td>
    <td class="tdc"> *</td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc"> o</td>
    <td class="tdc"> o</td>
    <td></td>
    <td class="tdc"> x</td>
  </tr>
  <tr>
    <td colspan="0"><strong>Marbled murrelet</strong> <em>(Brachyramphus marmoratus)</em><a name="FNanchor_24_24" id="FNanchor_24_24"></a><a href="#Footnote_24_24" class="fnanchor">[24]</a></td>
  </tr>
  <tr>
    <td class="tdp"> SE Alaska (Grinnell 1897)</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc"> o</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td class="tdp"> Langara Island (Sealy 1975)</td>
    <td class="tdc">x</td>
    <td class="tdc"> x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc"> *</td>
    <td></td>
    <td></td>
    <td class="tdc"> *</td>
    <td></td>
    <td class="tdc"> *</td>
    <td class="tdc"> *</td>
    <td class="tdc"> x</td>
    <td class="tdc"> *</td>
    <td></td>
  </tr>
  <tr>
    <td class="tdp"> Vancouver Island (Munro and Clemens 1931)</td>
    <td></td>
    <td></td>
    <td class="tdc"> x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc"> x</td>
    <td></td>
    <td></td>
    <td class="tdc"> x</td>
  </tr>
  <tr>
    <td class="tdp"> Olympic Peninsula (Cody 1973)</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc"> x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc"> x</td>
  </tr>
</table><span class="pagenum"><a name="Page_105" id="Page_105">[Pg 105]</a></span></div>

<div class="center">
<table border="0" cellpadding="4" cellspacing="0" summary="Table 6. Diets of auklets in different localities">
    <caption>Table 6. <em>Diets of auklets in different localities (x = major prey, o = minor prey, and * = incidental prey species).</em></caption>
  <tr>
    <th rowspan="3">Location</th>
    <th colspan="0">Diet</th>
  </tr>
  <tr>
    <th colspan="11">CRUSTACEAN</th>
    <th rowspan="2" class="tdt">PO&shy;LY&shy;CHAE&shy;TE</th>
    <th>SQUID</th>
    <th colspan="0">FISH</th>
  </tr>
  <tr>
    <th>Eu&shy;pha&shy;u&shy;si&shy;id</th>
    <th>Thy&shy;sa&shy;noes&shy;sa</th>
    <th>My&shy;sid</th>
    <th>Sty&shy;lo&shy;my&shy;sis</th>
    <th>Am&shy;phi&shy;pod</th>
    <th>Pa&shy;ra&shy;the&shy;mi&shy;sto</th>
    <th>Phro&shy;ne&shy;ma</th>
    <th>Gam&shy;ma&shy;rid</th>
    <th>Co&shy;pe&shy;pod</th>
    <th>Ca&shy;la&shy;nus</th>
    <th>Ca&shy;rid shrimp</th>
    <th>Lar&shy;vae</th>
    <th>Cot&shy;tid</th>
    <th>Lar&shy;vae</th>
  </tr>
  <tr>
    <td colspan="0"><strong>Cassin's auklet</strong> <em>(Ptychoramphus aleuticus)</em></td>
  </tr>
  <tr>
    <td class="tdp"> Forrester Island (Heath 1915)</td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc"> x</td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc"> x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td class="tdp"> Olympic Peninsula (Cody 1973)</td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc"> x</td>
  </tr>
  <tr>
    <td class="tdp"> Farallon Islands (Manuwal 1974)</td>
    <td class="tdc">x</td>
    <td class="tdc"> x</td>
    <td></td>
    <td></td>
    <td class="tdc"> x</td>
    <td></td>
    <td class="tdc"> x</td>
    <td></td>
    <td class="tdc"> x</td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc"> o</td>
    <td></td>
    <td class="tdc"> x</td>
  </tr>
  <tr>
    <td colspan="0"><strong>Parakeet auklet</strong> <em>(Cyclorrhynchus psittaculus)</em></td>
  </tr>
  <tr>
    <td class="tdp"> Chukhotsk Peninsula (Portenko 1934, <em>in</em> Dement'ev et al. 1968)</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td class="tdc">x</td>
    <td></td>
    <td class="tdc"> x x</td>
    <td></td>
    <td class="tdc"> x</td>
  </tr>
  <tr>
    <td class="tdp"> Aleutian Islands (Bent 1946)</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc"> x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td class="tdp"> St. Lawrence Island (B&eacute;dard 1969<em>a</em>)</td>
    <td class="tdc"> x</td>
    <td class="tdc"> x</td>
    <td class="tdc"> o</td>
    <td></td>
    <td class="tdc">x </td>
    <td class="tdc">x</td>
    <td></td>
    <td class="tdc">o </td>
    <td class="tdc">o</td>
    <td class="tdc"> * </td>
    <td class="tdc">o </td>
    <td class="tdc">o </td>
    <td class="tdc">o</td>
  </tr>
  <tr>
    <td colspan="0"><strong>Crested auklet</strong> <em>(Aethia cristatella)</em><a name="FNanchor_25_25" id="FNanchor_25_25"></a><a href="#Footnote_25_25" class="fnanchor">[25]</a></td>
  </tr>
  <tr>
    <td class="tdp"> W. Bering Sea (Portenko 1934, <em>in</em> Dement'ev et al. 1968)</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc"> x</td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td class="tdp"> Commander Islands (Stejneger 1885)</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc"> x</td>
    <td></td>
  </tr>
  <tr>
    <td class="tdp"> Amchitka (White et al. 1973)</td>
    <td class="tdc">x</td>
    <td class="tdc"> x</td>
    <td class="tdc"> x</td>
    <td class="tdc"> x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc"> x</td>
    <td></td>
  </tr>
  <tr>
    <td class="tdp"> St. Lawrence Island. (B&eacute;dard 1969<em>a</em>)</td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc"> x</td>
    <td class="tdc"> x</td>
    <td class="tdc"> x</td>
    <td></td>
    <td class="tdc">o o</td>
    <td class="tdc"> x</td>
    <td class="tdc"> x</td>
    <td class="tdc">*</td>
  </tr>
  <tr>
    <td class="tdp"> Pribilof Islands (Preble and McAtee 1923)</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc"> x</td>
    <td class="tdc"> x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td colspan="0"><strong>Least auklet</strong> <em>(A. pusilla)</em></td>
  </tr>
  <tr>
    <td class="tdp"> Commander Islands (Stejneger 1885)</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc"> x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td class="tdp"> Aleutian Islands (Bent 1946)</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc"> x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td class="tdp"> St. Lawrence Island (B&eacute;dard 1969<em>a</em>)</td>
    <td></td>
    <td></td>
    <td class="tdc"> o</td>
    <td class="tdc"> o</td>
    <td class="tdc"> *</td>
    <td class="tdc"> o</td>
    <td class="tdc">o</td>
    <td class="tdc">o </td>
    <td class="tdc">x</td>
    <td class="tdc"> x </td>
    <td class="tdc">x</td>
  </tr>
  <tr>
    <td colspan="0"><strong>Whiskered auklet</strong> <em>(A. pygmaea)</em></td>
  </tr>
  <tr>
    <td class="tdp"> Commander Islands (Stejneger 1885)</td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td class="tdc"> x</td>
    <td></td>
    <td></td>
    <td class="tdc"> *</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
</table></div>
<div class="pagenum"><a name="Page_106" id="Page_106">[Pg 106]</a></div>
<div class="center">
<table border="0" cellpadding="4" cellspacing="0" summary="Table 7. Diets of loons and grebes in different localities">
    <caption>Table 7. <em>Diets of loons and grebes in different localities (x = major prey, o = minor prey, and * = incidental prey species).</em></caption>
  <tr>
    <th rowspan="3">Location</th>
    <th colspan="0">Diet</th>
  </tr>
  <tr>
    <th colspan="4">CRUSTACEANS</th>
    <th rowspan="2" class="tdt">PO&shy;LY&shy;CHAE&shy;TE</th>
    <th colspan="0">FISH</th>
  </tr>
  <tr>
    <th>Eu&shy;pha&shy;us&shy;id</th>
    <th>Am&shy;phi&shy;pod</th>
    <th>My&shy;sid</th>
    <th>De&shy;ca&shy;pod</th>
    <th>An&shy;gu&shy;il&shy;la</th>
    <th>Cla&shy;pea</th>
    <th>Sar&shy;di&shy;nops</th>
    <th>Sal&shy;mo</th>
    <th>Tha&shy;le&shy;ich&shy;thys</th>
    <th>A&shy;the&shy;ri&shy;nops</th>
    <th>Zo&shy;ar&shy;chid</th>
    <th>Ga&shy;did</th>
    <th>Fun&shy;du&shy;lus</th>
    <th>Ga&shy;ste&shy;ro&shy;ste&shy;us</th>
    <th>Se&shy;ba&shy;stes</th>
    <th>Cat&shy;tid</th>
    <th>Cy&shy;ma&shy;to&shy;ga&shy;ster</th>
    <th>Sti&shy;chae&shy;id</th>
    <th>Am&shy;mo&shy;dy&shy;tes</th>
    <th>Go&shy;bi&shy;id</th>
  </tr>
  <tr>
    <td colspan="0"><strong>Common loon</strong> <em>(Gavia immer)</em></td>
  </tr>
  <tr>
    <td class="tdp"> Alaska (Palmer 1962)</td>
    <td></td>
    <td class="tdc"> *</td>
    <td></td>
    <td class="tdc"> *</td>
    <td></td>
    <td></td>
    <td class="tdc"> o</td>
    <td></td>
    <td class="tdc"> o</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc"> o</td>
    <td class="tdc"> o</td>
    <td class="tdc"> o</td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td class="tdp"> Vancouver Island (Munro and Clemens 1931)</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc"> x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td class="tdp"> Denmark (Madsen 1957)</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc"> *</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc"> o</td>
    <td class="tdc"> x</td>
    <td></td>
    <td class="tdc"> *</td>
    <td></td>
    <td class="tdc"> o</td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td colspan="0"><strong>Yellow-billed loon</strong> <em>(G. adamsii)</em><a name="FNanchor_26_26" id="FNanchor_26_26"></a><a href="#Footnote_26_26" class="fnanchor">[26]</a></td>
  </tr>
  <tr>
    <td class="tdp"> Alaska (Cottam and Knappen 1939)</td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc"> *</td>
    <td class="tdc"> *</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc"> o</td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc"> x</td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td class="tdp"> Alaska (Bailey 1922)</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc"> x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td colspan="0"><strong>Arctic loon</strong> <em>(G. arctica)</em><a name="FNanchor_27_27" id="FNanchor_27_27"></a><a href="#Footnote_27_27" class="fnanchor">[27]</a></td>
  </tr>
  <tr>
    <td class="tdp"> Vancouver Island (Palmer 1962)</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc"> x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td class="tdp"> Vancouver Island (Robertson, unpublished data)</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc"> x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td class="tdp"> California (Palmer 1962)</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc"> x</td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td class="tdp"> Denmark (Madsen 1957)</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc"> *</td>
    <td class="tdc"> o</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc"> *</td>
    <td class="tdc"> x</td>
    <td></td>
    <td class="tdc"> x</td>
    <td></td>
    <td class="tdc"> *</td>
    <td></td>
    <td></td>
    <td class="tdc"> x</td>
    <td class="tdc"> *</td>
  </tr>
  <tr>
    <td colspan="0"><strong>Red-throated loon</strong> <em>(G. stellata)</em><a name="FNanchor_28_28" id="FNanchor_28_28"></a><a href="#Footnote_28_28" class="fnanchor">[28]</a></td>
  </tr>
  <tr>
    <td class="tdp"> Oregon (Palmer 1962)</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc"> x</td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td class="tdp"> No. Atlantic (Palmer 1962)</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc"> x</td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc"> x</td>
    <td></td>
    <td></td>
    <td class="tdc"> o</td>
  </tr>
  <tr>
    <td class="tdp"> Denmark (Madsen 1957)</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc"> *</td>
    <td class="tdc"> o</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc"> *</td>
    <td class="tdc"> x</td>
    <td></td>
    <td class="tdc"> o</td>
    <td></td>
    <td class="tdc"> *</td>
    <td></td>
    <td></td>
    <td class="tdc"> *</td>
    <td class="tdc"> o</td>
  </tr>
  <tr>
    <td colspan="0"><strong>Western grebe</strong> <em>(Aechmophorus occidentalis)</em></td>
  </tr>
  <tr>
    <td class="tdp"> Vancouver Island (Munro 1941)</td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc"> o</td>
    <td class="tdc"> *</td>
    <td></td>
    <td class="tdc"> x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc"> x</td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td class="tdp"> Vancouver Island (Robertson, unpublished data)</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc"> x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc"> x</td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td class="tdp"> Puget Sound (Phillips and Carter 1957)</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc"> x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc"> *</td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc"> o</td>
    <td class="tdc"> o</td>
    <td class="tdc"> o</td>
    <td></td>
  </tr>
  <tr>
    <td class="tdp"> Washington (Chatwin 1956)</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc"> *</td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc"> x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td class="tdp"> California (Palmer 1962)</td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc"> o</td>
    <td class="tdc"> *</td>
    <td></td>
    <td class="tdc"> x</td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc"> x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc"> x</td>
    <td></td>
    <td class="tdc"> *</td>
    <td></td>
  </tr>
  <tr>
    <td colspan="0"><strong>Red-necked grebe</strong> <em>(Podiceps grisegena)</em></td>
  </tr>
  <tr>
    <td class="tdp"> Pribilof Islands (Preble and McAtee 1923)</td>
    <td></td>
    <td class="tdc"> o</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td class="tdp"> Vancouver Island (Wetmore 1924)</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc"> x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td class="tdp"> Vancouver Island (Munro 1941)</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc"> x</td>
    <td class="tdc"> o</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc"> x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td class="tdp"> E. No. America (Wetmore 1924)</td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc"> o</td>
    <td class="tdc"> *</td>
    <td class="tdc"> o</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc"> o</td>
    <td></td>
    <td></td>
    <td class="tdc"> x</td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td colspan="0"><strong>Horned grebe</strong> <em>(P. auritus)</em><a name="FNanchor_29_29" id="FNanchor_29_29"></a><a href="#Footnote_29_29" class="fnanchor">[29]</a></td>
  </tr>
  <tr>
    <td class="tdp"> Pribilof Islands (Preble and McAtee 1923)</td>
    <td></td>
    <td class="tdc"> x</td>
    <td></td>
    <td></td>
    <td class="tdc"> o</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td class="tdp"> W. No. America (Wetmore 1924)</td>
    <td></td>
    <td class="tdc"> x</td>
    <td class="tdc"> x</td>
    <td></td>
    <td class="tdc"> *</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc"> o</td>
    <td></td>
    <td class="tdc"> o</td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td class="tdp"> Vancouver Island (Munro 1941)</td>
    <td class="tdc"> x</td>
    <td></td>
    <td></td>
    <td class="tdc"> x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc"> o</td>
    <td class="tdc"> o</td>
    <td class="tdc"> *</td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td class="tdp"> Denmark (Madsen 1957)</td>
    <td></td>
    <td class="tdc"> o</td>
    <td class="tdc"> o</td>
    <td></td>
    <td class="tdc"> o</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc"> o</td>
  </tr>
  <tr>
    <td colspan="0"><strong>Eared grebe</strong> <em>(P. nigricollis)</em><a name="FNanchor_30_30" id="FNanchor_30_30"></a><a href="#Footnote_30_30" class="fnanchor">[30]</a></td>
  </tr>
  <tr>
    <td class="tdp"> W. No. America (Wetmore 1924)</td>
    <td></td>
    <td class="tdc"> *</td>
    <td class="tdc"> x</td>
    <td></td>
    <td class="tdc"> *</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc"> o</td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td class="tdp"> Vancouver Island (Munro 1941)</td>
    <td></td>
    <td class="tdc"> x</td>
    <td class="tdc"> x</td>
    <td></td>
    <td></td>
    <td class="tdc"> o</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td class="tdp"> Denmark (Madsen 1957)</td>
    <td></td>
    <td></td>
    <td class="tdc"> x</td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc"> *</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc"> o</td>
  </tr>
</table>
<span class="pagenum"><a name="Page_107" id="Page_107">[Pg 107]</a></span></div>

<div class="center">
<table border="0" cellpadding="4" cellspacing="0" summary="Table 8. Diets of albatrosses and petrels in different localities">
    <caption>Table 8. <em>Diets of albatrosses and petrels in different localities (x = major prey, o = minor prey, and * = incidental prey species).</em></caption>
  <tr>
    <th rowspan="3">Location</th>
    <th colspan="0">Diet</th>
  </tr>
  <tr>
    <th colspan="6">CRUSTACEAN</th>
    <th rowspan="2" class="tdt">COE&shy;LEN&shy;TE&shy;RA&shy;TE</th>
    <th rowspan="2" class="tdt">E&shy;CHI&shy;NO&shy;DERM</th>
    <th rowspan="2" class="tdt">CE&shy;PHA&shy;LO&shy;POD</th>
    <th colspan="0">FISH</th>
  </tr>
  <tr>
    <th>Eu&shy;pha&shy;u&shy;si&shy;id</th>
    <th>Am&shy;phi&shy;pod</th>
    <th>Co&shy;pe&shy;pod</th>
    <th>De&shy;ca&shy;pod</th>
    <th>Lar&shy;vae</th>
    <th>Bar&shy;na&shy;cle</th>
    <th>"Fish"</th>
    <th><em>En&shy;grau&shy;lis</em></th>
    <th>My&shy;cto&shy;phid</th>
    <th><em>Se&shy;ba&shy;stes</em></th>
    <th><em>Am&shy;mo&shy;dy&shy;tes</em></th>
    <th>Car&shy;ri&shy;on, fish of&shy;fal</th>
    <th>Fish eggs</th>
  </tr>
  <tr>
    <td colspan="0"><strong>Black-footed albatross</strong> <em>(Diomedea nigripes)</em></td>
  </tr>
  <tr>
    <td class="tdp">No. Pacific (Palmer 1962)</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
  </tr>
  <tr>
    <td class="tdp">Aleutian Islands (Cottam and Knappen 1939)</td>
    <td></td>
    <td class="tdc">o</td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">o</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td class="tdc">x</td>
  </tr>
  <tr>
    <td class="tdp">California (Miller 1936, 1940)</td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">o</td>
    <td></td>
    <td class="tdc">o</td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td class="tdc">x</td>
  </tr>
  <tr>
    <td colspan="0"><strong>Laysan albatross</strong> <em>(D. immutabilis)</em></td>
  </tr>
  <tr>
    <td class="tdp">No. Pacific (Palmer 1962; Bartsch 1922; Fisher 1904)</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td colspan="0"><strong>Northern fulmar</strong> <em>(Fulmarus glacialis)</em></td>
  </tr>
  <tr>
    <td class="tdp">Pribilof Islands (Preble and McAtee 1923)</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">o</td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td class="tdp">Alaska (Gabrielson and Lincoln 1959)</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
  </tr>
  <tr>
    <td class="tdp">Oregon (Gabrielson and Jewett 1940)</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td class="tdp">No. Atlantic (Hartley and Fisher 1936; Einarsson 1945; Fisher 1952)</td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
  </tr>
  <tr>
    <td colspan="0"><strong>Flesh-footed shearwater</strong> <em>(Puffinus carneipes)</em></td>
  </tr>
  <tr>
    <td class="tdp">Australia (Oliver 1955; Serventy et al. 1971)</td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td colspan="0"><strong>Pink-footed shearwater</strong> <em>(P. creatopus)</em></td>
  </tr>
  <tr>
    <td class="tdp">California (Murphy 1936; Ainley, personal observation)</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td class="tdp">E. Pacific (Cottam and Knappen 1939)</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td colspan="0"><strong>Buller's shearwater</strong> <em>(P. bulleri)</em></td>
  </tr>
  <tr>
    <td class="tdp">SW Pacific (Falla 1934; Serventy et al. 1971)</td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td class="tdp">Peru (Murphy 1936)</td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td colspan="0"><strong>Sooty shearwater</strong> <em>(P. griseus)</em></td>
  </tr>
  <tr>
    <td class="tdp">Aleutian Islands (Sanger, personal observation)</td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td class="tdp">British Columbia (Martin 1942; Sealy 1973<em>a</em>)</td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
  </tr>
  <tr>
    <td class="tdp">Oregon (Gabrielson and Jewett 1940)</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td class="tdp">California (Ainley, personal observation)</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td class="tdp">Peru (Murphy 1936)</td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td class="tdp">SW Pacific (Oliver 1955; Serventy et al. 1971)</td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td colspan="0"><strong>Short-tailed shearwater</strong> <em>(P. tenuirostris)</em></td>
  </tr>
  <tr>
    <td class="tdp">Bristol Bay (Bartonek, personal communication)</td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td class="tdp">Alaska (Cottam and Knappen 1939)</td>
    <td class="tdc">x</td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td class="tdc">o</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td class="tdp">No. Pacific (Palmer 1962; Kuroda 1955)</td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td class="tdp">Australia (Serventy et al. 1971)</td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td class="tdp">Bass Strait (Sheard 1953)</td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td><strong>Mottled petrel</strong> <em>(Pterodroma inexpectata)</em></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td class="tdp">Pacific Ocean (Imber 1973)</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td class="tdp">E. No. Pacific (Kuroda 1955)</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
  </tr>
  <tr>
    <td colspan="0"><strong>Fork-tailed storm-petrel</strong> <em>(Oceanodroma furcata)</em></td>
  </tr>
  <tr>
    <td class="tdp">Pribilof Islands (Preble and McAtee 1923)</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td class="tdp">SE Alaska (Heath 1915)</td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td class="tdp">British Columbia (Martin 1942)</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
  </tr>
  <tr>
    <td class="tdp">California (Ainley, personal observation)</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td colspan="0"><strong>Leach's storm-petrel</strong> <em>(O. leucorhoa)</em></td>
  </tr>
  <tr>
    <td class="tdp">SE Alaska (Heath 1915)</td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td class="tdp">California (PRBO, unpublished data)</td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
  </tr>
  <tr>
    <td class="tdp">So. California (Palmer 1962)</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
  </tr>
  <tr>
    <td class="tdp">No. Atlantic<a name="FNanchor_31_31" id="FNanchor_31_31"></a><a href="#Footnote_31_31" class="fnanchor">[31]</a> (Palmer 1962)</td>
    <td class="tdc">x</td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
  </tr>
</table><span class="pagenum"><a name="Page_108" id="Page_108">[Pg 108]</a></span></div>


<div class="center">
<table border="0" cellpadding="4" cellspacing="0" summary="Table 9. Diets of gulls in different localities">
    <caption>Table 9. <em>Diets of gulls in different localities (x = major prey, o = minor prey, and * = incidental prey species).</em></caption>
  <tr>
    <th rowspan="3">Location</th>
    <th colspan="0">Diet</th>
  </tr>
  <tr>
    <th colspan="3">CRUSTACEAN</th>
    <th rowspan="2" class="tdt">P<br />O<br />L<br />Y<br />C<br />H<br />A<br />E<br />T<br />E</th>
    <th colspan="2">MOLLUSC</th>
    <th rowspan="2" class="tdt">E<br />C<br />H<br />I<br />N<br />O<br />D<br />E<br />R<br />M</th>
    <th rowspan="2" class="tdt">C<br />O<br />E<br />L<br />E<br />N<br />T<br />E<br />R<br />A<br />T<br />E</th>
    <th colspan="16">FISH</th>
    <th rowspan="2" class="tdt">C<br />A<br />R<br />R<br />I<br />O<br />N<br />-<br />O<br />F<br />F<br />A<br />L</th>
    <th colspan="3">BIRD</th>
    <th rowspan="2" class="tdt">F<br />I<br />S<br />H<br /> <br />E<br />G<br />G<br />S</th>
  </tr>
  <tr>
    <th>E<br />u<br />p<br />h<br />a<br />u<br />s<br />i<br />i<br />d<br /></th>
    <th>B<br />a<br />r<br />n<br />a<br />c<br />l<br />e<br /></th>
    <th>D<br />e<br />c<br />a<br />p<br />o<br />d<br /></th>
    <th>S<br />h<br />e<br />l<br />l<br /> f<br />i<br />s<br />h<br /></th>
    <th>C<br />e<br />p<br />h<br />a<br />l<br />o<br />p<br />o<br />d<br /></th>
    <th>"<br />F<br />i<br />s<br />h<br />"</th>
    <th><em>C<br />l<br />u<br />p<br />e<br />a<br /></em></th>
    <th><em>E<br />n<br />g<br />r<br />a<br />u<br />l<br />i<br />s<br /></em></th>
    <th><em>O<br />s<br />m<br />e<br />r<br />u<br />s<br /></em></th>
    <th><em>P<br />o<br />r<br />i<br />c<br />h<br />i<br />h<br />y<br />s<br /></em></th>
    <th><em>O<br />t<br />a<br />p<br />h<br />i<br />d<br />i<br />u<br />m<br /></em></th>
    <th><em>M<br />a<br />l<br />l<br />o<br />t<br />u<br />s<br /></em></th>
    <th><em>B<br />o<br />r<br />r<br />o<br />g<br />a<br />d<br />u<br />s<br /></em></th>
    <th><em>M<br />i<br />c<br />r<br />o<br />g<br />a<br />d<br />u<br />s<br /></em></th>
    <th><em>G<br />a<br />d<br />u<br />s<br /></em></th>
    <th><em>L<br />y<br />c<br />o<br />d<br />e<br />s<br /></em></th>
    <th><em>S<br />e<br />b<br />a<br />s<br />t<br />e<br />s<br /></em></th>
    <th><em>M<br />y<br />x<br />o<br />c<br />e<br />p<br />h<br />a<br />l<br />u<br />s<br /></em></th>
    <th><em>G<br />e<br />n<br />y<br />o<br />n<br />e<br />m<br />u<br />s<br /></em></th>
    <th>E<br />m<br />b<br />i<br />o<br />t<br />o<br />c<br />i<br />d<br /></th>
    <th><em>A<br />m<br />m<br />o<br />d<br />y<br />t<br />e<br />s<br /></em></th>
    <th>E<br />g<br />g<br />s<br /></th>
    <th>C<br />h<br />i<br />c<br />k<br />s<br /></th>
    <th>A<br />d<br />u<br />l<br />t<br />s</th>
  </tr>
  <tr>
    <td colspan="0"><strong>Glaucous gull</strong> <em>(Larus hyperboreus)</em></td>
  </tr>
  <tr>
    <td class="tdp">St. Lawrence Island (Fay and Cade 1959)</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td class="tdc">x</td>
    <td class="tdc">x</td>
  </tr>
  <tr>
    <td class="tdp">Chukchi Sea (Swartz 1966)</td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td class="tdc">x</td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td class="tdp">Pribilof Islands (Preble and McAtee 1923)</td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td class="tdc">x</td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td class="tdp">Vancouver Island (Munro and Clemens 1931)</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
  </tr>
  <tr>
    <td colspan="0"><strong>Glaucous-winged gull</strong> <em>(L. glaucescens)</em><a name="FNanchor_32_32" id="FNanchor_32_32"></a><a href="#Footnote_32_32" class="fnanchor">[32]</a></td>
  </tr>
  <tr>
    <td class="tdp">Pribilof Islands (Preble and McAtee 1923)</td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td class="tdc">x</td>
    <td class="tdc">x</td>
    <td class="tdc">x</td>
  </tr>
  <tr>
    <td class="tdp">Alaska (Bent 1921)</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td class="tdp">No. Pacific (Sanger 1973)</td>
    <td></td>
    <td class="tdc">x</td>
    <td class="tdc">*</td>
    <td></td>
    <td class="tdc">*</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td class="tdp">Mandarte Island (Ward 1973)</td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td class="tdp">Vancouver Island (Munro and Clemens 1931; Robertson, unpublished data)</td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td colspan="0"><strong>Western gull</strong> <em>(L. occidentalis)</em><a name="FNanchor_33_33" id="FNanchor_33_33"></a><a href="#Footnote_33_33" class="fnanchor">[33]</a></td>
  </tr>
  <tr>
    <td class="tdp">Farallon Islands (PRBO, unpublished data)</td>
    <td class="tdc">x</td>
    <td class="tdc">x</td>
    <td class="tdc">o</td>
    <td></td>
    <td class="tdc">*</td>
    <td class="tdc">x</td>
    <td class="tdc">*</td>
    <td class="tdc">*</td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td class="tdc">x</td>
    <td class="tdc">o</td>
    <td></td>
    <td></td>
    <td class="tdc">o</td>
    <td class="tdc">o</td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td class="tdc">o</td>
    <td class="tdc">o</td>
    <td></td>
    <td class="tdc">x</td>
    <td class="tdc">o</td>
    <td class="tdc">o</td>
    <td class="tdc">o</td>
  </tr>
  <tr>
    <td colspan="0"><strong>Herring gull</strong> <em>(L. argentatus)</em></td>
  </tr>
  <tr>
    <td class="tdp">No. Atlantic (Zelikman 1961)</td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td class="tdp">E. No. America (Bent 1946; Ainley, personal observation)</td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td class="tdc">x</td>
    <td class="tdc">x</td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td class="tdc">*</td>
    <td></td>
    <td class="tdc">o</td>
    <td class="tdc">x</td>
  </tr>
  <tr>
    <td class="tdp">Vancouver Island (Munro and Clemens 1931)</td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td colspan="0"><strong>Mew gull</strong> <em>(L. canus)</em></td>
  </tr>
  <tr>
    <td class="tdp">Alaska (Bent 1921)</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
  </tr>
  <tr>
    <td class="tdp">Vancouver Island (Munro and Clemens 1931)</td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
  </tr>
  <tr>
    <td colspan="0"><strong>Heermann's gull</strong> <em>(L. heermanni)</em></td>
  </tr>
  <tr>
    <td class="tdp">California (Bent 1921)</td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td colspan="0"><strong>Bonaparte's gull</strong> <em>(L. philadelphia)</em></td>
  </tr>
  <tr>
    <td class="tdp">E. No. America (Bent 1921)</td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td colspan="0"><strong>Black-legged kittiwake</strong> <em>(Rissa tridactyla)</em><a name="FNanchor_34_34" id="FNanchor_34_34"></a><a href="#Footnote_34_34" class="fnanchor">[34]</a></td>
  </tr>
  <tr>
    <td class="tdp">Chukchi Sea (Swartz 1966)</td>
    <td></td>
    <td></td>
    <td class="tdc">o</td>
    <td class="tdc">*</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">o</td>
    <td></td>
    <td></td>
    <td class="tdc">o</td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td class="tdc">o</td>
    <td></td>
    <td class="tdc">o</td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td class="tdp">Pribilof Islands (Preble and McAtee 1923)</td>
    <td></td>
    <td></td>
    <td class="tdc">o</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">o</td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td class="tdp">Alaska (Bent 1921)</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">o</td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td class="tdp">Cook Inlet<a name="FNanchor_35_35" id="FNanchor_35_35"></a><a href="#Footnote_35_35" class="fnanchor">[35]</a> (Snarski, personal communication)</td>
    <td></td>
    <td></td>
    <td class="tdc">o</td>
    <td class="tdc">o</td>
    <td class="tdc">o</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td class="tdc">o</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td class="tdp">No. Atlantic (Hartley and Fisher 1936; Zelikman 1961)</td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td colspan="0"><strong>Red-legged kittiwake</strong> <em>(R. breuirostris)</em></td>
  </tr>
  <tr>
    <td class="tdp">Pribilof Islands (Preble and McAtee 1923)</td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td colspan="0"><strong>Sabine's gull</strong> <em>(Xenia sabini)</em></td>
  </tr>
  <tr>
    <td class="tdp">Pt. Barrow (Banner 1954)</td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
</table></div>

<div class="pagenum"><a name="Page_109" id="Page_109">[Pg 109]</a></div>


<div class="center">
<table border="0" cellpadding="4" cellspacing="0" summary="Table 10. Diets of terns in different localities">
    <caption>Table 10. <em>Diets of terns in different localities (x = major prey species).</em></caption>
  <tr>
    <th rowspan="3">Location</th>
    <th colspan="0">Diet</th>
  </tr>
  <tr>
    <th colspan="3">CRUSTACEAN</th>
    <th colspan="0">FISH</th>
  </tr>
  <tr>
    <th>"Crustacean"</th>
    <th><em>Mallotus</em></th>
    <th>Euphausiid</th>
    <th>Cottid</th>
    <th>Amphipod</th>
    <th><em>Ammodytes</em><br />larvae</th>
  </tr>
  <tr>
    <td colspan="0"><strong>Arctic tern</strong> <em>(Sterna paradisaea)</em></td>
  </tr>
  <tr>
    <td class="tdp">Pribilof Islands (Preble and McAtee 1923)</td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td class="tdc">x</td>
  </tr>
  <tr>
    <td class="tdp">Alaska (Bent 1921)</td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td class="tdc">x</td>
  </tr>
  <tr>
    <td class="tdp">No. Atlantic (Hartley and Fisher 1936)</td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td colspan="0"><strong>Common tern</strong> <em>(S. hirundo)</em></td>
  </tr>
  <tr>
    <td class="tdp">E. No. America (Bent 1921)</td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
  </tr>
</table></div>

<p>The oceanic habitat includes waters of the
photic zone overlying the deep ocean and continental
slopes beyond the continental or
insular shelves. The Bering Sea and central
subarctic domains are largely made up of
oceanic habitat. The other three domains include
both inshore and offshore neritic as well
as some oceanic habitat. The boundary between
the inshore and offshore neritic has yet
to be defined in terms of bird life, but it lies at
that line beyond which the bottom is too deep
for a diving bird to exploit. A depth contour
thus defines the boundary. In the antarctic
South Pacific, emperor penguins <em>(Aptenodytes
fosteri)</em> dive to depths of 275 m, but so
far as is known, no comparable bird exists in
the North Pacific. Some marine ducks and
loons reportedly dive to 50-60 m (Kooyman
1974). The inshore-offshore neritic boundary
for seabirds may lie near the 70-m depth
contour.</p>

<p>Food resource partitioning by seabirds in
the five oceanographic domains are shown in
Tables 11-15. Within each domain, the common
and usual members of the seabird community
are listed, and the major and minor
categories in each of their diets are shown (on
the basis of available literature, Tables 1-10).
The categories are grouped further, and
rather tenuously, according to the trophic
level at which a bird is presumably feeding: I
= herbivore, II = secondary carnivore, III =
tertiary carnivore, IV = final carnivore, and
Sc = scavengers (carnivorous) feeding at
many levels. Birds at level I feed on large
algae and seed plants and are not directly part
of the same food webs involving other species.
These food webs originate with phytoplankton
(Fig. 2). So far as is known, no bird feeds
on phytoplankton and few, if any, feed on
microzooplankton; hence birds do not
generally feed as primary carnivores. An exception
at times might be the least auklet
<em>(Aethia pusilla)</em> when it feeds on small copepods
(see B&eacute;dard 1969<em>b</em>).</p>

<p>The above groupings are "tenuous" because
prey in each category may represent more
than one trophic level, and a single prey
species could occur at one level one day or
place and at another level the next day or
place, depending upon what it happened to be
eating. This is shown in Fig. 2, where the
parakeet auklet <em>(Cyclorrhynchus psittaculus)</em>
can occur in the food web at different levels,
depending both on the prey it is eating and on
what its prey is eating. Even without this
complication, many seabirds feed at more
than one level in the food web. For instance,
murres eating euphausiids would be feeding
at a different level than murres feeding on
larger fish. It might be "safer" to regard prey
organisms in level II as macrozooplankton,
prey organisms in level III as micronekton,
and prey organisms (seabirds themselves) in
level IV as macronekton (after Sverdrup et al.
1942).</p>

<div class="figcenter" style="width: 700px;">
<img src="images/p_109.jpg" width="700" height="467" alt="" />
<div class="caption"><p>Fig. 2. Schematic food web of the parakeet auklet in
the eastern Bering Sea (based on B&eacute;dard 1969<em>a</em>
and Dunbar 1946). Arrow sizes indicate relative
importance of prey and Roman numerals refer to
prey sizes (see text).</p></div>
</div>

<p><span class="pagenum"><a name="Page_110" id="Page_110">[Pg 110]</a></span></p>

<div class="center">
<table border="0" cellpadding="4" cellspacing="0" summary="Table 11. Use of food resources by seabirds in the Bering Sea coastal domain.">
  <caption>Table 11. <em>Use of food resources by seabirds in the Bering
  Sea coastal domain.</em> Information is from Tables 1-10. (Trophic
  level I = plants, II = secondary carnivore, III = tertiary
  carnivore, IV = upper level carnivore [on birds only in this
  table], Sc = scavenger on carrion, offal, or detritus [II-IV]; x
  = major food in diet, o = minor food, * = incidental food, ? =
  probable food.)</caption>
  <tr>
    <th rowspan="4">Seabirds</th>
    <th colspan="0">Habitat, bird trophic levels (I-IV. Sc), and food categories</th>
  </tr>
  <tr>
    <th colspan="9">Oceanic and offshore neritic</th>
    <th colspan="0">Inshore neritic</th>
  </tr>
  <tr>
    <th colspan="4" class="br">II</th>
    <th colspan="2" class="br"> III</th>
    <th colspan="1"> IV</th>
    <th colspan="1"> Sc</th>
    <th colspan="1"> I</th>
    <th colspan="6" class="br">II</th>
    <th colspan="3" class="br"> III</th>
    <th colspan="1"> IV</th>
    <th colspan="1"> Sc</th>
  </tr>
  <tr>
    <th>C<br />r<br />u<br />s<br />t<br />a<br />c<br />e<br />a<br />n </th>
    <th>P<br />o<br />l<br />y<br />c<br />h<br />a<br />e<br />t<br />e </th>
    <th>C<br />o<br />e<br />l<br />e<br />n<br />t<br />e<br />r<br />a<br />t<br />e </th>
    <th>F<br />i<br />s<br />h<br />/<br />s<br />q<br />u<br />i<br />d<br /> <br />e<br />g<br />g<br />s<br /> <br />&amp;<br /> <br />l<br />a<br />r<br />v<br />a<br />e </th>

    <th>F<br />i<br />s<br />h </th>
    <th>C<br />e<br />p<br />h<br />a<br />l<br />o<br />p<br />o<br />d</th>
    <th>B<br />i<br />r<br />d<br />s</th>

    <th>C<br />a<br />r<br />r<br />i<br />o<br />n<br />/<br />o<br />f<br />f<br />a<br />l<br />/<br />d<br />e<br />t<br />r<br />i<br />t<br />u<br />s</th>

    <th>P<br />l<br />a<br />n<br />t</th>

    <th>C<br />r<br />u<br />s<br />t<br />a<br />c<br />e<br />a<br />n<br />,<br /> <br />m<br />i<br />d<br />w<br />a<br />t<br />e<br />r </th>
    <th>C<br />r<br />u<br />s<br />t<br />a<br />c<br />e<br />a<br />n<br />,<br /> <br />b<br />e<br />n<br />t<br />h<br />i<br />c </th>
    <th>C<br />o<br />e<br />l<br />e<br />n<br />t<br />e<br />r<br />a<br />t<br />e </th>
    <th>E<br />c<br />h<br />i<br />n<br />o<br />d<br />e<br />r<br />m </th>
    <th>M<br />o<br />l<br />l<br />u<br />s<br />c </th>
    <th>F<br />i<br />s<br />h<br />/<br />s<br />q<br />u<br />i<br />d<br /> <br />e<br />g<br />g<br />s<br /> <br />&amp;<br /> <br />l<br />a<br />r<br />v<br />a<br />e </th>

    <th>F<br />i<br />s<br />h<br />,<br /> <br />m<br />i<br />d<br />w<br />a<br />t<br />e<br />r</th>
    <th>F<br />i<br />s<br />h<br />,<br /> <br />b<br />e<br />n<br />t<br />h<br />i<br />c</th>
    <th>C<br />e<br />p<br />h<br />a<br />l<br />o<br />p<br />o<br />d</th>

    <th>B<br />i<br />r<br />d<br />s</th>

    <th>C<br />a<br />r<br />r<br />i<br />o<br />n<br />/<br />o<br />f<br />f<br />a<br />l<br />/<br />d<br />e<br />t<br />r<br />i<br />t<br />u<br />s</th>
  </tr>
  <tr>
    <td><em>Gavia adamsii</em></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc"> *</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">o</td>
    <td class="tdc"> x</td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td><em>G. arctica</em></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">o</td>
    <td class="tdc"> x</td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td><em>Podiceps grisegena</em></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc"> o</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">o</td>
    <td class="tdc"> x</td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td><em>Diomedea nigripes</em></td>
    <td class="tdc"> x</td>
    <td class="tdc"> o</td>
    <td class="tdc"> o</td>
    <td class="tdc">o</td>
    <td class="tdc"> x</td>
    <td class="tdc"> x</td>
    <td></td>
    <td class="tdc"> x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td><em>Fulmarus glacialis</em></td>
    <td class="tdc"> x</td>
    <td class="tdc"> o</td>
    <td class="tdc"> x</td>
    <td class="tdc">o</td>
    <td class="tdc"> x</td>
    <td class="tdc"> x</td>
    <td></td>
    <td class="tdc"> x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td><em>Puffinus griseus</em></td>
    <td class="tdc"> x</td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc"> x</td>
    <td class="tdc"> x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">o</td>
    <td></td>
    <td class="tdc"> o</td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td><em>P. tenuirostris</em></td>
    <td class="tdc"> x</td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc"> o</td>
    <td class="tdc"> x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">o</td>
    <td></td>
    <td class="tdc"> o</td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td><em>Oceanodroma furcata</em></td>
    <td class="tdc"> x</td>
    <td class="tdc"> o</td>
    <td class="tdc"> o</td>
    <td class="tdc">x</td>
    <td class="tdc"> x</td>
    <td class="tdc"> x</td>
    <td></td>
    <td class="tdc"> x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td><em>Phalacrocorax auritus</em></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc"> o</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td class="tdc"> o</td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td><em>P. pelagicus</em></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc"> x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc"> x</td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td><em>P. urile</em></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc"> x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc"> x</td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td><em>Branta bernicla</em></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc"> x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td><em>Philacte canagica</em></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc"> x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td><em>Clangula hyemalis</em></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc"> o</td>
    <td class="tdc"> x</td>
    <td></td>
    <td></td>
    <td class="tdc">o</td>
    <td></td>
    <td></td>
    <td class="tdc"> o</td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td><em>Histrionicus histrionicus</em></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc"> o</td>
    <td class="tdc"> x</td>
    <td></td>
    <td></td>
    <td class="tdc">o</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td><em>Polysticta stelleri</em></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc"> o</td>
    <td class="tdc"> x</td>
    <td class="tdc"> o</td>
    <td></td>
    <td></td>
    <td class="tdc">o</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td><em>Samateria mollissima</em></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc"> x</td>
    <td></td>
    <td class="tdc">o</td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td><em>S. spectabilis</em></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc"> o</td>
    <td></td>
    <td class="tdc"> o</td>
    <td></td>
    <td class="tdc">o</td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td><em>S. fischeri</em></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc"> x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td><em>Melanitta deglandi</em></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td class="tdc"> o</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td><em>M. nigra</em></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc"> o</td>
    <td></td>
    <td class="tdc"> o</td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td><em>Haliaeetus leucocephalus</em></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td class="tdc"> x</td>
  </tr>
  <tr>
    <td><em>Falco peregrinus</em></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
  </tr>
  <tr>
    <td><em>Phalaropus fulicarius</em></td>
    <td class="tdc"> x</td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc"> o</td>
    <td></td>
    <td class="tdc"> x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc"> x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td><em>Lobipes lobatus</em></td>
    <td class="tdc"> x</td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc"> o</td>
    <td></td>
    <td class="tdc"> x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc"> x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td><em>Stercorarius</em> spp.</td>
    <td class="tdc"> o</td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc"> x</td>
    <td class="tdc"> x</td>
    <td class="tdc">?</td>
    <td class="tdc"> x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td class="tdc"> x</td>
  </tr>
  <tr>
    <td><em>Larus hyperboreus</em></td>
    <td class="tdc"> o</td>
    <td></td>
    <td></td>
    <td class="tdc">o</td>
    <td class="tdc"> o</td>
    <td class="tdc"> o</td>
    <td></td>
    <td></td>
    <td class="tdc"> o</td>
    <td></td>
    <td class="tdc"> o</td>
    <td class="tdc">o</td>
    <td class="tdc">o</td>
    <td class="tdc">o</td>
    <td class="tdc"> o</td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td class="tdc"> x</td>
  </tr>
  <tr>
    <td><em>L. glaucescens</em></td>
    <td class="tdc"> o</td>
    <td></td>
    <td></td>
    <td class="tdc">o</td>
    <td class="tdc"> o</td>
    <td class="tdc"> o</td>
    <td></td>
    <td></td>
    <td class="tdc"> o</td>
    <td></td>
    <td class="tdc"> o</td>
    <td class="tdc">o</td>
    <td class="tdc">o</td>
    <td class="tdc">o</td>
    <td class="tdc"> o</td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td class="tdc"> x</td>
  </tr>
  <tr>
    <td><em>L. argentatus</em></td>
    <td class="tdc"> o</td>
    <td></td>
    <td></td>
    <td class="tdc">o</td>
    <td class="tdc"> o</td>
    <td class="tdc"> o</td>
    <td></td>
    <td></td>
    <td class="tdc"> o</td>
    <td></td>
    <td class="tdc"> o</td>
    <td class="tdc">o</td>
    <td class="tdc">o</td>
    <td class="tdc">o</td>
    <td class="tdc"> o</td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td class="tdc">o</td>
    <td class="tdc"> x</td>
  </tr>
  <tr>
    <td><em>L. canus</em></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc"> x</td>
    <td class="tdc">o</td>
    <td class="tdc">o</td>
    <td class="tdc">o</td>
    <td class="tdc"> x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc"> x</td>
  </tr>
  <tr>
    <td><em>Rissa tridactyla</em></td>
    <td class="tdc"> x</td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc"> x</td>
    <td class="tdc"> x</td>
    <td></td>
    <td></td>
    <td class="tdc"> o</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td><em>Xema sabini</em></td>
    <td class="tdc"> x</td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td class="tdc"> o</td>
    <td class="tdc"> o</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td><em>Sterna paradisaea</em></td>
    <td class="tdc"> x</td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td class="tdc"> o</td>
    <td class="tdc"> o</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td><em>Uria aalge</em></td>
    <td class="tdc"> x</td>
    <td class="tdc"> o</td>
    <td></td>
    <td></td>
    <td class="tdc"> x</td>
    <td class="tdc"> x</td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc"> o</td>
    <td class="tdc"> o</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td class="tdc"> o</td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td><em>U. lomvia</em></td>
    <td class="tdc"> x</td>
    <td class="tdc"> o</td>
    <td></td>
    <td></td>
    <td class="tdc"> x</td>
    <td class="tdc"> x</td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc"> o</td>
    <td class="tdc"> x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">o</td>
    <td class="tdc"> x</td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td><em>Lunda cirrhata</em></td>
    <td class="tdc"> ?</td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc"> x</td>
    <td class="tdc"> x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td><em>Fratercula corniculata</em></td>
    <td></td>
    <td class="tdc"> *</td>
    <td></td>
    <td></td>
    <td class="tdc"> x</td>
    <td class="tdc"> x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td><em>Cepphus columba</em></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc"> o</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc"> x</td>
    <td class="tdc"> o</td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td><em>Synthliboramphus antiquus</em></td>
    <td class="tdc"> x</td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc"> x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc"> o</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td><em>Brachyramphus brevirostris</em></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc"> x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">o</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td><em>Cyclorrhynchus psittaculus</em></td>
    <td class="tdc"> x</td>
    <td class="tdc"> o</td>
    <td></td>
    <td class="tdc">*</td>
    <td class="tdc"> *</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td><em>Aethia cristatella</em></td>
    <td class="tdc"> x</td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td><em>A. pusilla</em></td>
    <td class="tdc"> x</td>
    <td></td>
    <td></td>
    <td class="tdc">o</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
</table><span class="pagenum"><a name="Page_111" id="Page_111">[Pg 111]</a></span></div>


<div class="center">
<table border="0" cellpadding="4" cellspacing="0" summary="Table 12. Use of food resources by seabirds in the oceanic and offshore neritic habitats, Bering Sea domain.">
  <caption>Table 12. <em>Use of food resources by seabirds in the oceanic
  and offshore neritic habitats, Bering Sea domain.</em> Information
  is from Tables 1-10. (Trophic level I = plants, II = secondary
  carnivore, III = tertiary carnivore, IV = upper level carnivore;
  Sc = scavenger on carrion, offal, or detritus [II-IV]; x = major
  food in diet, o = minor food, * = incidental food, ? = probable
  food.)</caption>
  <tr>
    <th rowspan="3">Seabirds</th>
    <th colspan="0">Bird trophic levels and food categories</th>
  </tr>
  <tr>
    <th colspan="4" class="br">II</th>
    <th colspan="2" class="br">III </th>
    <th colspan="1">IV </th>
    <th colspan="1">Sc</th>
  </tr>
  <tr>
    <th class="tdp">Cru&shy;sta&shy;ce&shy;an</th>
    <th class="tdp">Po&shy;ly&shy;chae&shy;te</th>
    <th class="tdp">Coe&shy;len&shy;te&shy;ra&shy;te</th>
    <th class="tdp">Fish/Squid eggs &amp; Lar&shy;vae</th>
    <th class="tdp">Fish</th>
    <th class="tdp">Ce&shy;pha&shy;lo&shy;pod</th>
    <th class="tdp">Birds</th>
    <th class="tdp">Car&shy;ri&shy;on/of&shy;fal/de&shy;tri&shy;tus</th>
  </tr>
  <tr>
    <td><em>Diomedea nigripes</em></td>
    <td class="tdc">x</td>
    <td class="tdc"> o</td>
    <td class="tdc"> o</td>
    <td class="tdc"> o</td>
    <td class="tdc"> x</td>
    <td class="tdc"> x</td>
    <td></td>
    <td class="tdc">x</td>
  </tr>
  <tr>
    <td><em>D. immutabilis</em></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc"> x</td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td><em>Fulmarus glacialis</em></td>
    <td class="tdc">x</td>
    <td class="tdc"> o</td>
    <td class="tdc"> x</td>
    <td class="tdc"> o</td>
    <td class="tdc"> x</td>
    <td class="tdc"> x</td>
    <td></td>
    <td class="tdc">x</td>
  </tr>
  <tr>
    <td><em>Puffinus griseus</em></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc"> x</td>
    <td class="tdc"> x</td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td><em>P. tenuirostris</em></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc"> o</td>
    <td class="tdc"> x</td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td><em>Pterodroma inexpectata</em></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc"> x</td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
  </tr>
  <tr>
    <td><em>Oceanodroma furcata</em></td>
    <td class="tdc">x</td>
    <td class="tdc"> o</td>
    <td class="tdc"> o</td>
    <td class="tdc"> x</td>
    <td class="tdc"> x</td>
    <td class="tdc"> x</td>
    <td></td>
    <td class="tdc">x</td>
  </tr>
  <tr>
    <td><em>Phalaropus fulicarius</em></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td class="tdc"> x</td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">o</td>
  </tr>
  <tr>
    <td><em>Lobipes lobatus</em></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td class="tdc"> x</td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">o</td>
  </tr>
  <tr>
    <td><em>Stercorarius</em> spp.</td>
    <td class="tdc">o</td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc"> x</td>
    <td class="tdc"> x</td>
    <td class="tdc"> ?</td>
    <td class="tdc">x</td>
  </tr>
  <tr>
    <td><em>Larus hyperboreus</em></td>
    <td class="tdc">x</td>
    <td class="tdc"> o</td>
    <td class="tdc"> o</td>
    <td class="tdc"> o</td>
    <td class="tdc"> x</td>
    <td class="tdc"> x</td>
    <td class="tdc"> ?</td>
    <td class="tdc">x</td>
  </tr>
  <tr>
    <td><em>L. glaucescens</em></td>
    <td class="tdc">x</td>
    <td class="tdc"> o</td>
    <td class="tdc"> o</td>
    <td class="tdc"> o</td>
    <td class="tdc"> x</td>
    <td class="tdc"> x</td>
    <td class="tdc"> ?</td>
    <td class="tdc">x</td>
  </tr>
  <tr>
    <td><em>Rissa tridactyla</em></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc"> x</td>
    <td class="tdc"> x</td>
    <td></td>
    <td class="tdc">o</td>
  </tr>
  <tr>
    <td><em>R. brevirostris</em></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc"> x</td>
    <td class="tdc"> x</td>
    <td></td>
    <td class="tdc">o</td>
  </tr>
  <tr>
    <td><em>Xema sabini</em></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td class="tdc"> x</td>
    <td class="tdc"> o</td>
    <td class="tdc"> o</td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td><em>Sterna paradisaea</em></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td class="tdc"> x</td>
    <td class="tdc"> o</td>
    <td class="tdc"> o</td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td><em>Uria aalge</em></td>
    <td class="tdc">x</td>
    <td class="tdc"> o</td>
    <td></td>
    <td></td>
    <td class="tdc"> x</td>
    <td class="tdc"> x</td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td><em>U. lomvia</em></td>
    <td class="tdc">x</td>
    <td class="tdc"> o</td>
    <td></td>
    <td></td>
    <td class="tdc"> x</td>
    <td class="tdc"> x</td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td><em>Lunda cirrhata</em></td>
    <td class="tdc">?</td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc"> x</td>
    <td class="tdc"> x</td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td><em>Fratercula corniculata</em></td>
    <td></td>
    <td class="tdc"> *</td>
    <td></td>
    <td></td>
    <td class="tdc"> x</td>
    <td class="tdc"> x</td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td><em>Synthliboramphus antiquus</em></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc"> x</td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td><em>Cyclorrhynchus psittaculus</em></td>
    <td class="tdc">x</td>
    <td class="tdc"> o</td>
    <td></td>
    <td class="tdc"> *</td>
    <td class="tdc"> *</td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td><em>Aethia cristatella</em></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td class="tdc"> x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td><em>A. pusilla</em></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td class="tdc"> o</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td><em>A. pygmaea</em></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
</table></div>

<p>Information contained in Tables 11-15 can
be summarized to show characteristics of seabird
trophic relations. One such characteristic
is the range of diet breadth or diet complexity
(Table 16). Few species (about 6%) feed on
only one type of prey and might, therefore, be
referred to as "specialists." Included are
eared grebe <em>(Podiceps caspicus)</em>, Laysan albatross,
brown pelican, emperor goose <em>(Philacte
canagica)</em>, black brant <em>(Bernicia bernicla)</em>,
peregrine falcon, and whiskered auklet
<em>(Aethia pygmaea)</em>. Consideration of these
species as specialists may require revision
when more data become available. Except for
the albatross and auklet, these species are
members of the inshore neritic cohort. Food
specialization does not seem to be characteristic
of oceanic birds in particular or of most
seabirds in general.</p>

<p><span class="pagenum"><a name="Page_112" id="Page_112">[Pg 112]</a></span></p>


<div class="center">
<table border="0" cellpadding="4" cellspacing="0" summary="Table 13. Use of food resources by seabirds in the Alaska Stream domain.">
  <caption>Table 13. <em>Use of food resources by seabirds in the
  Alaska Stream domain.</em> Information is from Tables 1-10.
  (Trophic level I = plants, II = secondary carnivore, III = tertiary
  carnivore, IV = upper level carnivore, Sc = scavenger on carrion,
  offal, or detritus [II-IV]; x = major food in diet, o = minor food,
  * = incidental food, ? = probable food.)</caption>
  <tr>
    <th rowspan="4">Seabirds</th>
    <th colspan="0">Habitat, bird trophic levels (I-IV. Sc), and food categories</th>
  </tr>
  <tr>
    <th colspan="9" class="br">Oceanic and offshore neritic</th>
    <th colspan="0">Inshore neritic Inshore neritic</th>
  </tr>
  <tr>
    <th colspan="4" class="br">II</th>
    <th colspan="2" class="br">III</th>
    <th colspan="1">IV</th>
    <th colspan="1">Sc</th>
    <th colspan="1">I</th>
    <th colspan="6" class="br">II</th>
    <th colspan="3" class="br">III</th>
    <th colspan="1">IV</th>
    <th colspan="1">Sc</th>
  </tr>
  <tr>
    <th>C<br />r<br />u<br />s<br />t<br />a<br />c<br />e<br />a<br />n </th>
    <th>P<br />o<br />l<br />y<br />c<br />h<br />a<br />e<br />t<br />e </th>
    <th>C<br />o<br />e<br />l<br />e<br />n<br />t<br />e<br />r<br />a<br />t<br />e </th>
    <th>F<br />i<br />s<br />h<br />/<br />s<br />q<br />u<br />i<br />d<br /> <br />e<br />g<br />g<br />s<br /> <br />&amp;<br /> <br />l<br />a<br />r<br />v<br />a<br />e </th>
    <th>F<br />i<br />s<br />h </th>
    <th>C<br />e<br />p<br />h<br />a<br />l<br />o<br />p<br />o<br />d</th>
    <th>B<br />i<br />r<br />d<br />s</th>
    <th>C<br />a<br />r<br />r<br />i<br />o<br />n<br />/<br />o<br />f<br />f<br />a<br />l<br />/<br />d<br />e<br />t<br />r<br />i<br />t<br />u<br />s</th>
    <th>P<br />l<br />a<br />n<br />t</th>
    <th>C<br />r<br />u<br />s<br />t<br />a<br />c<br />e<br />a<br />n<br />,<br /> <br />m<br />i<br />d<br />w<br />a<br />t<br />e<br />r </th>
    <th>C<br />r<br />u<br />s<br />t<br />a<br />c<br />e<br />a<br />n<br />,<br /> <br />b<br />e<br />n<br />t<br />h<br />i<br />c </th>
    <th>C<br />o<br />e<br />l<br />e<br />n<br />t<br />e<br />r<br />a<br />t<br />e </th>
    <th>E<br />c<br />h<br />i<br />n<br />o<br />d<br />e<br />r<br />m </th>
    <th>M<br />o<br />l<br />l<br />u<br />s<br />c</th>
    <th>F<br />i<br />s<br />h<br />,<br /> <br />m<br />i<br />d<br />w<br />a<br />t<br />e<br />r</th>
    <th>F<br />i<br />s<br />h<br />,<br /> <br />b<br />e<br />n<br />t<br />h<br />i<br />c</th>
    <th>C<br />e<br />p<br />h<br />a<br />l<br />o<br />p<br />o<br />d</th>
    <th>B<br />i<br />r<br />d<br />s</th>
    <th>F<br />i<br />s<br />h<br />/<br />s<br />q<br />u<br />i<br />d<br /> <br />e<br />g<br />g<br />s<br /> <br />&amp;<br /> <br />l<br />a<br />r<br />v<br />a<br />e </th>
    <th>C<br />a<br />r<br />r<br />i<br />o<br />n<br />/<br />o<br />f<br />f<br />a<br />l<br />/<br />d<br />e<br />t<br />r<br />i<br />t<br />u<br />s</th>
  </tr>
  <tr>
    <td><em>Gavia immer</em></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">*</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td><em>G. adamsii</em></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">*</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">o</td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td><em>G. stellata</em></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">*</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">o</td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td><em>Podiceps grisegena</em></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">o</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td class="tdc">o</td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td><em>Diomedea nigripes</em></td>
    <td class="tdc"> x</td>
    <td class="tdc">o</td>
    <td class="tdc">o</td>
    <td class="tdc">o</td>
    <td class="tdc"> x</td>
    <td class="tdc"> x</td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td><em>Fulmarus glacialis</em></td>
    <td class="tdc"> x</td>
    <td class="tdc">o</td>
    <td class="tdc">x</td>
    <td class="tdc">o</td>
    <td class="tdc"> x</td>
    <td class="tdc"> x</td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td><em>Puffinus griseus</em></td>
    <td class="tdc"> x</td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc"> x</td>
    <td class="tdc"> x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">o</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td><em>P. tenuirostris</em></td>
    <td class="tdc"> x</td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc"> o</td>
    <td class="tdc"> x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">o</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td><em>Pterodroma inexpectata</em></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc"> x</td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td><em>Oceanodroma furcata</em></td>
    <td class="tdc"> x</td>
    <td class="tdc">o</td>
    <td class="tdc">o</td>
    <td class="tdc">x</td>
    <td class="tdc"> x</td>
    <td class="tdc"> x</td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td><em>Phalacrocorax auritus</em></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">o</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td class="tdc">o</td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td><em>P. pelagicus</em></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td><em>P. urile</em></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td><em>Philacte canagica</em></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc"> x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td><em>Clangula hyemalis</em></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc"> o</td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td class="tdc">o</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td><em>Histrionicus histrionicus</em></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc"> x</td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td class="tdc">o</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td><em>Polysticta stelleri</em></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc"> o</td>
    <td class="tdc"> x</td>
    <td class="tdc">o</td>
    <td></td>
    <td></td>
    <td class="tdc">o</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td><em>Somateria mollissima</em></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td class="tdc">o</td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td><em>S. spectabilis</em></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">o</td>
    <td></td>
    <td class="tdc">o</td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td><em>S. fischeri</em></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc"> x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td><em>Melanitta deglandi</em></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td class="tdc">o</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td><em>M. perspicillata</em></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc"> o</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td class="tdc">o</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td><em>M. nigra</em></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc"> o</td>
    <td></td>
    <td class="tdc">o</td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td><em>Mergus serrator</em></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">o</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td><em>Haliaeetus leucocephalus</em></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td class="tdc">x</td>
  </tr>
  <tr>
    <td><em>Falco peregrinus</em></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
  </tr>
  <tr>
    <td><em>Phalaropus fulicarius</em></td>
    <td class="tdc"> x</td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">o</td>
    <td class="tdc"> x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td><em>Lobipes lobatus</em></td>
    <td class="tdc"> x</td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">o</td>
    <td class="tdc"> x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td><em>Stercorarius</em> spp.</td>
    <td class="tdc"> o</td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc"> x</td>
    <td class="tdc"> x</td>
    <td class="tdc"> ?</td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td class="tdc">x</td>
  </tr>
  <tr>
    <td><em>Larus hyperboreus</em></td>
    <td class="tdc"> o</td>
    <td></td>
    <td></td>
    <td class="tdc">o</td>
    <td class="tdc"> o</td>
    <td class="tdc"> o</td>
    <td></td>
    <td class="tdc">o</td>
    <td></td>
    <td></td>
    <td class="tdc">o</td>
    <td class="tdc">o</td>
    <td class="tdc">o</td>
    <td class="tdc">o</td>
    <td class="tdc">o</td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td class="tdc">x</td>
  </tr>
  <tr>
    <td><em>L. glaucescens</em></td>
    <td class="tdc"> o</td>
    <td></td>
    <td></td>
    <td class="tdc">o</td>
    <td class="tdc"> o</td>
    <td class="tdc"> o</td>
    <td></td>
    <td class="tdc">o</td>
    <td></td>
    <td></td>
    <td class="tdc">o</td>
    <td class="tdc">o</td>
    <td class="tdc">o</td>
    <td class="tdc">o</td>
    <td class="tdc">o</td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td class="tdc">x</td>
  </tr>
  <tr>
    <td><em>L. argentatus</em></td>
    <td class="tdc"> o</td>
    <td></td>
    <td></td>
    <td class="tdc">o</td>
    <td class="tdc"> o</td>
    <td class="tdc"> o</td>
    <td></td>
    <td class="tdc">o</td>
    <td></td>
    <td></td>
    <td class="tdc">o</td>
    <td class="tdc">o</td>
    <td class="tdc">o</td>
    <td class="tdc">o</td>
    <td class="tdc">o</td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td class="tdc">o</td>
    <td class="tdc">x</td>
  </tr>
  <tr>
    <td><em>L. canus</em></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td class="tdc">o</td>
    <td class="tdc">o</td>
    <td class="tdc">o</td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
  </tr>
  <tr>
    <td><em>Rissa tridactyla</em></td>
    <td class="tdc"> x</td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc"> x</td>
    <td class="tdc"> x</td>
    <td></td>
    <td class="tdc">o</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td><em>R. brevirostris</em></td>
    <td class="tdc"> x</td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc"> x</td>
    <td class="tdc"> x</td>
    <td></td>
    <td class="tdc">o</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td><em>Sterna paradisaea</em></td>
    <td class="tdc"> x</td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td class="tdc"> o</td>
    <td class="tdc"> o</td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc"> x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">o</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td><em>S. aleutica</em></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc"> x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">o</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td><em>Uria aalge</em></td>
    <td class="tdc"> x</td>
    <td class="tdc">o</td>
    <td></td>
    <td></td>
    <td class="tdc"> x</td>
    <td class="tdc"> x</td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc"> o</td>
    <td class="tdc">o</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td class="tdc">o</td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td><em>U. lomvia</em></td>
    <td class="tdc"> x</td>
    <td class="tdc">o</td>
    <td></td>
    <td></td>
    <td class="tdc"> x</td>
    <td class="tdc"> x</td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc"> o</td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">o</td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td><em>Lunda cirrhata</em></td>
    <td class="tdc"> ?</td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc"> x</td>
    <td class="tdc"> x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td><em>Fratercula corniculata</em></td>
    <td></td>
    <td class="tdc">*</td>
    <td></td>
    <td></td>
    <td class="tdc"> x</td>
    <td class="tdc"> x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td><em>Cepphus columba</em></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">o</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td class="tdc">o</td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td><em>Brachyramphus marmoratus</em></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc"> x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">o</td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td><em>B. brevirostris</em></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc"> x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">o</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td><em>Synthliboramphus antiquus</em></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc"> x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc"> x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td><em>Cyclorrhynchus psittaculus</em></td>
    <td class="tdc">x </td>
    <td class="tdc">o</td>
    <td></td>
    <td class="tdc">*</td>
    <td class="tdc"> *</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td><em>Aethia cristatella</em></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td><em>A. pusilla</em></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td class="tdc">o</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td><em>A. pygmaea</em></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
</table><span class="pagenum"><a name="Page_113" id="Page_113">[Pg 113]</a></span></div>


<div class="center">
<table border="0" cellpadding="4" cellspacing="0" summary="Table 14. Use of food resources by seabirds in the oceanic habitat, central subarctic domain.">
  <caption>Table 14. <em>Use of food resources by seabirds in the
  oceanic habitat, central subarctic domain.</em> Information is from
  Tables 1-10. (Trophic level I = plants, II = secondary carnivore,
  III = tertiary carnivore, IV = upper level carnivore, Sc =
  scavenger on carrion, offal, or detritus [II-IV]; x = major food in
  diet, o = minor food, * = incidental food, ? = probable food.)</caption>
  <tr>
    <th rowspan="3">Seabirds</th>
    <th colspan="0">Bird trophic levels and food categories</th>
  </tr>
  <tr>
    <th colspan="4" class="br">II</th>
    <th colspan="2" class="br">III</th>
    <th colspan="1">IV</th>
    <th colspan="0">Sc</th>
  </tr>
  <tr>
    <th>Crustacean</th>
    <th>Polychaete</th>
    <th>Coelenterate</th>
    <th>Fish/squid eggs &amp; larvae</th>
    <th>Fish</th>
    <th>Cephalopod</th>
    <th>Birds</th>
    <th>Carrion/offal/detritus</th>
  </tr>
  <tr>
    <td><em>Diomedea nigripes</em></td>
    <td class="tdc">x</td>
    <td class="tdc">o</td>
    <td class="tdc">o</td>
    <td class="tdc">o</td>
    <td class="tdc">x</td>
    <td class="tdc">x</td>
    <td></td>
    <td class="tdc">x</td>
  </tr>
  <tr>
    <td><em>D. immutabilis</em></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
  </tr>
  <tr>
    <td><em>Fulmarus glacialis</em></td>
    <td class="tdc">x</td>
    <td class="tdc">o</td>
    <td class="tdc">x</td>
    <td class="tdc">o</td>
    <td class="tdc">x</td>
    <td class="tdc">x</td>
    <td></td>
    <td class="tdc">x</td>
  </tr>
  <tr>
    <td><em>Puffinus carneipes</em></td>
    <td class="tdc">o</td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td class="tdc">x</td>
    <td></td>
  </tr>
  <tr>
    <td><em>P. griseus</em></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td class="tdc">x</td>
    <td></td>
  </tr>
  <tr>
    <td><em>P. tenuirostris</em></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">o</td>
    <td class="tdc">x</td>
    <td></td>
  </tr>
  <tr>
    <td><em>Pterodroma inexpectata</em></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
  </tr>
  <tr>
    <td><em>Oceanodroma furcata</em></td>
    <td class="tdc">x</td>
    <td class="tdc">o</td>
    <td class="tdc">o</td>
    <td class="tdc">o</td>
    <td class="tdc">x</td>
    <td class="tdc">x</td>
    <td></td>
    <td class="tdc">x</td>
  </tr>
  <tr>
    <td><em>O. leucorhoa</em></td>
    <td class="tdc">x</td>
    <td class="tdc">o</td>
    <td class="tdc">o</td>
    <td class="tdc">o</td>
    <td class="tdc">x</td>
    <td class="tdc">x</td>
    <td></td>
    <td class="tdc">x</td>
  </tr>
  <tr>
    <td><em>Phalaropus fulicarius</em></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">o</td>
  </tr>
  <tr>
    <td><em>Lobipes lobatus</em></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">o</td>
  </tr>
  <tr>
    <td><em>Stercorarius</em> spp.</td>
    <td class="tdc">o</td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td class="tdc">x</td>
    <td class="tdc">?</td>
    <td class="tdc">x</td>
  </tr>
  <tr>
    <td><em>Larus hyperboreus</em></td>
    <td class="tdc">x</td>
    <td class="tdc">o</td>
    <td class="tdc">o</td>
    <td class="tdc">o</td>
    <td class="tdc">x</td>
    <td class="tdc">x</td>
    <td class="tdc">?</td>
    <td class="tdc">x</td>
  </tr>
  <tr>
    <td><em>L. glaucescens</em></td>
    <td class="tdc">x</td>
    <td class="tdc">o</td>
    <td class="tdc">o</td>
    <td class="tdc">o</td>
    <td class="tdc">x</td>
    <td class="tdc">x</td>
    <td class="tdc">?</td>
    <td class="tdc">x</td>
  </tr>
  <tr>
    <td><em>L. argentatus</em></td>
    <td class="tdc">x</td>
    <td class="tdc">o</td>
    <td class="tdc">o</td>
    <td class="tdc">o</td>
    <td class="tdc">x</td>
    <td class="tdc">x</td>
    <td></td>
    <td class="tdc">x</td>
  </tr>
  <tr>
    <td><em>Rissa tridactyla</em></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td class="tdc">x</td>
    <td></td>
    <td class="tdc">o</td>
  </tr>
  <tr>
    <td><em>Xema sabini</em></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td class="tdc">o</td>
    <td class="tdc">o</td>
    <td></td>
  </tr>
  <tr>
    <td><em>Sterna paradisaea</em></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td class="tdc">o</td>
    <td class="tdc">o</td>
    <td></td>
  </tr>
  <tr>
    <td><em>Uria aalge</em></td>
    <td class="tdc">x</td>
    <td class="tdc">*</td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td class="tdc">x</td>
    <td></td>
  </tr>
  <tr>
    <td><em>U. lomvia</em></td>
    <td class="tdc">x</td>
    <td class="tdc">*</td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td class="tdc">x</td>
    <td></td>
  </tr>
  <tr>
    <td><em>Lunda cirrhata</em></td>
    <td class="tdc">o</td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td class="tdc">x</td>
    <td></td>
  </tr>
  <tr>
    <td><em>Fratercula corniculata</em></td>
    <td></td>
    <td class="tdc">*</td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td class="tdc">x</td>
    <td></td>
  </tr>
  <tr>
    <td><em>Cerorhinca monocerata</em></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td><em>Synthliboramphus antiquus</em></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td><em>Cyclorrhynchus psittaculus</em></td>
    <td class="tdc">x</td>
    <td class="tdc">o</td>
    <td></td>
    <td class="tdc">*</td>
    <td class="tdc">*</td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td><em>Ptychoramphus aleuticus</em></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td class="tdc">o</td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
</table></div>

<p>Most species (roughly 53% in any community)
include two or three prey categories in
their diets&mdash;usually midwater schooling fish,
squid, and crustaceans. These birds include
the most numerous in the communities&mdash;the
shearwaters and some alcids&mdash;which feed
largely on three prey types, and also include
some of the less abundant birds, the marine
ducks, which feed mostly on two prey
categories.</p>

<p>The remaining seabirds are more general in
their feeding. Many have large populations,
but are not as abundant as shearwaters or
most alcids. The true "generalists" are the
species that feed on as many as eight or more
types of prey, and relatively few (12%) such
species exist in each avian community. These
birds, the scavengers, include black-footed
albatross, fulmar, storm-petrels, and large
gulls. The petrels are the scavengers of the
oceanic habitat and the gulls are their counterparts
in the neritic habitat (but see Sanger
1973).</p>

<p>Another comparison is shown in Table 17,
where the species in each community are categorized
according to the number feeding at
each trophic level. If a species feeds at more
than one level, it is tallied once in each level.
Most seabirds (66-77%) feed at the second and
third levels as secondary and tertiary carnivores.
Few feed as terminal carnivores, and
relatively few are scavengers. Actually, most
scavenging occurs at levels II and III, so
about 90% of the seabirds in each community
feed at levels II and III. Communities including
an inshore neritic feeding element are the
only ones that include herbivores, and even
then, few of these species exist in significant
numbers in the marine environment (discounting
estuaries and sheltered bays).</p>

<p><span class="pagenum"><a name="Page_114" id="Page_114">[Pg 114]</a></span></p>


<div class="center">
<table border="0" cellpadding="4" cellspacing="0" summary="Table 15. Use of food resources by seabirds in the North American coastal domain.">
  <caption>Table 15. <em>Use of food resources by seabirds in the
  North American coastal domain.</em> Information is from Tables
  1-10. (Trophic level I = plants, II = secondary carnivore, III =
  tertiary carnivore, IV = upper level carnivore, Sc = scavenger on
  carrion, offal, or detritus [II-IV]; x = major food in diet, o =
  minor food, * = incidental food, ? = probable food.)</caption>
  <tr>
    <th rowspan="4">Seabirds</th>
    <th colspan="0">Habitat, bird trophic levels (I-IV, Sc), and food categories</th>
  </tr>
  <tr>
    <th colspan="9">Oceanic and offshore neritic</th>
    <th colspan="0">Inshore neritic</th>
  </tr>
  <tr>
    <th colspan="4">II</th>
    <th colspan="2">III </th>
    <th colspan="1">IV </th>
    <th colspan="1">Sc </th>
    <th colspan="1">I </th>
    <th colspan="6">II </th>
    <th colspan="3">III </th>
    <th colspan="1">IV </th>
    <th colspan="0">Sc</th>
  </tr>
  <tr>
    <th>C<br />r<br />u<br />s<br />t<br />a<br />c<br />e<br />a<br />n</th>
    <th>P<br />o<br />l<br />y<br />c<br />h<br />a<br />e<br />t<br />e</th>
    <th>C<br />o<br />e<br />l<br />e<br />n<br />t<br />e<br />r<br />a<br />t<br />e</th>
    <th>F<br />i<br />s<br />h<br />-<br />s<br />q<br />u<br />i<br />d<br /> <br />e<br />g<br />g<br />s<br /> <br />&amp;<br /> <br />l<br />a<br />r<br />v<br />a<br />e</th>
    <th>F<br />i<br />s<br />h</th>
    <th>C<br />e<br />p<br />h<br />a<br />l<br />o<br />p<br />o<br />d</th>
    <th>B<br />i<br />r<br />d<br />s</th>
    <th>C<br />a<br />r<br />r<br />i<br />o<br />n<br />/<br />o<br />f<br />f<br />a<br />l<br />/<br />d<br />e<br />t<br />r<br />i<br />t<br />u<br />s</th>
    <th>P<br />l<br />a<br />n<br />t</th>
    <th>C<br />r<br />u<br />s<br />t<br />a<br />c<br />e<br />a<br />n<br />,<br /> <br />m<br />i<br />d<br />w<br />a<br />t<br />e<br />r</th>
    <th>C<br />r<br />u<br />s<br />t<br />a<br />c<br />e<br />a<br />n<br />,<br /> <br />b<br />e<br />n<br />t<br />h<br />i<br />c</th>
    <th>C<br />o<br />e<br />l<br />e<br />n<br />t<br />e<br />r<br />a<br />t<br />e</th>
    <th>E<br />c<br />h<br />i<br />n<br />o<br />d<br />e<br />r<br />m</th>
    <th>M<br />o<br />l<br />l<br />u<br />s<br />c</th>
    <th>F<br />i<br />s<br />h<br />/<br />s<br />q<br />u<br />i<br />d<br /> <br />e<br />g<br />g<br />s<br /> <br />&amp;<br /> <br />l<br />a<br />r<br />v<br />a<br />e</th>
    <th>F<br />i<br />s<br />h<br />,<br /> <br />m<br />i<br />d<br />w<br />a<br />t<br />e<br />r</th>
    <th>F<br />i<br />s<br />h<br />,<br /> <br />b<br />e<br />n<br />t<br />h<br />i<br />c</th>
    <th>C<br />e<br />p<br />h<br />a<br />l<br />o<br />p<br />o<br />d</th>
    <th>B<br />i<br />r<br />d<br />s</th>
    <th>C<br />a<br />r<br />r<br />i<br />o<br />n<br />/<br />o<br />f<br />f<br />a<br />l<br />/<br />d<br />e<br />t<br />r<br />i<br />t<br />u<br />s</th>
  </tr>
  <tr>
    <td><em>Gavia immer</em></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">*</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td><em>G. adamsii</em></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">*</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">o</td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td><em>G. arctica</em></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">o</td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td><em>G. stellata</em></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">o</td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td><em>Podiceps grisegena</em></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">o</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td class="tdc">o</td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td><em>P. nigricollis</em></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">o</td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td><em>P. auritus</em></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">o</td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td><em>Aechmophorus occidentalis</em></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">o</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td><em>Diomedea nigripes</em></td>
    <td class="tdc">x</td>
    <td class="tdc">o</td>
    <td class="tdc">o</td>
    <td class="tdc">o</td>
    <td class="tdc">x</td>
    <td class="tdc">x</td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td><em>Fulmarus glacialis</em></td>
    <td class="tdc">x</td>
    <td class="tdc">o</td>
    <td class="tdc">x</td>
    <td class="tdc">o</td>
    <td class="tdc">x</td>
    <td class="tdc">x</td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td><em>Puffinus creatopus</em></td>
    <td class="tdc">o</td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td><em>P. carneipes</em></td>
    <td class="tdc">o</td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td><em>P. bulleri</em></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td><em>P. griseus</em></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">o</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">o</td>
    <td class="tdc">o</td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td><em>P. tenuirostris</em></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">o</td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">o</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">o</td>
    <td class="tdc">o</td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td><em>Oceanodroma furcata</em></td>
    <td class="tdc">x</td>
    <td class="tdc">o</td>
    <td class="tdc">o</td>
    <td class="tdc">o</td>
    <td class="tdc">x</td>
    <td class="tdc">x</td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td><em>Pelecanus occidentalis</em></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td><em>Phalacrocorax auritus</em></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">o</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td class="tdc">o</td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td><em>P. penicillatus</em></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">o</td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td><em>P. pelagicus</em></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td><em>Branta bernicla</em></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td><em>Clangula hyemalis</em></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">o</td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td class="tdc">o</td>
    <td></td>
    <td></td>
    <td class="tdc">o</td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td><em>Histrionicus histrionicus</em></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">o</td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td class="tdc">o</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td><em>Melanitta deglandi</em></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td class="tdc">o</td>
    <td></td>
    <td class="tdc">o</td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td><em>M. perspicillata</em></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">o</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td class="tdc">o</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td><em>M. nigra</em></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">*</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">o</td>
    <td></td>
    <td class="tdc">o</td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td><em>Mergus serrator</em></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">*</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td><em>Haliaeetus leucocephalus</em></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td class="tdc">x</td>
  </tr>
  <tr>
    <td><em>Falco peregrinus</em></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
  </tr>
  <tr>
    <td><em>Phalaropus fulicarius</em></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">o</td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">o</td>
  </tr>
  <tr>
    <td><em>Lobipes lobatus</em></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">o</td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">o</td>
  </tr>
  <tr>
    <td><em>Stercorarius</em> spp.</td>
    <td class="tdc">o</td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td class="tdc">x</td>
    <td class="tdc">?</td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td class="tdc">x</td>
    <td></td>
    <td class="tdc">o</td>
    <td class="tdc">x</td>
  </tr>
  <tr>
    <td><em>Larus hyperboreus</em></td>
    <td class="tdc">o</td>
    <td></td>
    <td></td>
    <td class="tdc">o</td>
    <td class="tdc">o</td>
    <td class="tdc">o</td>
    <td class="tdc">?</td>
    <td class="tdc">o</td>
    <td></td>
    <td></td>
    <td class="tdc">o</td>
    <td class="tdc">o</td>
    <td class="tdc">o</td>
    <td class="tdc">o</td>
    <td class="tdc">o</td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td class="tdc">x</td>
  </tr>
  <tr>
    <td><em>L. glaucescens</em></td>
    <td class="tdc">o</td>
    <td></td>
    <td></td>
    <td class="tdc">o</td>
    <td class="tdc">o</td>
    <td class="tdc">o</td>
    <td class="tdc">?</td>
    <td class="tdc">o</td>
    <td></td>
    <td></td>
    <td class="tdc">o</td>
    <td class="tdc">o</td>
    <td class="tdc">o</td>
    <td class="tdc">o</td>
    <td class="tdc">x</td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td class="tdc">x</td>
  </tr>
  <tr>
    <td><em>L. occidentalis</em></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td class="tdc">x</td>
    <td class="tdc">x</td>
    <td></td>
    <td class="tdc">o</td>
    <td></td>
    <td class="tdc">x</td>
    <td class="tdc">o</td>
    <td class="tdc">o</td>
    <td class="tdc">o</td>
    <td class="tdc">*</td>
    <td class="tdc">*</td>
    <td class="tdc">x</td>
    <td></td>
    <td class="tdc">x</td>
    <td class="tdc">o</td>
    <td class="tdc">x</td>
  </tr>
  <tr>
    <td><em>L. argentatus</em></td>
    <td class="tdc">o</td>
    <td></td>
    <td></td>
    <td class="tdc">o</td>
    <td class="tdc">o</td>
    <td class="tdc">o</td>
    <td></td>
    <td class="tdc">o</td>
    <td></td>
    <td class="tdc">x</td>
    <td class="tdc">o</td>
    <td class="tdc">o</td>
    <td class="tdc">o</td>
    <td class="tdc">o</td>
    <td class="tdc">o</td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td class="tdc">o</td>
    <td class="tdc">x</td>
  </tr>
  <tr>
    <td><em>L. heermanni</em></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
  </tr>
  <tr>
    <td><em>L. canus</em></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td class="tdc">o</td>
    <td class="tdc">o</td>
    <td class="tdc">o</td>
    <td class="tdc">x</td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
  </tr>
  <tr>
    <td><em>L. philadelphia</em></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td class="tdc">o</td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td><em>Rissa tridactyla</em></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td class="tdc">x</td>
    <td></td>
    <td class="tdc">o</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td><em>Xema sabini</em></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td class="tdc">o</td>
    <td class="tdc">o</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td><em>Sterna paradisaea</em></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td class="tdc">o</td>
    <td class="tdc">o</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td><em>S. hirundo</em></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">o</td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td><em>Uria aalge</em></td>
    <td class="tdc">x</td>
    <td class="tdc">o</td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">o</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td class="tdc">o</td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td><em>U. lomvia</em></td>
    <td class="tdc">x</td>
    <td class="tdc">o</td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">o</td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">o</td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td><em>Lunda cirrhata</em></td>
    <td class="tdc">?</td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td><em>Fratercula corniculata</em></td>
    <td></td>
    <td class="tdc">*</td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td><em>Cerorhinca monocerata</em></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td><em>Cepphus columba</em></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">o</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td class="tdc">o</td>
    <td></td>
  </tr>
  <tr>
    <td><em>Brachyramphus marmoratus</em></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">o</td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td><em>B. brevirostris</em></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">o</td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td><em>Synthliboramphus antiquus</em></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">o</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td><em>Ptychoramphus aleuticus</em></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td class="tdc">x</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
  </tr>
</table><span class="pagenum"><a name="Page_115" id="Page_115">[Pg 115]</a></span></div>

<div class="center">
<table border="0" cellpadding="4" cellspacing="0" summary="Table 16. Number of seabirds of different oceanographic regions having different numbers of categories of food in their diets.">
  <caption>Table 16. <em>Number of seabirds of different
  oceanographic regions having different numbers of categories of
  food in their diets.</em></caption>
  <tr>
    <th rowspan="2">Oceanographic region (domain)</th>
    <th colspan="0">Number of categories in the diets<a name="FNanchor_36_36" id="FNanchor_36_36"></a><a href="#Footnote_36_36" class="fnanchor">[36]</a></th>
  </tr>
  <tr>
    <th>1</th>
    <th>2</th>
    <th>3</th>
    <th>4</th>
    <th>5-7</th>
    <th>7</th>
    <th>8+</th>
  </tr>
  <tr>
    <td>Bering Sea coastal</td>
    <td class="tdn">3</td>
    <td class="tdn">11</td>
    <td class="tdn">9</td>
    <td class="tdn">6</td>
    <td class="tdn">5</td>
    <td class="tdn">4</td>
    <td class="tdn">5</td>
  </tr>
  <tr>
    <td>Bering Sea</td>
    <td class="tdn">2</td>
    <td class="tdn">6</td>
    <td class="tdn">5</td>
    <td class="tdn">7</td>
    <td class="tdn">0</td>
    <td class="tdn">5</td>
    <td class="tdn">0</td>
  </tr>
  <tr>
    <td>Alaskan Stream</td>
    <td class="tdn">3</td>
    <td class="tdn">14</td>
    <td class="tdn">14</td>
    <td class="tdn">5</td>
    <td class="tdn">4</td>
    <td class="tdn">4</td>
    <td class="tdn">5</td>
  </tr>
  <tr>
    <td>Central Subarctic</td>
    <td class="tdn">1</td>
    <td class="tdn">6</td>
    <td class="tdn">8</td>
    <td class="tdn">4</td>
    <td class="tdn">0</td>
    <td class="tdn">7</td>
    <td class="tdn">0</td>
  </tr>
  <tr>
    <td>North American Coastal</td>
    <td class="tdn">3</td>
    <td class="tdn">14</td>
    <td class="tdn">17</td>
    <td class="tdn">6</td>
    <td class="tdn">3</td>
    <td class="tdn">4</td>
    <td class="tdn">6</td>
  </tr>
  <tr>
    <td class="tdp">Total</td>
    <td class="tdn">12</td>
    <td class="tdn">51</td>
    <td class="tdn">53</td>
    <td class="tdn">28</td>
    <td class="tdn">12</td>
    <td class="tdn">24</td>
    <td class="tdn">16</td>
  </tr>
  <tr>
    <td>Percent total species (196)</td>
    <td class="tdn">6</td>
    <td class="tdn">26</td>
    <td class="tdn">27</td>
    <td class="tdn">14</td>
    <td class="tdn">6</td>
    <td class="tdn">12</td>
    <td class="tdn">8</td>
  </tr>
</table></div>

<p>It is readily apparent from the foregoing
comparisons that much overlap exists in the
prey eaten by seabirds within each community.
The question whether real competition
ever exists is academic. Competition perhaps
exists only rarely because seabirds partition
resources through use of different feeding
methods, selection of different-sized prey, and
habitat zonation. Table 18 lists feeding methods
(after Ashmole 1971 and Ainley 1977) and
the body size and bill length of each species
considered in this review. Bill length is
usually related directly to body size (Ashmole
1968; B&eacute;dard 1969<em>b</em>), but note, for instance,
that the longer species of the two kittiwakes
has the shorter bill. Body weight would be a
better measure of relative size than body size,
but few reliable weight data are available for
seabirds.</p>

<p>The use of different feeding methods by
species in each community grossly assigns
birds to feeding at different depths. Thus,
whereas shearwaters, puffins, and small gulls
(<em>Xema</em> sp., <em>Rissa</em> spp.) overlap almost entirely
in prey categories and even prey species, the
gulls can capture these organisms only at the
surface; the shearwaters capture them at shallow
depths; and the puffins capture them at
much deeper depths. Direct field observations
of this phenomenon are few but Gould (1971)
and Sealy (1973<em>a</em>) compared the diets of birds
feeding in mixed-species flocks. An example of
how even finer divergence in feeding methods
helps to partition food resources has been provided
by Spring (1971) in his comparison of
the two murres. Both species feed by diving to
great depths, but the thick-billed murre is
able to hover over the bottom and thereby is
better able to capture benthic organisms.</p>



<div class="center">
<table border="0" cellpadding="4" cellspacing="0" summary="Table 17. Number of species feeding at different trophic
  levels within seabird communities and habitats of the northeastern
  North Pacific Ocean and Bering Sea.">

  <caption>Table 17. <em>Number of species feeding at different trophic
  levels within seabird communities and habitats of the northeastern
  North Pacific Ocean and Bering Sea.</em> A single species can be
  represented in more than one level. (Trophic level I = vegetarian,
  II = secondary carnivore, III = tertiary carnivore, IV = upper
  level carnivore, Sc = scavenger [II-IV].)</caption>
  <tr>
    <th rowspan="2">Domain</th>
    <th colspan="4">Oceanic/offshore neritic</th>
    <th colspan="0">Inshore neritic</th>
  </tr>
  <tr>
    <th>II</th>
    <th>III</th>
    <th>IV</th>
    <th>Sc</th>
    <th>I</th>
    <th>II</th>
    <th>III</th>
    <th>IV</th>
    <th>Sc</th>
  </tr>
  <tr>
    <td>Bering Sea Coastal</td>
    <td class="tdn">11</td>
    <td class="tdn">17</td>
    <td class="tdn">1?</td>
    <td class="tdn">10</td>
    <td class="tdn">6</td>
    <td class="tdn">23</td>
    <td class="tdn">18</td>
    <td class="tdn">6</td>
    <td class="tdn">6</td>
  </tr>
  <tr>
    <td>Bering Sea</td>
    <td class="tdn">22</td>
    <td class="tdn">21</td>
    <td class="tdn">3?</td>
    <td class="tdn">11</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
  </tr>
  <tr>
    <td>Alaska Stream</td>
    <td class="tdn">21</td>
    <td class="tdn">19</td>
    <td class="tdn">1?</td>
    <td class="tdn">12</td>
    <td class="tdn">5</td>
    <td class="tdn">28</td>
    <td class="tdn">21</td>
    <td class="tdn">6</td>
    <td class="tdn">6</td>
  </tr>
  <tr>
    <td>Central Subarctic</td>
    <td class="tdn">23</td>
    <td class="tdn">22</td>
    <td class="tdn">3?</td>
    <td class="tdn">12</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
  </tr>
  <tr>
    <td>North American Coastal</td>
    <td class="tdn">25</td>
    <td class="tdn">24</td>
    <td class="tdn">3?</td>
    <td class="tdn">11</td>
    <td class="tdn">3</td>
    <td class="tdn">28</td>
    <td class="tdn">35</td>
    <td class="tdn">7</td>
    <td class="tdn">10</td>
  </tr>
  <tr>
    <td class="tdp">Total</td>
    <td class="tdn">102</td>
    <td class="tdn">103</td>
    <td class="tdn">11?</td>
    <td class="tdn">56</td>
    <td class="tdn">14</td>
    <td class="tdn">79</td>
    <td class="tdn">74</td>
    <td class="tdn">19</td>
    <td class="tdn">22</td>
  </tr>
  <tr>
    <td>Proportion</td>
    <td class="tdn">0.38</td>
    <td class="tdn">0.39</td>
    <td class="tdn">0.02<a name="FNanchor_37_37" id="FNanchor_37_37"></a><a href="#Footnote_37_37" class="fnanchor">[37]</a> </td>
    <td class="tdn">0.21</td>
    <td class="tdn">0.07</td>
    <td class="tdn">0.38</td>
    <td class="tdn">0.28</td>
    <td class="tdn">0.09</td>
    <td class="tdn">0.10</td>
  </tr>
</table><span class="pagenum"><a name="Page_116" id="Page_116">[Pg 116]</a></span></div>

<div class="center">
<table border="0" cellpadding="4" cellspacing="0" summary="Table 18.
Size relationships and feeding methods of major species in the eastern
North Pacific and Bering Sea.">
    <caption>Table 18. <em>Size relationships and feeding methods of
    major species in the eastern North Pacific and Bering Sea.</em>
    (D = dive, SS = surface seize, PP = pursuit plunge, Di = dip, P
    = plunge, T = tip, x = eats seabirds, A = piracy, SP = shallow
    plunge.)</caption>
<thead>
  <tr>
    <th>Species</th>
    <th>Body length<a name="FNanchor_38_38" id="FNanchor_38_38"></a><a href="#Footnote_38_38" class="fnanchor">[38]</a> (cm)</th>
    <th>Bill length<a name="FNanchor_39_39" id="FNanchor_39_39"></a><a href="#Footnote_39_39" class="fnanchor">[39]</a> (mm)</th>
    <th>Feeding<a name="FNanchor_40_40" id="FNanchor_40_40"></a><a href="#Footnote_40_40" class="fnanchor">[40]</a> method</th>
  </tr>
</thead>
<tbody>
  <tr>
    <td><em>Gavia adamsii</em></td>
    <td class="tdr">63.5</td>
    <td class="tdc">90-91</td>
    <td class="tdc">D</td>
  </tr>
  <tr>
    <td><em>G. immer</em></td>
    <td class="tdr">61.0</td>
    <td class="tdc">80-82</td>
    <td class="tdc">D</td>
  </tr>
  <tr>
    <td><em>G. arctica</em></td>
    <td class="tdr">45.7</td>
    <td class="tdc">51-52</td>
    <td class="tdc">D</td>
  </tr>
  <tr>
    <td><em>G. stellata</em></td>
    <td class="tdr">43.5</td>
    <td class="tdc">51-52</td>
    <td class="tdc">D</td>
  </tr>
  <tr>
    <td><em>Podiceps grisegena</em></td>
    <td class="tdr">33.0</td>
    <td class="tdc">48-50</td>
    <td class="tdc">D</td>
  </tr>
  <tr>
    <td><em>P. nigricollis</em></td>
    <td class="tdr">22.9</td>
    <td class="tdc">24-26</td>
    <td class="tdc">D</td>
  </tr>
  <tr>
    <td><em>P. auritus</em></td>
    <td class="tdr">24.1</td>
    <td class="tdc">23-24</td>
    <td class="tdc">D</td>
  </tr>
  <tr>
    <td><em>Aechmophorus occidentalis</em></td>
    <td class="tdr">45.7</td>
    <td class="tdc">65-76</td>
    <td class="tdc">D</td>
  </tr>
  <tr>
    <td><em>Diomedea nigripes</em></td>
    <td class="tdr">71.1</td>
    <td class="tdc">141-144</td>
    <td class="tdc">SS</td>
  </tr>
  <tr>
    <td><em>D. immutabilis</em></td>
    <td class="tdr">71.1</td>
    <td class="tdc">102-112</td>
    <td class="tdc">SS</td>
  </tr>
  <tr>
    <td><em>Fulmarus glacialis</em></td>
    <td class="tdr">45.7</td>
    <td class="tdc">36-37</td>
    <td class="tdc">SS</td>
  </tr>
  <tr>
    <td><em>Puffinus carneipes</em></td>
    <td class="tdr">45.7</td>
    <td class="tdc">41-46</td>
    <td class="tdc">PP</td>
  </tr>
  <tr>
    <td><em>P. creatopus</em></td>
    <td class="tdr">45.7</td>
    <td class="tdc">41-46</td>
    <td class="tdc">PP</td>
  </tr>
  <tr>
    <td><em>P. bulleri</em></td>
    <td class="tdr">38.1</td>
    <td class="tdc">38-45</td>
    <td class="tdc">PP</td>
  </tr>
  <tr>
    <td><em>P. griseus</em></td>
    <td class="tdr">40.3</td>
    <td class="tdc">41-42</td>
    <td class="tdc">PP</td>
  </tr>
  <tr>
    <td><em>P. tenuirostris</em></td>
    <td class="tdr">38.1</td>
    <td class="tdc">31-32</td>
    <td class="tdc">PP</td>
  </tr>
  <tr>
    <td><em>Oceanodroma furcata</em></td>
    <td class="tdr">19.0</td>
    <td class="tdc">15</td>
    <td class="tdc">Di,SS</td>
  </tr>
  <tr>
    <td><em>O. leucorhoa</em></td>
    <td class="tdr">19.0</td>
    <td class="tdc">16</td>
    <td class="tdc">Di,SS</td>
  </tr>
  <tr>
    <td><em>Pterodroma inexpectata</em></td>
    <td class="tdr">29.2</td>
    <td class="tdc">26-27</td>
    <td class="tdc">SS</td>
  </tr>
  <tr>
    <td><em>Phalacrocorax auritus</em></td>
    <td class="tdr">68.6</td>
    <td class="tdc">55-57</td>
    <td class="tdc">D</td>
  </tr>
  <tr>
    <td><em>P. penicillatus</em></td>
    <td class="tdr">73.7</td>
    <td class="tdc">66-71</td>
    <td class="tdc">D</td>
  </tr>
  <tr>
    <td><em>P. urile</em></td>
    <td class="tdr">71.1</td>
    <td class="tdc">54-55</td>
    <td class="tdc">D</td>
  </tr>
  <tr>
    <td><em>P. pelagicus</em></td>
    <td class="tdr">55.9</td>
    <td class="tdc">47-50</td>
    <td class="tdc">D</td>
  </tr>
  <tr>
    <td><em>Pelecanus occidentalis</em></td>
    <td class="tdr">104.0</td>
    <td class="tdc">294-319</td>
    <td class="tdc">P</td>
  </tr>
  <tr>
    <td><em>Branta</em> spp. <em>(bernicla)</em></td>
    <td class="tdr">43.5</td>
    <td class="tdc">33-36</td>
    <td class="tdc">T</td>
  </tr>
  <tr>
    <td><em>Philacte canagica</em></td>
    <td class="tdr">45.7</td>
    <td class="tdc">37-42</td>
    <td class="tdc">T</td>
  </tr>
  <tr>
    <td><em>Anas</em> spp.</td>
    <td class="tdr">40.0</td>
    <td class="tdc">32-35</td>
    <td class="tdc">T</td>
  </tr>
  <tr>
    <td><em>Clangula hyemalis</em></td>
    <td class="tdr">38.1</td>
    <td class="tdc">25-27</td>
    <td class="tdc">D</td>
  </tr>
  <tr>
    <td><em>Histrionicus histrionicus</em></td>
    <td class="tdr">30.5</td>
    <td class="tdc">25-28</td>
    <td class="tdc">D</td>
  </tr>
  <tr>
    <td><em>Polysticta stelleri</em></td>
    <td class="tdr">30.5</td>
    <td class="tdc">37-43</td>
    <td class="tdc">D</td>
  </tr>
  <tr>
    <td><em>Somateria mollisima</em></td>
    <td class="tdr">43.5</td>
    <td class="tdc">45-55</td>
    <td class="tdc">D</td>
  </tr>
  <tr>
    <td><em>S. spectabilis</em></td>
    <td class="tdr">40.3</td>
    <td class="tdc">31-33</td>
    <td class="tdc">D</td>
  </tr>
  <tr>
    <td><em>S. fischeri</em></td>
    <td class="tdr">38.1</td>
    <td class="tdc">22-26</td>
    <td class="tdc">D</td>
  </tr>
  <tr>
    <td><em>Melanitta deglandi</em></td>
    <td class="tdr">35.6</td>
    <td class="tdc">41-44</td>
    <td class="tdc">D</td>
  </tr>
  <tr>
    <td><em>M. perspicillata</em></td>
    <td class="tdr">40.3</td>
    <td class="tdc">ca. 40</td>
    <td class="tdc">D</td>
  </tr>
  <tr>
    <td><em>M. nigra</em></td>
    <td class="tdr">35.6</td>
    <td class="tdc">42-47</td>
    <td class="tdc">D</td>
  </tr>
  <tr>
    <td><em>Mergus serrator</em></td>
    <td class="tdr">40.3</td>
    <td class="tdc">45-54</td>
    <td class="tdc">D</td>
  </tr>
  <tr>
    <td><em>Haliaeetus leucocephalus</em></td>
    <td class="tdr">80.0</td>
    <td class="tdc">52-54</td>
    <td class="tdc">X</td>
  </tr>
  <tr>
    <td><em>Falco peregrinus</em></td>
    <td class="tdr">37.5</td>
    <td class="tdc">21-25</td>
    <td class="tdc">X</td>
  </tr>
  <tr>
    <td><em>Phalaropus fulicarius</em></td>
    <td class="tdr">16.5</td>
    <td class="tdc">22</td>
    <td class="tdc">SS</td>
  </tr>
  <tr>
    <td><em>Lobipes lobatus</em></td>
    <td class="tdr">15.2</td>
    <td class="tdc">22</td>
    <td class="tdc">SS</td>
  </tr>
  <tr>
    <td><em>Stercorarius pomarinus</em></td>
    <td class="tdr">43.5</td>
    <td class="tdc">40</td>
    <td class="tdc">SS,A</td>
  </tr>
  <tr>
    <td><em>S. parasiticus</em></td>
    <td class="tdr">40.3</td>
    <td class="tdc">32</td>
    <td class="tdc">SS,A</td>
  </tr>
  <tr>
    <td><em>S. longicaudus</em></td>
    <td class="tdr">38.1</td>
    <td class="tdc">29</td>
    <td class="tdc">SS,A</td>
  </tr>
  <tr>
    <td><em>Larus hyperboreus</em></td>
    <td class="tdr">61.0</td>
    <td class="tdc">55-60</td>
    <td class="tdc">SS</td>
  </tr>
  <tr>
    <td><em>L. glaucescens</em></td>
    <td class="tdr">55.9</td>
    <td class="tdc">54-58</td>
    <td class="tdc">SS</td>
  </tr>
  <tr>
    <td><em>L. occidentalis</em></td>
    <td class="tdr">53.0</td>
    <td class="tdc">54-57</td>
    <td class="tdc">SS,Di</td>
  </tr>
  <tr>
    <td><em>L. argentatus</em></td>
    <td class="tdr">50.8</td>
    <td class="tdc">48-54</td>
    <td class="tdc">SS,Di</td>
  </tr>
  <tr>
    <td><em>L. californicus</em></td>
    <td class="tdr">43.5</td>
    <td class="tdc">45-50</td>
    <td class="tdc">SS,Di</td>
  </tr>
  <tr>
    <td><em>L. heermanni</em></td>
    <td class="tdr">38.1</td>
    <td class="tdc">42-46</td>
    <td class="tdc">SS,Di</td>
  </tr>
  <tr>
    <td><em>L. canus</em></td>
    <td class="tdr">35.6</td>
    <td class="tdc">34-36</td>
    <td class="tdc">SS,Di</td>
  </tr>
  <tr>
    <td><em>L. philadelphia</em></td>
    <td class="tdr">27.9</td>
    <td class="tdc">30-31</td>
    <td class="tdc">Di</td>
  </tr>
  <tr>
    <td><em>Rissa tridactyla</em></td>
    <td class="tdr">34.2</td>
    <td class="tdc">39-40</td>
    <td class="tdc">Di</td>
  </tr>
  <tr>
    <td><em>R. brevirostris</em></td>
    <td class="tdr">38.1</td>
    <td class="tdc">29-30</td>
    <td class="tdc">Di</td>
  </tr>
  <tr>
    <td><em>Xema sabini</em></td>
    <td class="tdr">27.9</td>
    <td class="tdc">26-27</td>
    <td class="tdc">Di</td>
  </tr>
  <tr>
    <td><em>Sterna paradisaea</em></td>
    <td class="tdr">38.1</td>
    <td class="tdc">31-33</td>
    <td class="tdc">Di,SP</td>
  </tr>
  <tr>
    <td><em>S. hirundo/forsteri</em></td>
    <td class="tdr">35.6</td>
    <td class="tdc">36-39</td>
    <td class="tdc">Di,SP</td>
  </tr>
  <tr>
    <td><em>S. aleutica</em></td>
    <td class="tdr">33.0</td>
    <td class="tdc">33</td>
    <td class="tdc">Di,SP</td>
  </tr>
  <tr>
    <td><em>Uria aalge</em></td>
    <td class="tdr">35.6</td>
    <td class="tdc">43-47</td>
    <td class="tdc">D</td>
  </tr>
  <tr>
    <td><em>U. lomvia</em></td>
    <td class="tdr">35.6</td>
    <td class="tdc">39-42</td>
    <td class="tdc">D</td>
  </tr>
  <tr>
    <td><em>Lunda cirrhata</em></td>
    <td class="tdr">31.8</td>
    <td class="tdc">57-60</td>
    <td class="tdc">D</td>
  </tr>
  <tr>
    <td><em>Fratercula corniculata</em></td>
    <td class="tdr">29.2</td>
    <td class="tdc">49-51</td>
    <td class="tdc">D</td>
  </tr>
  <tr>
    <td><em>Cerorhinca monocerata</em></td>
    <td class="tdr">29.2</td>
    <td class="tdc">34-35</td>
    <td class="tdc">D</td>
  </tr>
  <tr>
    <td><em>Cepphus columba</em></td>
    <td class="tdr">26.7</td>
    <td class="tdc">32-33</td>
    <td class="tdc">D</td>
  </tr>
  <tr>
    <td><em>Brachyramphus marmoratus</em></td>
    <td class="tdr">20.3</td>
    <td class="tdc">15</td>
    <td class="tdc">D</td>
  </tr>
  <tr>
    <td><em>B. brevirostris</em></td>
    <td class="tdr">19.0</td>
    <td class="tdc">10</td>
    <td class="tdc">D</td>
  </tr>
  <tr>
    <td><em>Synthliboramphus antiquus</em></td>
    <td class="tdr">20.3</td>
    <td class="tdc">13</td>
    <td class="tdc">D</td>
  </tr>
  <tr>
    <td><em>Ptychoramphus aleuticus</em></td>
    <td class="tdr">17.8</td>
    <td class="tdc">19</td>
    <td class="tdc">D</td>
  </tr>
  <tr>
    <td><em>Aethia pygmaea</em></td>
    <td class="tdr">16.5</td>
    <td class="tdc">8-9</td>
    <td class="tdc">D</td>
  </tr>
  <tr>
    <td><em>A. pusilla</em></td>
    <td class="tdr">13.3</td>
    <td class="tdc">8</td>
    <td class="tdc">D</td>
  </tr>
  <tr>
    <td><em>A. cristatella</em></td>
    <td class="tdr">17.8</td>
    <td class="tdc">11</td>
    <td class="tdc">D</td>
  </tr>
  <tr>
    <td><em>Cyclorrhynchus psittaculus</em></td>
    <td class="tdr">18.4</td>
    <td class="tdc">15</td>
    <td class="tdc">D</td>
  </tr>
</tbody>
</table></div>

<p>The scavengers (generalists) offer a good
example of how a range of bird and bill sizes is
usually represented among species having
similar diets and feeding methods. The progression
of oceanic scavenger sizes is graded
rather evenly from the black-footed albatross
down to the northern fulmar, to the scaled
petrel, to the storm-petrel. All these species
capture prey that occur only at or near the
water surface. Recently Sanger (1973) reported
appreciable numbers of glaucous-winged
gulls <em>(Larus glaucescens)</em> and herring
gulls <em>(L. argentatus)</em>, noted neritic scavengers,
out in the oceanic realm of the petrel. He
presented limited data that suggested an
overlap between the diet of these gulls and
that of black-footed albatrosses, as noted by
Miller (1940). It would not be surprising if
these gulls were as much generalists in the
oceanic habitat as they are in the neritic. Interestingly,
their bill and body sizes fall between
those of the albatross and the fulmar,
thus in theory enabling them to invade the
oceanic habitat without great competition. It
is likely that their invasion occurred during
historical times and is related to their habit of
following fishing boats from shore out to sea
(Sanger 1973). If so, the gulls might be assuming
from other species part of a previously uncontested
resource.</p>

<p>Another interesting group of species that
shows close similarities in diet consists of the
piscivorous loons, grebes, and mergansers.
All these birds, including seven or eight
species, apparently feed on fish occurring on
or near the bottom in the inshore neritic habitat.
Again, however, an even progression in
size exists: yellow-billed loon <em>(Gavia adamsii)</em>,
common loon <em>(G. immer)</em>, arctic loon <em>(G. arctica)</em>,
red-throated loon <em>(G. stellata)</em>, western
grebe <em>(Aechmophorus occidentalis)</em>, red-necked
grebe <em>(Podiceps grisegena)</em>, and com<span class="pagenum"><a name="Page_118" id="Page_118">[Pg 118]</a></span>mon
merganser <em>(Mergus merganser)</em>. Most
likely then, they select different-sized fish.
Another example of this phenomenon is provided
by the eight neritic gulls, which are
largely scavengers and show a remarkably
even progression in bill and body size. Finally,
as shown clearly by B&eacute;dard (1969<em>a</em>, 1969<em>b</em>)
and Harris (1970), alcids of different sizes
select different-sized prey, often of the same
species.</p>

<p>A final important way in which seabirds
partition available resources is by inhabiting
different zones. Zonation is especially evident
during the breeding season when species common
to the same breeding site sort themselves
out according to the distances they range for
food. This phenomenon was discussed by
Murphy (1936), Shuntov (1974), Sealy (1972),
Cody (1973), and Scott (1973).</p>


<h4>Trophic Relations and
Seabird Conservation</h4>

<p>The species that appear to have specialized
food habits (if further research confirms that
indeed they do) are probably very sensitive to
vagaries in food availability or are, at least,
much more sensitive than other species. Some
specialists which also have very restricted distributions
would, therefore, be susceptible to
localized catastrophes occurring where
specialists are concentrated around the food
resource. This is proved in the case of the
scoters, which are both specialized and rather
restricted to nearshore beds of molluscs and
have fallen victim to local oil slicks (Smail
et al. 1972). An example of another potentially
critical situation is that of the black
brant, which at certain times of the year concentrate
their entire population around eelgrass
beds in Bristol Bay, Alaska, where
much offshore oil drilling may soon occur.</p>

<p>Birds adapted to feed by diving, with the
exception of cormorants, spend most of their
time in the water. These species are therefore
most susceptible to oiling (Smail et al. 1972),
but pursuit plungers (the shearwaters) are
also highly susceptible (Point Reyes Bird Observatory,
unpublished data). A characteristic
of polar and subpolar seabird communities is
the high percentage of birds that feed by diving
and pursuit plunging. These birds are
mostly absent from tropical and subtropical
communities because feeding by these methods
is not adaptive there (Ainley 1977).
Hence, oil pollution has all the potential of
rendering maladaptive the principal feeding
methods of many polar seabirds.</p>

<p>Another way in which seabird feeding relates
to conservation problems concerns competition
between birds and man for commercially
valuable fishes. A related problem is the
mass mortality of seabirds due to man's fishing
gear. An acute situation is the drowning
of seabirds caught in salmon gill nets (Bartonek
et al. 1974; Pacific Seabird Group 1975;
Ripley 1975; King et al., this volume). Immediate
action is definitely required.</p>

<p>Further, competition between birds and
man for the same resource has the potential
for disastrous effects on bird populations if
humans out-compete the birds and overfish
the resource. A classic example, reviewed by
Idyll (1973), is the possible collapse of the
Peruvian anchovy <em>(Engraulis ringens)</em> fishery;
if overfishing and an El Ni&ntilde;o should coincide,
the Peruvian seabird populations could collapse
as well. The California fisheries and apparently
the double-crested cormorants that
nest on the Farallon Islands have both suffered
from the demise of the Pacific sardine
<em>(Sardinops caerulea)</em> in the California current
(Ainley and Lewis 1974). In regulating fish
harvests, fishery organizations should include
in their calculations the harvest by creatures
other than man (Schaefer 1970), rather than
evading the issue by referring to a vague
"natural mortality."</p>

<hr class="tb" />

<p>Finally, fishing by humans can benefit seabirds
by removing fish (or whales) that compete
with birds for food (Laws 1977). A potential
example is that of northern California,
where salmon and seabirds both feed heavily
on juvenile rockfishes (Fitch and Lavenberg
1971; Point Reyes Bird Observatory, unpublished
data). Harvest of salmon should theoretically
leave more rockfish available for
birds to eat. This sort of situation has not yet
been fully documented and definitely warrants
further study, especially in such areas
as the Bering Sea, where some fish stocks
have become depressed due to overfishing
(Gulland 1970).</p>

<p><span class="pagenum"><a name="Page_119" id="Page_119">[Pg 119]</a></span></p>


<h4>Recommendations for
Further Research</h4>

<p>Many people realize intuitively that seabirds
are important members of marine ecosystems.
Although the supporting evidence is
not now available, it will be needed if seabirds
are to be protected. Emotion alone will not
justify the protection of seabirds in an age
when the human race moves steadily toward
global famine. The job at hand is, in part, to
sell seabirds, not just to the public, government
officials, executives of oil companies, or
fish-packing concerns, but also to marine
biologists and oceanographers, for the scientists
have the best means to study organisms
at sea. We must move away from the concept
that seabirds are merely yo-yos of various
sizes, shapes, and colors on strings of various
lengths that venture forth to sea from the
land, grab a quick lunch, and then return to
the safety of terra firma. Seabirds are marine
organisms and deserve at least as much research
attention as that currently given
marine mammals.</p>

<p>The information now available on seabird
diets is largely presented in terms of the number
and volume of various prey species taken.
Whereas these data provide the relative importance
of prey, fishery data on prey stocks
are usually measured in terms of biomass.
Thus, it is difficult to relate seabird data to
the immense wealth of information on biological
oceanography. If we are to recognize the
importance of seabirds in the nutrient and
energy cycling of marine ecosystems, rather
than considering them merely as "yo-yo
predators," we must relate them to the total
marine community.</p>

<p>The goal of marine ornithologists should be
to refine and broaden considerably in detail
such studies as those by Sanger (1972), Shuntov
(1974), and Laws (1977), who attempted to
assess the relations between seabird populations
and stocks of other marine organisms
for the northern North Pacific, the world
oceans, and the Antarctic, respectively. The
trophic roles played by seabirds must be
studied in detail at the community level year-round
before those analyses can be properly
refined. Another exemplary work is that done
by Brownell (1974), who studied trophic relations
of higher vertebrates off Uruguay, including
dolphins, pinnipeds, seabirds, and
some large fish. In a review study, Sanger
(1974) considered the food-chain relations of
similar vertebrates in the Bering Sea. These
sorts of studies will serve to bring the role of
seabirds into perspective with other upper
trophic level feeders.</p>


<h4>Acknowledgments</h4>

<p>We much appreciate the opportunity to participate
in the symposium at which this paper
was presented. The encouragement and help
given by J. C. Bartonek was indispensable.
D. G. Ainley's participation in the symposium
was supported by the Point Reyes
Bird Observatory. This is contribution No.
124 of the Point Reyes Bird Observatory.</p>


<h4>References</h4>

<div class="hangindent">

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<div class="footnotes"><h4>FOOTNOTES:</h4>

<div class="footnote">

<p><a name="Footnote_15_15" id="Footnote_15_15"></a><a href="#FNanchor_15_15"><span class="label">[15]</span></a> Present address: U.S. Fish and Wildlife Service,
Office of Biological Services, 1011 East Tudor
Road, Anchorage, Alaska 99503.</p></div>

<div class="footnote">

<p><a name="Footnote_16_16" id="Footnote_16_16"></a><a href="#FNanchor_16_16"><span class="label">[16]</span></a> Other incidental prey were squid and atherinid fishes, both at the Farallon Islands.</p></div>

<div class="footnote">

<p><a name="Footnote_17_17" id="Footnote_17_17"></a><a href="#FNanchor_17_17"><span class="label">[17]</span></a> Other incidental prey were squid and such fishes as atherinids, <em>Zaniolepis</em>, <em>Genyonemus</em> and <em>Peprilus</em> at the Farallones, and atherinids,
<em>Trachurus</em> and <em>Heterostichus</em> at San Diego.</p></div>

<div class="footnote">

<p><a name="Footnote_18_18" id="Footnote_18_18"></a><a href="#FNanchor_18_18"><span class="label">[18]</span></a> Other incidental prey were polychaetes at Netarts and the Farallon Islands.</p></div>

<div class="footnote">

<p><a name="Footnote_19_19" id="Footnote_19_19"></a><a href="#FNanchor_19_19"><span class="label">[19]</span></a> Principal sources: Bent 1925; Cleaver and Franett 1945; Cottam 1939; Cottam and Knappen 1939; Kortright 1942; Mabbot 1920; McGilvrey
1967; Munro and Clemens 1939; Roberts and Huntington 1959.</p></div>

<div class="footnote">

<p><a name="Footnote_20_20" id="Footnote_20_20"></a><a href="#FNanchor_20_20"><span class="label">[20]</span></a> Other incidental items were the fish <em>Cololabis</em> and <em>Peprilus</em> at the Farallon Islands.</p></div>

<div class="footnote">

<p><a name="Footnote_21_21" id="Footnote_21_21"></a><a href="#FNanchor_21_21"><span class="label">[21]</span></a> Other incidental items were myctophid fish in northeastern Canada.</p></div>

<div class="footnote">

<p><a name="Footnote_22_22" id="Footnote_22_22"></a><a href="#FNanchor_22_22"><span class="label">[22]</span></a> Other incidental items included the lamprey <em>(Lampetra)</em> at the Farallon Islands.</p></div>

<div class="footnote">

<p><a name="Footnote_23_23" id="Footnote_23_23"></a><a href="#FNanchor_23_23"><span class="label">[23]</span></a> Bent (1946) listed "fish" as prey.</p></div>

<div class="footnote">

<p><a name="Footnote_24_24" id="Footnote_24_24"></a><a href="#FNanchor_24_24"><span class="label">[24]</span></a> Grinnell (1897) listed "fish" as the major dietary component.</p></div>

<div class="footnote">

<p><a name="Footnote_25_25" id="Footnote_25_25"></a><a href="#FNanchor_25_25"><span class="label">[25]</span></a> B&eacute;dard (1969<em>a</em>) also listed "fish" as an incidental item.</p></div>

<div class="footnote">

<p><a name="Footnote_26_26" id="Footnote_26_26"></a><a href="#FNanchor_26_26"><span class="label">[26]</span></a> Other incidental prey were copepods and isopods.</p></div>

<div class="footnote">

<p><a name="Footnote_27_27" id="Footnote_27_27"></a><a href="#FNanchor_27_27"><span class="label">[27]</span></a> Other incidental prey were pholids in Denmark.</p></div>

<div class="footnote">

<p><a name="Footnote_28_28" id="Footnote_28_28"></a><a href="#FNanchor_28_28"><span class="label">[28]</span></a> Other incidental prey were copepods and cephalopods in North Atlantic areas.</p></div>

<div class="footnote">

<p><a name="Footnote_29_29" id="Footnote_29_29"></a><a href="#FNanchor_29_29"><span class="label">[29]</span></a> Other incidental prey were isopods in western North America and fish eggs near Vancouver Island.</p></div>

<div class="footnote">

<p><a name="Footnote_30_30" id="Footnote_30_30"></a><a href="#FNanchor_30_30"><span class="label">[30]</span></a> Other incidental prey were fish eggs in Denmark.</p></div>

<div class="footnote">

<p><a name="Footnote_31_31" id="Footnote_31_31"></a><a href="#FNanchor_31_31"><span class="label">[31]</span></a> Offal from wounded whales and seals, and bits of food, primarily crustaceans and fish, from feeding whales are
important scavenger foods (Bent 1922).</p></div>

<div class="footnote">

<p><a name="Footnote_32_32" id="Footnote_32_32"></a><a href="#FNanchor_32_32"><span class="label">[32]</span></a> Other incidental prey were isopods in the North Pacific.</p></div>

<div class="footnote">

<p><a name="Footnote_33_33" id="Footnote_33_33"></a><a href="#FNanchor_33_33"><span class="label">[33]</span></a> Other incidental prey were the fish <em>Merluccius</em> at the Farallon Islands.</p></div>

<div class="footnote">

<p><a name="Footnote_34_34" id="Footnote_34_34"></a><a href="#FNanchor_34_34"><span class="label">[34]</span></a> Other incidental prey were isopods near the Pribilofs and in the Chukchi Sea, and amphipods in the latter area; Bent (1921) considered "crustaceans"
to be major prey.</p></div>

<div class="footnote">

<p><a name="Footnote_35_35" id="Footnote_35_35"></a><a href="#FNanchor_35_35"><span class="label">[35]</span></a> Study conducted during period of breeding failure.</p></div>

<div class="footnote">

<p><a name="Footnote_36_36" id="Footnote_36_36"></a><a href="#FNanchor_36_36"><span class="label">[36]</span></a> These are the "food categories" of Tables 11-15. Items included in diets are not included here.</p></div>

<div class="footnote">

<p><a name="Footnote_37_37" id="Footnote_37_37"></a><a href="#FNanchor_37_37"><span class="label">[37]</span></a> Proportion based on the arbitrary assumption that half (5) of the 11 species in question catch and eat
birds at sea.</p></div>

<div class="footnote">

<p><a name="Footnote_38_38" id="Footnote_38_38"></a><a href="#FNanchor_38_38"><span class="label">[38]</span></a> Information on body sizes (length) is from Robbins et al. (1966).</p></div>

<div class="footnote">

<p><a name="Footnote_39_39" id="Footnote_39_39"></a><a href="#FNanchor_39_39"><span class="label">[39]</span></a> Information on bill lengths is from Palmer (1962), Dement'ev et al. (1968), and Friedmann (1950).</p></div>

<div class="footnote">

<p><a name="Footnote_40_40" id="Footnote_40_40"></a><a href="#FNanchor_40_40"><span class="label">[40]</span></a> Feeding methods are from Ashmole (1971) as adapted by Ainley (unpubl. manuscr.).</p>

<p><span class="pagenum"><a name="Page_123" id="Page_123">[Pg 123]</a></span></p></div></div>
<div class="chapter"></div>
<hr class="chap" />




<h3>Population Dynamics in Northern Marine Birds</h3>


<p class="center">by</p>

<p class="center">William H. Drury</p>

<p class="center">College of the Atlantic<br />
Bar Harbor, Maine 04609
</p>


<h4>Abstract</h4>

<div class="blockquot">

<p>It seems only reasonable to assume that populations of marine birds fluctuate
even when not disturbed by man; such fluctuations would result both from the
secondary effects of species adaptive tactics and from changes in the marine
environment. I briefly review some human activities and some other natural processes
that have resulted in changes in numbers and distribution of seabirds and
present a short discussion of theoretical models which emphasizes that conclusions
drawn or predictions made from models of the dynamics of populations
depend upon the assumptions about stability that were used in preparing the
models. I then review those special characteristics of seabirds which are directly
relevant to planning programs intended to protect seabirds or encourage their
increase and identify several goals for improving our understanding of the population
dynamics and biology of marine birds. My general conclusion is that
enough is already known to undertake effective conservation programs, and that
time is pressing.</p></div>

<p>Seabirds have been categorized as renewable
resources in only a few places, although
their symbolic value has been recognized for
centuries (for example, the medieval poem
"The Seafarer" and the designs on Saint
Cuthbert's tunic). With the exception of the
Russians (Belopol'skii 1961; Uspenski 1956),
the Australians (Serventy 1967), and the Icelanders,
industrialized peoples have not considered
seabirds to be salable and therefore
worth managing. Yet during many centuries
the seabirds of the northern seas were a major
food for coastal and island villages (Bent
1919, 1921, 1922; Fisher and Lockley 1954).</p>

<p>Some biological principles that affect the
dynamics of seabird populations are identified
in this paper. I believe these principles must
form the basis of plans to maintain and increase
seabird numbers.</p>

<p>I describe some observations of population
changes, review briefly the conflicting theoretical
frameworks for population dynamics,
and identify some of the biological characteristics
of marine birds that affect the way in
which population changes occur. The terms
"seabirds" and "marine birds" are used interchangeably
for those bird species which depend
upon salt water for some part of their annual
cycle (c.f., the Pacific Seabird Group).</p>


<h4>Population Fluctuations</h4>

<p>Broadly stated, the populations of northern
seabirds have shown marked short-and long-term
fluctuations. Most authors have assumed
that all such fluctuations reflect
human disturbance of the natural system, because
of the obvious effects of human predation
during the last 200 years.</p>


<h5><em>Human Impact</em></h5>

<p>In the centuries before people traveled extensively
between islands, seabirds were
taken in ways that we judge must have
allowed the survival of the colonies (e.g., those
at the Faroes or Saint Kilda, those in Iceland
and Greenland, or those in the Aleutian Islands
and the Bering Strait). We presume
either that the populations of island peoples
were regulated by shortage of resources other<span class="pagenum"><a name="Page_124" id="Page_124">[Pg 124]</a></span>
than seabirds or that those who overcropped
and eliminated the seabirds suffered the
consequences.</p>


<h6>Negative Effects</h6>

<p>When a sea-going, commodity-oriented way
of life evolved, seabirds were killed in huge
numbers for such uses as the plumage trade,
fish bait, or rendering into oil (Tuck 1960;
Fisher and Lockley 1954). Even the elimination
of several colonies&mdash;e.g., Funk Island,
Newfoundland (Tuck 1960); Seal Island, Eastern
Egg Rock, Maine (Norton 1921); Muskeget,
Massachusetts (Forbush 1929)&mdash;may
have had little effect on the rate of cropping
because those who killed off one source could
probably seek out another. As the colonial
seabirds became scarce they became more
valuable, which stimulated more intensive
pursuit of the remnants (Dutcher 1901, 1904).</p>

<p>In some places where seabird colonies did
not supply a croppable economic resource, the
islands were used for alternative crops with at
least temporary commodity value (e.g., foxes
were introduced in the Aleutian Islands; Bent
1919). Large herbivores were introduced to
supply meat for island residents (e.g., Saint
Matthew Island; Klein 1959), as well as pigs,
cattle, sheep, goats, and rabbits on islands in
the North Atlantic and southern oceans
(many authors). Increases in many seabird
populations over the last 75 years have been
generally associated with relief from predation
by humans such as the fowlers, eggers,
and plume hunters of the 19th century. Such
relief may have been partly responsible for the
increase of North Atlantic gannets, <em>Sula bassana</em>,
and common murres or guillemots, <em>Uria
aalge</em> (Fisher and Vevers 1943, 1944; Cramp
et al. 1974). On a smaller scale, several population
increases along the coast of New England
have been recorded following the enactment
of protective legislation (Dutcher 1901, 1904;
Norton 1921, 1924; Palmer 1949; Drury 1973).</p>

<p>Coulson (1974) argued that in addition to relief
from predation, the explosion of the population
of kittiwakes <em>(Rissa tridactyla)</em> in this
century resulted from access to previously un-occupiable
breeding sites. Nesting cliffs and
buildings suitable for kittiwake nesting are
abundant and now protected from egging or
fowling.</p>


<h6>Positive Effects</h6>

<p>There can be little doubt that human activities
have also had marked positive effects in
some cases. For example, Fisher (1952) suggested
that the North Atlantic fulmar <em>(Fulmarus
glacialis)</em> was provided food first by
whaling, then by commercial fishing, and that
this food allowed the species to increase
steadily over the last 3 centuries.</p>

<p>The worldwide increase of gulls (<em>Larus
argentatus</em>, <em>L. fuscus</em>, <em>L. dominicanus</em>, <em>L. ridibundus</em>,
<em>L. novae-hollandii</em>) has been credited
to availability of food from wasteful human
garbage disposal (Murray and Carrick 1964;
Fordham 1968, 1970; Harris 1964; Harris and
Plumb 1965; Kadlec and Drury 1968; Brown
1967; Mills 1973; Vermeer 1963).</p>

<p>It is hard to dismiss the evidence pointing
to the impact of human activities on seabird
populations during the last 3 centuries. Yet it
would be misleading to assume that without
man's interference seabird populations would
have remained stable. Success in designing
programs of protection and population enhancement
must allow for the realities&mdash;that
seabird populations fluctuate inherently, and
that secular changes occur regularly in their
environment.</p>


<h5><em>Impact of Natural Events</em></h5>

<p>Some population changes appear to result
from sudden impacts; other changes are
gradual.</p>


<h6>Sudden Disasters</h6>

<p>Gromme (1927) reported windrows of dead
murres in the Unimak Pass and Alaska Peninsula;
die-offs of murres in winter storms in the
Atlantic and Arctic Oceans were reported by
Tuck (1960) and Dement'ev et al. (1968).</p>

<p>Recently some mass mortalities have been
associated with specific causes. Bailey and
Davenport (1972) reported that starvation
caused the die-off of common murres in the
southern Bering Sea&mdash;Bristol Bay area. Foul
weather, which apparently inhibited feeding
between 19 and 23 April 1970, culminated in
an intense storm. Similarly in late winter 1969
bad weather in the Irish Sea, combined with
strains of molt and perhaps contamination<span class="pagenum"><a name="Page_125" id="Page_125">[Pg 125]</a></span>
with industrial chemicals, seems to have contributed
to mass mortality of the same
species (called common guillemot in Britain;
Holdgate 1971). The seabird victims of this
event had metabolized their body fat and as a
result, polychlorinated biphenyls (PCB) and
other industrial chemicals passed into livers,
kidneys, and brains. Again, a storm at the end
of a period of stress seems to have been more
than the birds could tolerate.</p>

<p>A further example of a die-off of waterfowl
apparently brought on by starvation was
given by Barry (1968), who estimated that
about 100,000 king eiders <em>(Somateria spectabilis)</em>
died when they arrived before the ice
broke up in the Beaufort Sea in spring 1964.</p>

<p>Diseases have produced massive die-offs in
marine birds. Fowl cholera caused high mortality
in nesting common eiders <em>(Somateria
mollissima)</em> in the Gulf of St. Lawrence in
Quebec (Reed and Cousineau 1967) and in
Penobscot Bay, Maine, in the early 1960's
(H. Mendall, personal communication).
Poisoning from a "red tide" (a bloom of the
dinoflagellate <em>Gonyaulax tamerensis</em>) caused
a die-off of black ducks <em>(Anas rubripes)</em> and
herring gulls on the coast of New England in
1972. Similarly a die-off of shags <em>(Phalacrocorax
aristotelis)</em> on the east coast of England
was caused by a "red tide" (Coulson et al.
1968). During a period of 1 week 90% of the
shag nests on the Farne Islands in Northumberland
were deserted and about 80% of
the breeding population died.</p>


<h6>Gradual Declines</h6>

<p>When the new volcanic island of Bogoslov
emerged in the western Aleutians, Preble and
McAtee (1923) reported that it was colonized
by large numbers of pigeon guillemots
<em>(Cepphus columba)</em>, but in the following
decades the guillemots have steadily decreased
(G. J. Divoky, personal communication).
As a further example, the nesting population
of Atlantic puffins <em>(Fratercula arctica)</em>
in the Atlantic has declined over the past
several years, especially those nesting on the
Outer Hebrides (Flegg 1972; Harris 1976).</p>

<p>It is difficult to find seabird species whose
nesting grounds have not been affected by
humans but whose numbers have been censused.
The best illustrations of secular
changes in relatively constant habitats are
probably those available in the British Trust
for Ornithology's breeding censuses of songbirds.
Songbirds are short-lived and their
populations change on relatively short time
scales. The northwestern European landscape
has remained relatively constant for the last
75 years, yet there are observable decade-long
trends&mdash;for example, of willow warblers <em>(Phylloscopus
trochilus)</em> and dunnock <em>(Prunella
modularis)</em>. There are detailed data on population
changes in great tits <em>(Parus major)</em>
through the work of Kluyver (1951), Lack
(1964), and Perrins (1965).</p>


<h6>Effects Reflecting Environmental Change</h6>

<p>Nelson (1966) argued that the increase of
gannets in the North Atlantic during this century
has been related to increasing temperatures
rather than (as usually ascribed) to increased
food from fish damaged or escaped
during commercial fishing.</p>

<p>Ainley and Lewis (1974) described a particularly
interesting example of the effects of environmental
change on seabird populations.
The events begin with the decrease of seabirds
on the Farallon Islands off California as
a result of human depredations. Even after
fowling was made illegal, the populations of
murres, double-crested cormorants <em>(Phalacrocorax
auritus)</em>, and especially of tufted puffins
<em>(Lunda cirrhata)</em> and pigeon guillemots continued
to decline as a result of oil pollution.
During the last 3 decades the smaller species
of seabirds nesting on the Farallons, such as
rhinoceros auklets <em>(Cerorhinca monocerata)</em>,
have increased rapidly and the authors suggest
that their increase was abetted by an increase
in the small prey fish, northern anchovy
<em>(Engraulis mordax)</em>. One of course expects
predators to be affected by changes in
the abundance of their prey. During this same
period, larger species of seabirds such as
double-crested cormorants and tufted puffins
have failed to recover their numbers, and the
authors speculate that this failure is related
to a decrease of the larger prey fish, Pacific
sardine <em>(Sardinops caerulea)</em>.</p>

<p>A widely publicized impact of environmental
fluctuation upon seabird populations
is that of the northeast wind, El Ni&ntilde;o, off the
Peruvian coast. This wind pushes the upwell<span class="pagenum"><a name="Page_126" id="Page_126">[Pg 126]</a></span>ing
Humboldt Current water offshore and
causes mass mortality in the Peruvian anchovies
<em>(Engraulis ringens)</em> and, as a consequence,
a die-off among the millions of seabirds
such as Peruvian guanay cormorants
<em>(Phalacrocorax bougainvillii)</em> and Peruvian
boobies or piquero <em>(Sula variegata)</em> which feed
upon them (Murphy 1936).</p>


<h4>Theoretical
Considerations</h4>

<p>Can useful generalizations be drawn from
these observations on population changes?
Can a model be constructed of the forces
which drive population changes or of population-habitat
interactions which keep populations
from extinction? Some conflicting theories
and assumptions of population dynamics
are examined and discussed below.</p>


<h5><em>The Assumption of Population
Stability and of Closely Attuned
Density-dependent Mortality</em></h5>

<p>During the 5 decades before 1970, it was
widely accepted that most animal populations
were generally stable and saturated before the
arrival of the white man. Although a few field
biologists vigorously dissented, "establishment"
ecologists regarded fluctuations as a
departure from the norm, and as such, a
hazard to the population. Many theorists of
both evolution and ecology argued that adaptations
were required to damp fluctuations or
the fluctuations would become "random
walks" and the population would rapidly become
extinct. As a consequence, relatively all
theoretical models included stability as a central
assumption.</p>

<p>&#8226; The basic element of this theoretical complex
has been the Lotka-Volterra formula for
a logistic curve of population growth and stabilization.
According to this formula it has
been reasoned that by establishing the inherent
rate of increase of a population (i.e., its
average natality relative to mortality, or <em>r</em>)
and by measuring the carrying capacity of the
environment (which is the density of the population
at saturation, or <em>K</em>), one can predict the
maximally productive population size, and
maximum rate of production of new individuals
(or maximum sustained yield). These
assumptions have supplied the theoretical
framework for virtually all game management
and many fisheries practices.</p>

<p>Once stability was assumed, a mechanism
for maintaining stability was necessary. This
mechanism was found in an interaction between
the population and the environment,
called density-dependent mortality (Nicholson
1933). The impact of this feedback has
been assumed to cause the point of inflection
of the "sigmoid curve" and to regulate the
density "at equilibrium."</p>

<p>Populations growing in relatively isolated
or closed systems have been observed to follow
a sigmoid curve toward a steady state.
We have data on the growth of several Massachusetts
gull colonies which show this type of
short-period rapid increase followed by a long
sequence of shallow oscillations (Drury and
Nisbet 1972). But usually observations have
been terminated at about the time the population
passed through the point of inflection.</p>

<p>&#8226; Lack (1954) accepted the principles formulated
by Lotka-Volterra and hence viewed
Nicholson's (1933) density-dependence as logically
necessary. Lack (1948, 1954) argued that
reproductive effort (clutch size or litter size
times the number of broods) must be as large
as the parents can successfully raise to independence
because these biological characteristics
are directly subject to natural selection.
He argued that because reproductive potential
is excessive (Darwin 1859), mortality
must be density-dependent if a population is
to avoid fluctuations. The only adequately
density-dependent regulating process he accepted
was the population's response to its
food supply (Lack 1954). In fact, for many
years Lack rejected Kluyver and Tinbergen's
(1953) hypothesis that territory could act as a
control on population size in birds because, he
argued, territories were compressible and
therefore allowed wide fluctuations. To his
credit, however, Lack eventually acknowledged
this mistake.</p>

<p>The first defect in the concept of "carrying
capacity" is the idea that populations have
"mechanisms" or "institutions" (Wynne-Edwards
1959) by which the population is
kept stable at the carrying capacity in a
stable habitat.</p>

<p><span class="pagenum"><a name="Page_127" id="Page_127">[Pg 127]</a></span></p>

<p>The second defect in the concept of carrying
capacity is that it presupposes a stable environment.
During the early decades of the 20th
century most climatologists believed that a
departure from the norms of a regional climate
set processes in motion which would return
the climate to normal. During the last
decades, however, climatologists and
oceanographers have shown clearly that environments
are continuously in flux.</p>


<h5><em>An Attack on Density-dependent
Mortality</em></h5>

<p>Some theorists rejected the concept of
carrying capacity as soon as it was formulated.
Andrewartha and Birch (1954) predicted
fluctuations would be undamped by inherent
population mechanisms but rather
would be controlled by external forces indifferent
to the density. Their supporting data
were drawn from field studies of insects in
arid climates. Some of their ideas are directly
relevant to seabirds; for example, their assertion
that in many cases limits to carrying
capacity of the habitat are not set in a way responsive
to the density of the population. The
number of occupiable ledges on a seabird cliff
are fixed and when they are full no more birds
can breed there regardless of the amount of
food available. For another example, some
biological processes act in a way that reinforces
fluctuations. Predation can act in this
way in the relatively closed system of a seabird
colony; i.e., the smaller the prey population
the larger the percentage taken by the
predators. The importance of predation as a
selecting factor is shown by the adaptations
marine birds and waterfowl make to avoid it.
The fact that large colonies of seafowl are
usually concentrated on isolated, predator-free
islands is one obvious case (Lack 1966).</p>

<p>Although their ideas are useful in understanding
changes in many species, primarily
insect populations, the generality of Andrewartha
and Birch's (1954) hypothesis is
weakened because it conflicts with detailed
studies of seabirds which show that in many
cases local food resources do limit breeding
success. Ashmole (1963) showed this for tropical
terns, and Hunt (1972) for some colonies of
herring gulls on the New England coast.
Nettleship (1972), studying the effects of
herring gulls on Atlantic puffins, showed that
the effect of harassment and stealing food
from the parents was to reduce the amount of
food brought to the young and thus reproductive
success. In those parts of the colony
where gulls were numerous or where the
puffins were at a disadvantage in escaping
from gulls (i.e., flat rather than steep slopes)
the reproductive success of puffins was significantly
lower than in areas away from the
gulls.</p>

<p>The literal application of Andrewartha and
Birch's general ideas also conflicts with observations
on subtle adaptations some waterfowl
have made to counter predation.</p>

<p>Barry (1967) described the density-avoiding
adaptations of arctic-nesting geese to evade
predation&mdash;specifically by foxes. Black brant
<em>(Branta nigricans)</em> nest on low coastal or delta
islands seeking to escape by remoteness.
Snow geese <em>(Chen caerulescens)</em> are colonial
on large, flat areas, seeking protection in numbers.
White-fronted geese <em>(Anser albifrons)</em>
are solitary nesters on inland swamps, seeking
to be "over-dispersed" among scrub
willow.</p>

<p>Common eiders, black scoters <em>(Melanitta
nigra)</em>, tufted ducks <em>(Aythya fuligula)</em>, and
other ducks select gull colonies as nesting
habitat. Although there is little doubt that
the ducks choose gull colonies for nesting,
there is some doubt as to the reasons. Finnish
biologists (summarized by Bergman 1957;
Hild&eacute;n 1965) have concluded generally that
gulls protect the duck nests from predation
by hooded crows <em>(Corvus corone)</em>.</p>


<h5><em>The Assumption that Fluctuations
Are Generally Present</em></h5>

<p>Recently theorists have built models based
on assumptions that fluctuations are a
general characteristic of population dynamics,
such as Gilpin's (1975) model describing
multi-phased oscillations. He took account
of the fact that fluctuations (and models) become
more complex as more species and nonlinear
effects are included. May and Leonard
(1975) emphasized that the effect of nonlinearities
is to make it impossible to speak
even in principle of the equilibrium point of a<span class="pagenum"><a name="Page_128" id="Page_128">[Pg 128]</a></span>
community. They pointed out that even
though the model is deterministic (i.e., assumes
that the system will come to equilibrium)
the oscillations are so complex that they
may appear to be random, and it may be a
very long time before the system returns to a
position near its starting point. "On the other
hand a truly random ecological system could
always be fitted by a suitably ingenious limit
cycle. This suggests that ecological analysis
which does not consider component processes
must be viewed with great suspicion" (Gilpin
1975). May and Leonard (1975) and Gilpin are
both making a familiar point&mdash;that neither
the logic nor the interactions described in a
formula will describe biological reality unless
the assumptions are correct. They are also
making a different point&mdash;that an ingenious
mathematician can create a formula to describe
almost any operation (whether its
workings are systematic or random), and the
formula may seem to work.</p>

<p>Gilpin's moral is that one cannot learn very
much that is helpful by studying fluctuations
as such. One must study the factors controlling
populations. This is a very old idea.</p>

<p>It would appear that defining carrying
capacity and inherent rate of increase will not
be very instructive in managing seabird populations
other than in speculating upon what
might be ideal upper limits. It can also encourage
the musty sophistry that when a population
increases beyond this abstract carrying
capacity it "needs" to be hunted to prevent
overcropping resources and damage to itself
through a population decline. But we will not
have the time to carry out detailed studies of
life histories seeking for critical population-habitat
interactions over several fluctuations
for each species involved in a disaster before
designing programs to help seabird populations
to build up their numbers.</p>


<h4>General Characteristics of
Marine Birds and Waterfowl</h4>

<p>Because general theory does not seem to
work and because detailed studies take too
much time, I conclude that it is necessary to
identify certain general principles upon which
to base applied programs. These categories of
knowledge include: (1) how vulnerable certain
categories of seabirds, waterfowl, and shorebirds
are to specific types of disasters, (2) how
quickly their numbers build up after they
have been reduced, and (3) at what stages we
can help them best (i.e., at the breeding
grounds, at the winter gathering grounds, or
on migration). I believe that we already know
enough to design effective programs and to
begin work. To this end some characteristics
of seabirds are identified which determine the
population structures and ways in which their
numbers respond to changes in the environment.</p>


<h5><em>Habitat</em></h5>

<p>Although the shallow oceans, islands, and
seashores are among the most permanent features
of the earth in general, the details of
their numbers and distribution change
rapidly. Sandy shores are obviously being reworked
even in the short span of a single lifetime.
Distribution of islands and the sediment
load, extent, and strengths of currents vary
constantly in space and change with time.</p>

<p>The food that seabirds use is patchy and
subject to both short-and long-term fluctuations
in numbers and shifts in geography.
Suitable breeding habitat is scattered, and in
many places where oceanic conditions provide
a good food supply there are no nesting sites.
Consequently, seabirds aggregate in colonies,
often dense, and the colonies are clumped for
geographical as well as biological reasons.</p>

<p>Lack (1966) discussed some general features
of how the breeding adaptations of seabirds
are adjusted to the distances the birds must
go to find food. The species which feed close to
the nest characteristically establish isolated
territories or nest in small groups, and they
accept many different kinds of nesting substrate.
Their clutch sizes are large, individuals
move nesting sites readily, and their young
grow rapidly compared to the species which
feed far at sea. Species which feed far at sea
aggregate in large colonies. These species are
often rigid in their requirements for suitable
nesting sites, their clutches are usually
limited to one egg per season, their young
grow slowly, and there seems to be strong attachment
to traditional colony sites.</p>

<p><span class="pagenum"><a name="Page_129" id="Page_129">[Pg 129]</a></span></p>


<h5><em>Breeding</em></h5>

<p>Ashmole (1963) suggested that the clutch
size of some oceanic birds is small and colonies
occupy only part of the available habitat
because food resources within efficient commuting
distance of the breeding site are
limited. We can see this effect in the usual failure
of common terns to raise a third chick,
even in the colonies that are surrounded by
favorable habitat (Nisbet 1973). Herring gulls
whose colonies are close to sources of human
refuse raise more young than do those whose
colonies are at some distance (Drury 1963;
Kadlec and Drury 1968; Hunt 1972).</p>

<p>Ashmole (1963) suggested that during the
course of the breeding season the birds exhaust
the available food supply. The validity
of this suggestion is reflected in the long distances
some species (petrels, boobies, murres,
dovekies) go for food to feed their young. One
would therefore expect that early nesting
pairs would be more successful, and this
seems to be the case in herring gulls (Nisbet
and Drury 1972), kittiwakes (Coulson 1966),
and red-billed gulls, <em>Larus novaehollandiae</em>
(Mills 1973).</p>

<p>If food is in short supply and parents have
to seek over a wide area for food so that they
can bring back only a little food at long time
intervals, one would expect these birds to
have a small clutch and their young to grow
slowly, as is the case. One would also expect
seabird colonies situated near oceanic currents
to be larger and more successful because
food is continuously renewed. Conversely, one
would expect colonies next to still waters to
be smaller and less successful.</p>

<p>The small clutch size of seabirds means that
when a population has been reduced, it will
grow slowly toward its former abundance.
The growth rates of seabird populations on
the New England coast since their release
from human predation reflects this. Species
such as black guillemots with only two eggs
per clutch and herring gulls with three eggs
per clutch have increased more slowly than
have the populations of common eiders or
double-crested cormorants both with three to
six eggs per clutch (Drury 1973).</p>

<p>If the species that nest in colonies show a
high degree of site tenacity, they are not likely
to reestablish a colony after it has been eliminated.
An exception to this is the food subsidy
provided by man, which seems to have
been important in creating a nonbreeding
population of herring gulls large enough to
form a "critical mass" for the formation of a
new gullery.</p>


<h5><em>Age Structure</em></h5>

<p>Because the main element of population
size&mdash;the number of breeding adults&mdash;is
limited by the number of breeding colonies
and the food available to those colonies, one
assumes that the total numbers of seabirds is
much less than could be supported by the
larger areas of productive oceans. Hence one
suspects that there is lessened competition
for food outside the breeding season and that
lack of competition for food is a major reason
for seabirds being long-lived, often to extremes
little suspected until recently. Mortalities
of 10-12% per year are common, and some
as low as 4% (wandering albatross, <em>Diomedea
exulans</em>; Tickell 1968) have been recorded.</p>

<p>In contrast, songbirds with large clutches,
such as the titmice studied by Kluyver (1951),
produce a large number of young with whom
they and other adults must compete for food
during the winter period of food shortage. Because
the titmice are permanent residents,
they occupy all of the available habitat
throughout the year. Hence titmice suffer intense
intraspecific competition, which
shortens the survival of adults. Kluyver's experiments
(1966) with nest boxes used by a
closed population of great tits on Vlieland,
The Netherlands, showed that by artificially
reducing clutch size the survival of adults was
increased.</p>

<p>Similar competition for the few territories
available on marshes and consequent shortened
life expectancy, can be expected in
waterfowl with large broods. The effect
should be less marked for geese with smaller
clutches that nest in less confined habitats.</p>

<p>The long life span of seabirds means that a
population will have a large component of
older age categories; this characteristic has
several implications:</p>

<p>&#8226; It means that the population can survive
years of reproductive failure without the observable
immediate effects that would be<span class="pagenum"><a name="Page_130" id="Page_130">[Pg 130]</a></span>
manifest in titmice, grouse, or rabbits. Near
failure of reproduction during a breeding season
among arctic seabirds at Bear Island was
reported by Bertram et al. (1934). Many similar
observations have been made since then:
Pitelka et al. (1955) reported such a case
among skuas and gulls at Point Barrow,
Drury (1961) for greater snow geese <em>(Chen
cerulescens atlantica)</em> at Bylot Island, Jones
(1970) for black brant gathering at Isambek
Lagoon on the Alaska Peninsula, and D. A.
Snarski (personal communication) for kittiwakes
at Cook Inlet. Reproductive failure can
sometimes be chronic, as observed by Nisbet
(1972) for terns at Cape Cod, Massachusetts,
or by Drury (1963) and Hunt (1972) for
herring gulls on the outer islands on the coast
of Maine.</p>

<p>When reproductive failure becomes chronic
as observed on peregrine falcons <em>(Falco peregrinus)</em>
by Hickey (1969) and in ospreys
<em>(Pandion haliaetus)</em> by Ames and Mersereau
(1964), the population of adults may hold on
for a number of years without evident decline.
Damage to the structure of the whole population
may be serious before any numerical results
are evident.</p>

<p>&#8226; Although there may not be intensive competition
for food in the habitat away from
breeding colonies, there is intense competition
for food and breeding sites at and around the
colonies. Hence age and previous experience
in seabirds assume importance in establishing
territory and in breeding success. Associated
with this is the tendency for immature birds
to delay breeding until they are several years
old and for the immatures to remain on feeding
grounds at some distance from the colonies.
In some cases young birds may "hang
around" breeding colonies and even feed some
of the young. When young birds do first breed
they usually lay smaller clutches and raise
fewer young than do older birds. The importance
of age and experience upon breeding
success has been well documented for kittiwakes
(Coulson 1966) and red-billed gulls
(Mills 1973).</p>

<p>The fundamental biological importance of
this delayed maturity seems to be emphasized
by the persistence for several years of immature
plumages, so clearly identifiable that
even a human observer can recognize the age
of an individual. One assumes such an evident
feature must have adaptive significance.</p>


<h5><em>Wintering Grounds</em></h5>

<p>When colonial nesting seabirds leave their
breeding islands for their wintering grounds,
their identification with that island is lost as
far as population effects are concerned, because
birds from many colonies mingle on the
wintering grounds. Major mortality takes
place on the wintering grounds and must
therefore act on the species population as a
whole rather than differentially on individuals
associated with especially dense colonies.
Such a direct relation between colony density
and mortality would be necessary for density-dependent
mortality to regulate the number
of birds on a breeding colony. Conversely, one
cannot expect that all colonies will decrease
equally because mortality should be equally
distributed if all the population gathers on a
common wintering ground. Thus density-dependence
acts only in a very general way upon
the sum of animals considered as an abstract
entity&mdash;the population.</p>

<p>In fact, on the wintering grounds, as shown
by a graph of numbers of gulls reported on
Christmas Counts on Cape Cod, Massachusetts
(Kadlec and Drury 1968), herring gulls
are very responsive to local conditions and
move several tens of miles to gather at favorable
feeding sites. An aerial survey of the
gulls on the East Coast of the United States
(Kadlec and Drury 1968) showed that more
than half of the gulls were gathered near
major food sources in large metropolitan districts.
Most of the remainder were gathered
near small fishing ports. Very few were
scattered along the shoreline in what one assumes
is the traditional gull habitat. Later
analyses of the relation between the distribution
of banding recoveries of birds in their
first winter and the distribution of immatures
as found on this winter census (Drury and
Nisbet 1972) suggested that proportionately
more first-year gulls died in those areas where
the birds were sparsely distributed than died
in the crowded metropolitan areas.</p>

<p>These results suggest both that there is not
a direct feedback between reproductive rate
and mortality, and that mortality may even
be inversely density-dependent on wintering
grounds. This last runs counter to traditional<span class="pagenum"><a name="Page_131" id="Page_131">[Pg 131]</a></span>
ecological ideas that density causes a change
in mortality rate. The idea that individuals
gather where "living is easy" and mortalities
are low is consistent with the theory of natural
selection. One would not expect the food of
the gulls to be evenly distributed, and one
would expect individuals to move away from
areas where food is scarce and mortality is
high.</p>


<h5><em>Differences in Breeding
Success Between Colonies</em></h5>

<p>Breeding success has been shown to vary
among individual pairs of gulls (Drost et al.
1961). Certain groups of individuals nesting in
patches within a single colony have greater
breeding success than do others (Coulson
1968; Drury and Nisbet, in preparation). Differences
in breeding success also occur between
colonies (Frazer-Darling 1938; Kadlec
and Drury 1968; Drury and Nisbet 1972).
Some colonies reproduce consistently better
than others&mdash;for example, the gull colonies
close to fishing ports and metropolitan areas.
Other colonies produce consistently fewer
young, such as the colonies on the outer islands
in the Gulf of Maine (Drury 1963; Kadlec
and Drury 1968; Hunt 1972). The populations
of successful colonies grow while the
numbers of unsuccessful colonies decline,
even during a period of general population increase
(Kadlec and Drury 1968).</p>

<p>The difference between success and failure,
growth and decline, appears to lie in the food
available. Colonies increase where breeding
success is high and decrease where breeding
success is low. One important reason seems to
be that adult gulls may move to a more productive
colony even after they have nested
with another colony (Drury and Nisbet 1972;
Kadlec 1971). Such adaptations can be viewed
as adjustments by which individuals meet the
requirements of an environment in which the
availability of food and other necessities is
patchy and shifting.</p>


<h5><em>Dispersal</em></h5>

<p>In general terms, the willingness of some individuals
to disperse while the majority of
individuals remain loyal to a colony can be
considered a major mechanism of population
maintenance. If conditions deteriorate seriously
at one place so that the local populations
decline or disappear, dispersal from
other centers can be expected to repopulate
the area as soon as local conditions again become
suitable. This subject has been treated
in more detail by Drury and Nisbet (1972) and
Drury (1974<em>b</em>).</p>

<p>Occupation of new, or return to former,
nesting sites has been recorded in detail for
fulmars <em>(Fulmaris glacialis)</em> by Fisher (1952)
and for herring gulls by Kadlec and Drury
(1968). Dispersal is also known for waterfowl.
Hansen and Nelson (1957) reported that of
some 8,000 brant banded in midsummer on
the Yukon delta 8 were recovered in northern
Siberia and 28 in northern Alaska and arctic
Canada. They suspected that pairing on the
wintering grounds was responsible for the
change in breeding areas, a change that would
not be expected among other North American
species of geese. Similarly, wide dispersal
seems to occur in pintails <em>(Anas acuta)</em>,
mallards <em>(Anas platyrhynchos)</em>, and wood
ducks <em>(Aix sponsa)</em>.</p>

<p>The general tendency for some individuals
to disperse and the frequency of "extra
limital" breeding attempts is especially well
established in the Bering Sea region, in part
at least because vagrants from Siberia or
North America are readily identified as such.
In the Aleutian Islands, Emison et al. (1971)
and Byrd et al. (1974) have enumerated the
nesting vagrants. For the Pribilof Islands,
Kenyon and Phillips (1965), Sladen (1966), and
Thompson and DeLong (1969) have recorded
the repeated appearance of birds of Siberian
distribution, and Fay and Cade (1959) and
Sealy et al. (1971) did the same for St.
Lawrence Island.</p>

<p>One can conclude that a few individuals are
constantly trying to settle in new geographical
areas. As climatic and habitat conditions
change, some populations are able to become
established; for example, southern species
such as mockingbirds <em>(Mimus polyglottus)</em>,
cardinals <em>(Cardinalis cardinalis)</em>, and tufted
titmice <em>(Parus bicolor)</em> have settled in southeastern
New England during the last 2
decades. These southern species have received
much publicity. But at the same time, a less
publicized dispersal of white-throated spar<span class="pagenum"><a name="Page_132" id="Page_132">[Pg 132]</a></span>rows
<em>(Zonotrichia albicollis)</em>, hermit thrushes
<em>(Catharus guttatus)</em>, and dark-eyed juncos
<em>(Junco hyemalis)</em> has resulted in new nesting
records of more northerly species, also in
southeastern New England.</p>

<p>The ability (or lack of ability) of some organisms
to expand their ranges over time has
been a subject of consideration for a number
of years by plant and animal geographers. An
important botanical paper on this subject in
the Bering Sea region was presented by
Hult&eacute;n (1937), who analyzed the ranges of
plants of the area of Kamchatka, eastern Siberia,
Alaska, and northwest Canada, showing
that diverse floras occur in some restricted
geographic areas. He called these
areas "refugia," and postulated that many
species had survived Pleistocene glaciations
in them because these refugia remained ice-free.
He, like Fernald (1925), was puzzled as to
why only certain species had been able to expand
their ranges outward from these "areas
of persistence," while other apparently more
"conservative" species were unable to do so.
Similarly, there appear to be conservative endemic
bird species of the Bering Sea region:
the extinct Commander Islands cormorant
<em>(Phalacrocorax perspicillatus)</em>, Steller's eider
<em>(Polysticta stelleri)</em>, spectacled eider <em>(Lampronetta
fisheri)</em>, emperor goose <em>(Philacte canagica)</em>,
whiskered auklet <em>(Aethia pygmaea)</em>,
least auklet <em>(A. pusilla)</em>, parakeet auklet <em>(Cyclorrhynchus
psittacula)</em>, Aleutian tern <em>(Sterna
aleutica)</em>, red-legged kittiwake <em>(Rissa brevirostris)</em>,
bristle-thighed curlew <em>(Numenius
tahitiensis)</em>, long-billed dowitcher <em>(Limnodromus
scolopaceus)</em>, surfbird <em>(Aphriza virgata)</em>,
black turnstone <em>(Arenaria melanocephala)</em>,
rock sandpiper <em>(Calidris ptilocnemis)</em>,
and western sandpiper <em>(C. mauri)</em>.</p>

<p>The ranges of horned puffins <em>(Fratercula
corniculata)</em>, Kittlitz's murrelet <em>(Brachyramphus
brevirostris)</em> and, perhaps, crested
auklet <em>(Aethia cristatella)</em> suggest that some
species of "Beringian" seabirds have expanded
their ranges from Hult&eacute;n's (1937)
"refugia."</p>


<h5><em>Dispersal and Regional Persistence
of Marginal Populations</em></h5>

<p>The presence of several sub-elements of a
species population and, therefore, the opportunity
for dispersion among alternative breeding
sites may be an important factor in the regional
persistence of a species on the margin
of its range, as illustrated by the history of
laughing gulls <em>(Larus atricilla)</em> in New
England.</p>

<p>Between 1875 and 1900 there were fewer
than 50 laughing gulls in Massachusetts
(Mackay 1893) and about 35 in Maine (Norton
1924). In Massachusetts the laughing gulls all
settled on one large island, Muskeget, where
by 1940 there were about 20,000 pairs (Noble
and W&uuml;rm 1943). Meanwhile, in Maine the
population had been disturbed by sheep and
men and had shifted about among seven islands.
The Maine population grew to only
about 350 pairs by 1940 (Palmer 1949).</p>

<p>The laughing gull population in both States
has decreased since 1940. In Massachusetts,
where all pairs occupied one island, the population
had fallen to about 250 pairs by 1972,
but the Maine population, still divided into
five colonies, stabilized at 250 pairs (i.e., the
same as instead of only 1% of the Massachusetts
population).</p>


<h4>Use of General Principles in
Solving Conservation Problems</h4>

<p>Game biologists have successfully maintained
the populations of hunted animals by
using a number of classical principles of game
management. They have controlled mortality
by regulating kill and have increased standing
stock by improving habitat on a local scale.
This seems to have worked in species which
are short-lived, have large clutch sizes or
litters, and which occupy mosaics of highly
productive "successional habitat." Seabirds,
however, contrast with these species in a number
of important biological characteristics.
They have small clutches, postpone breeding
until they are several years old, and are subject
to periodic or chronic reproductive failures.
Therefore, their populations are skewed
toward older animals and replacement of lost
individuals is slow. Many seabirds, like some
geese, have a high level of site tenacity and
thus may resist recolonization or fail in the attempt
to recolonize a breeding site once eliminated
from it. In those species studied it appears
that the breeding birds at a small per<span class="pagenum"><a name="Page_133" id="Page_133">[Pg 133]</a></span>centage
of colonies are responsible for a large
proportion of the annual crop of young. It is
probably dangerous, therefore, to risk either
damage to or elimination of well-established
colonies.</p>

<p>Studies of kittiwakes by Coulson and White
(1958, 1961), sooty terns <em>(Sterna fuscata)</em> by
Ashmole (1963) and Harrington (1974), Atlantic
puffins by Nettleship (1972), and
Cassin's auklets <em>(Ptychoramphus aleutica)</em> by
Manuwal (1974), and the practice of eider
"farming" in Iceland indicate that the number
of available territories or breeding sites
may limit the size of a population and that
populations can be increased by increasing
the number of sites available. This suggests
one way in which direct steps can be taken to
encourage the numbers of breeding seabirds.
Other studies indicate that seabirds will move
into synthetic habitat such as created by the
window ledges on buildings (Coulson and
White 1958) or the islands created by dumping
spoil from channel dredging operations
(Buckley and Buckley 1971, 1975; Soots and
Parnell 1975).</p>

<p>Most generalizations of population biology
have been derived from the study of insects,
songbirds, or game species. It seems inadvisable
to assume that those principles will apply
to seabirds without modification. For
example, predation by gulls and ravens may
have a disastrous effect on a seabird colony at
low colony density but have progressively less
impact as the colony size and density increase.
Fox predation may have important effects
over most ranges of prey density because
the presence of foxes has important
psychological effects.</p>

<p>The habitats of seabirds include elements in
which birds are widely dispersed (feeding
areas) and others in which birds are crowded
and narrowly localized (nesting sites). Thus effective
programs of conservation should include
guarantees that a number of colony
sites be available in as widely dispersed a pattern
as possible. Each productive feeding
ground should, if possible, have several
colony sites available.</p>

<p>We have argued elsewhere (Drury and Nisbet
1972; Drury 1974<em>a</em>) that one of the chief defenses
any population has against extinction
is the combination of being divided into a
number of population centers with having
some movement of individuals between the
centers, but not too much. Because it is highly
improbable that a single catastrophe will affect
more than a part of a species' range at
any time, the more numerous and widely
scattered the partially independent segments
of a population are, the better the species is insured
against extinction. This, of course, suggests
that the size of each colony may be less
important for long-term survival than is the
total number of colonies.</p>

<p>One intuitively concludes that "conservative"
species, such as those endemic to the
Bering Sea region (whose dispersal and colonizing
mechanisms seem to be poorly developed),
are especially vulnerable to the effects
of local population crashes. These
"local" species therefore deserve special
consideration.</p>

<p>I would like to emphasize two points to be
included in designing a "management"
program:</p>

<p>&#8226; It seems that the most promising management
techniques will be built upon ensuring
the health of colonies and the associated feeding
areas at which reproductive success is
high enough to "export" young. Thus it is
useful to identify those colonies which are exporting
young and to give special care to their
preservation. As populations of prey species
change locally, so will the success of the local
nesting birds. A colony which is thriving at
one time may be barely maintaining itself at
another (Ainley and Lewis 1974), or it may decrease,
as in the case of "guano birds" during
El Ni&ntilde;o years in the Peru Current.</p>

<p>&#8226; Because centers of abundance of marine
birds shift (Fisher 1952; Drury 1963, 1974<em>a</em>),
it will be prudent to plan for large areas and
over long periods of time. Harrison Lewis, a
pioneer in seabird management in eastern
Canada, said (personal communication) that
just as soon as he got approval of a new seabird
sanctuary through the long corridors of
the distant government bureaucracies in Ottawa,
the birds would move to a new island
and he had to start the process all over again.</p>

<p>The objective is to maintain a variety of
colony sites for populations to move among as
local patterns of productivity in the shallow
sea shift.</p>

<p><span class="pagenum"><a name="Page_134" id="Page_134">[Pg 134]</a></span></p>


<h4>Goals for Research on
Population Dynamics of Seabirds
for Purposes of Conservation</h4>

<p>1. To learn the distribution and relative importance
of seabird colonies, the number of
pairs nesting and nonbreeding individuals at
each colony, and the timing of breeding activities
for each geographical region. The most
important step toward conserving marine
birds is to get public ownership and protection
for their breeding grounds.</p>

<p>2. To understand the life cycle of key
species. Three needs are clear:</p>

<p>a. To identify key species whose biological
characteristics can conveniently be studied
and measured. Studies of these species may
be useful in monitoring the "health" of seabird
breeding areas.</p>

<p>If it is established that the reproductive
success of certain species varies similarly in
response to changes in their marine habitat
(such as black-legged kittiwakes and horned
puffins), one could use key species (black-legged
kittiwake) to assess the performance of
those species in a colony whose breeding success
is difficult to measure (horned puffin).</p>

<p>b. To develop efficient and practical ways
of censusing and measuring productivity of
crevice-, scree-, and hole-nesting species such
as puffins and auklets.</p>

<p>c. To establish annual differences in reproductive
success and mortality rates by age
classes of the key species, and from these to
identify rates of population turnover so as to
be able to predict the effects of mass mortalities.</p>

<p>3. To learn enough about the differences in
behavior and productivity among colonies to
establish which colonies produce surplus
young and which have low productivity. At
first, maximum efforts for conservation
should be concentrated at those sites which
produce surplus young.</p>

<p>4. To learn about colonial behavior. Two
needs are apparent:</p>

<p>a. To know enough about the lives of individually
marked birds of known age so as to
be able to infer the behavior of population elements
at all stages of their life cycle.</p>

<p>b. To know enough about the lives of subadult
birds to understand what proportion of
subadults visit and become established at
breeding sites, why the subadults visit the
breeding sites and what effect their presence
has on the territories and breeding success of
their neighbors and biological relatives.</p>

<p>5. To know enough about places where seabirds,
waterfowl, and shorebirds gather on migration
and during the winter to identify
those areas which need special protection
from effects of economic development.</p>

<p>a. It is important to determine the areas
where marine birds gather at sea when they
are away from their breeding grounds. What
factors of habitat and food supply make certain
places preferable to others? What is the
relation between gathering grounds and
underwater topography (banks and edges of
the continental shelf)? What are the seasonal
and annual differences in preferred gathering
grounds? What special hazards exist, such as
unusual extent of sea ice or exceptional
storms?</p>

<p>b. It is important to plot coastal areas
where waterfowl and shorebirds gather on migration,
for molting, and during the winter.
Which open leads in the ice and patches of
open water at the mouths of rivers are of
especial importance in spring? What shorelines
and beaches act as "leading lines" during
migration? Which capes and points result
in concentrated overflights of migrating
waterfowl, and hence are locations of unusually
high kills by hunters? What wetlands,
bogs, coastal ponds, lakes, and lagoons are
used as gathering grounds and to what extent
do waterfowl and shorebirds exchange between
gathering grounds? How much redundancy
of wetlands is needed to make the wetlands
system maximally productive for waterfowl
and shorebirds?</p>

<p>Answers to these questions will identify
which geographic areas deserve special protection
during development. The answers will
also identify the kinds of influences which
might lower the contribution of each critical
area to the survival of seabirds, waterfowl,
and shorebirds. Areas identified as important
under these categories must be included in
policy decisions related to land-use planning
and management.</p>

<p>6. To learn more about the effects of varying
quantities of food on breeding behavior
and performance:</p>

<p><span class="pagenum"><a name="Page_135" id="Page_135">[Pg 135]</a></span></p>

<p>a. What are the effects of food abundance
in early spring on date of laying, clutch size,
and egg size?</p>

<p>b. What effects do storms have at different
stages of the reproductive schedule?</p>

<p>c. What effects do quantitative and qualitative
(species composition of prey) changes in
food supply have on the survival of chicks?</p>

<p>d. What are the similarities and differences
between what parents eat and what
they feed their chicks?</p>

<p>Although this is important basic biological
knowledge, it contributes little to conservation
efforts because food differences result
from changes in the ocean over which humans
can have little effect.</p>

<p>7. To learn more about prey species and
their availability to marine birds:</p>

<p>a. To know more about the breeding
areas, reproductive rates, growth rates, and
routes of dispersal of the major prey food
species. In most areas a few species of teleost
fish (e.g., <em>Ammodytes</em>) or Crustacea (e.g.,
copepods, euphausids, mycids, or amphipods)
make up most of the food of marine birds. Yet,
the barest minimum is known about the biology
of such species. A good first estimate of
the "condition" of the marine environment
can probably be made by measuring reproductive
rates and growth rates of these key prey
species. Hence an efficient (though indirect)
way to measure those rates may be by monitoring
reproduction of birds.</p>

<p>b. To know more about the density and
distribution of key prey items season by season,
and to learn more about the relation of
their abundance and distribution to their
availability to birds, as B&eacute;dard (1969) showed
for <em>Calanus</em> to least auklets, and <em>Thysanoessa</em>
to crested auklets.</p>

<p>There are some indications that the population
size of prey items can vary widely without
having a marked effect on the numbers of
their predators. Does commercial fishing for
the large, predatory fish have a measurable effect
on the food available to marine fish? Do
the large pollock and salmon fisheries (high
seas) make zooplankton available to smaller
alcids? Do marine birds affect a fishery?</p>

<p>c. To know more about the oceanography
of continental shelf waters, more specifically
the waters between 6 and 60 m deep. The shallow
waters of continental shelves are some of
the most productive of sea waters, but are
among the least studied. Although some
species (black-legged kittiwakes, tufted
puffins, and fulmars) move into deep waters,
many species of marine birds of northern
waters gather in large numbers on preferred
feeding grounds at or near the edges of continental
shelves during their winter season
(Fisher 1952; Tuck 1960).</p>

<p>8. To know more about the potential effects
of proposed developments on seabirds and
waterfowl.</p>

<p>a. To prepare models which will predict
probabilities of contamination of breeding
and feeding areas (summer, winter, and during
migration) using existing knowledge of</p>

<p>(1) areas of proposed mineral development;</p>

<p>(2) areas that will be influenced by secondary
development such as dredging new
harbors, laying subsurface pipelines;</p>

<p>(3) tidal and oceanic currents;</p>

<p>(4) numbers of marine birds or waterfowl
using specific geographic areas and habitats
(e.g., waters below nesting cliffs, feeding
grounds, wintering grounds, and gatherings
during migration);</p>

<p>(5) the distribution and patchiness of
habitats (i.e., the redundancy among and
within habitats and the degree to which populations
exchange between alternative
habitats);</p>

<p>(6) the biological importance of species
in local ecosystems (Are they predators whose
effects increase diversity?);</p>

<p>(7) the human importance of the species
(Are they endemics? Do they have unusual
"charisma" for the public?);</p>

<p>(8) the vulnerability of the species (Is its
distribution restricted? Is it subject to oil pollution?
Are their preferred grounds near areas
of high development potential?);</p>

<p>(9) the types of biological effects (e.g.,
oil contamination of plumage, PCB contamination
of food chains); and</p>

<p>(10) whether the potential impacts are
reversible or irreversible and to what degree.</p>

<p>b. To understand more of the effects of
hunting on the behavior of marine birds and
waterfowl on their breeding grounds, and to
assess the effects on breeding performance of
changes in behavior which result from human
activities (such as hunting or studying the<span class="pagenum"><a name="Page_136" id="Page_136">[Pg 136]</a></span>
birds).</p>

<p>c. To understand the effects of the presence
of predators (whether introduced or native)
on breeding colonies in order to assess
the importance of removing the predators or
preventing their access to breeding grounds.</p>


<h4>The Relation of the Products
of Biological Research to
Programs for Conservation of
Marine Bird Resources</h4>

<p>Although peaceful coexistence of wildlife
populations and economic development are
here assumed to be practical, some new social
institutions are needed to control damaging
activities of people during economic development.
Human activities and industrial products
which damage wildlife or their habitat
must be identified, as must the space and resources
which wildlife require for survival and
health.</p>

<p>1. What seabird cliffs, islands, lagoons, wetlands,
river mouths, and other habitat features
are of first importance for breeding or
for maintaining the populations? Some small
areas of habitat are critical for the survival of
some species during periods of stress. Those
habitats need official recognition. Steps are
needed to ensure that the habitats are
maintained.</p>

<p>2. What physical expressions of economic
development are of little, modest, or serious
impact on wildlife and its habitat? These activities
and constructions include harbors,
storage sites, transshipment facilities, roads,
pipelines, summer camps, and suburban or
vacation developments.</p>

<p>3. What kinds of human activities will disturb,
damage, or change the behavior or accessibility
of wildlife? Many activities of one
group of people have secondary effects which
affect the enjoyment of resources for other
groups. These include</p>

<p>a. gill netting for salmon, which may kill
large numbers of murres and diving ducks;</p>

<p>b. release of predators on seabird nesting
islands, which may kill adults or inhibit their
feeding their young;</p>

<p>c. free running of pets (such as dogs and
cats) over wetlands or wildlife habitat, because
pets are predators and harass the wildlife
which may be feeding;</p>

<p>d. flights of aircraft, especially helicopters,
near or over seabird cliffs because
such flights may cause serious damage to
eggs and young;</p>

<p>e. hunting, because the game becomes
timid and flees from those who might enjoy
watching wildlife;</p>

<p>f. snow machines, because their presence
is disagreeable to many and they provide easy
access by which disturbing activities may
reach into areas where wildlife would otherwise
be undisturbed.</p>

<p>4. What limitations or alterations are
needed in the existing legal institutions, such
as the Marine Mammals Protection Act, the
instruments implementing native land claims,
the process of Alaska State lands withdrawal,
the conditions for leasing State and Federal
lands for development of mineral resources,
and traditional rights of private property? All
of these legal institutions are relevant to
problems of wildlife survival and restoration,
and within most of these institutions there
exist conflicts between rights and benefits of
special political interests and the husbanding
of renewable common property resources.</p>

<p>Experience in Europe and in New England
suggests that if reasonable limitations are set
on human activities and that if adequate
money charge is made against those who
profit by economic development to defray full
social costs, wildlife can continue to do well.
In most cases where damage has occurred it is
because those who administer the public institutions
have failed to include consideration of
the common property resources.</p>


<h4>References</h4>

<div class="hangindent">

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<p>Ames, P. L., and G. S. Mersereau. 1964. Some factors
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<p>Ashmole, N. P. 1963. The regulation of numbers of
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<p>B&eacute;dard, J. 1969. Feeding of the least, crested, and
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<p>Brown, R. G. B. 1967. Breeding success and population
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<p>Buckley, F. G., and P. A. Buckley. 1971. The breeding
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<p>Buckley, P. A., and F. G. Buckley. 1975. The significance
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<p>Byrd, G. V., D. D. Gibson, and D. L. Johnson. 1974.
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<p>Coulson, J. C. 1968. Differences in the quality of
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<p>Coulson, J. C. 1974. Kittiwakes <em>Rissa tridactyla</em>.
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<p>Coulson, J. C., G. A. Potts, I. R. Deans, and S. M.
Fraser. 1968. Exceptional mortality of shags and
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<p>Coulson, J. C., and E. White. 1958. The effect of age
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<p>Coulson, J. C., and E. White. 1961. An analysis of
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<p>Darwin, C. 1859. On the origin of species by means
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<div class="chapter"></div>
<hr class="chap" />




<h3><a name="Time-Energy_Use_and_Life_History" id="Time-Energy_Use_and_Life_History">Time-Energy Use and Life History
Strategies of Northern Seabirds</a></h3>


<p class="center">by</p>

<p class="center">Erica H. Dunn</p>

<p class="center">Long Point Bird Observatory</p>

<p class="center">Point Rowan, Ontario, Canada
</p>


<h4>Abstract</h4>

<div class="blockquot">

<p>Time and energy budgets can be compared among species of birds with very
different ecology as a way of summarizing differences and as an approach to
determining selective pressures on each species. This paper reviews time-energy
use of northern seabirds. Energetic cost of maintenance (basal metabolism,
thermoregulation, and procurement and processing of food) depends largely on
the following factors: (1) small birds have higher metabolic costs per unit size
than do larger ones; (2) body structure affects the cost of locomotion as well as of
food procurement; (3) climate affects metabolic costs; and (4) food availability
and nutrition vary among food types, and throughout the year within a food
type. Little is known of maintenance energetics in seabirds. Time and energy
allocations to items beyond basic maintenance are also compared. Patterns and
costs of molt and migration are known only in a general way, and the variety of
possible patterns suggests that more research would be of value. Almost nothing
is known of location and daily activities of seabirds outside the breeding season.
The review of breeding season activities is more comprehensive, and stresses the
variety of factors known to affect timing, and the total time devoted to and the
energetic costs of various aspects of reproduction. Some of these factors are
weather, year, geographic location, feeding conditions, age, sex, and distance of
food source from the breeding colony. Species characteristics such as clutch size,
egg and yolk size, developmental type, growth rate, food type, and behavior combine
with environmental variables to make seabirds a very diverse group in time
and energy budgeting. Time-energy studies and determination of productive
energy (energy remaining after maintenance needs have been met) can be useful
in pinpointing those groups of birds and the times of year when birds are most
vulnerable to environmental stress. Life history considerations suggest that
most seabirds are adapted to low population turnover and would not be able to
recover quickly from sudden increases in mortality.</p></div>

<p>Effective management of a population requires
manipulation of the factors most critical
in causing population increase or decrease.
Deciding what these factors are and which are
most suitable for effective manipulation is
very difficult due to the complexity of life
cycles and possible factors affecting demography.
It takes a thorough knowledge of a
species and of its relationships with the biotic
and abiotic environment to make effective
management decisions. The following review
of seabird time and energy use is meant to emphasize
the wide variation of species ecology
within this avian group.</p>

<p>Time and energy patterning is being used as
the basis of ecological comparison for the following
reason. Any activity of an animal requires
time and energy use; therefore, the patterning
of use makes a common thread to
which allocation to all activities in a bird's life
cycle can be related. Time-energy patterns can
be compared among birds with diverse food
types, habitats, life cycles, and life expectancy,
and therefore offer an opportunity for<span class="pagenum"><a name="Page_142" id="Page_142">[Pg 142]</a></span>
comparison not available through other kinds
of analysis (King 1974).</p>

<p>The amounts of time and energy allocated
by an organism to different aspects of survival
and reproduction should be regarded as
being molded by natural selection to optimize
(not necessarily to maximize) lifetime reproductive
output (Fisher 1958; Williams 1966;
Schoener 1971). Thus, differences in time and
energy use between species should reflect
adaptation to different biotic and abiotic environments.
By comparing time and energy use,
one can gain insight into the selective pressures
on each species and have a basis on
which to compare complex ecology more
meaningfully than if one listed other types of
differences.</p>

<p>This review of time-energy use in northern
seabirds cannot be comprehensive, largely because
many of the necessary data are lacking.
It stresses major areas of difference, however,
and points out aspects about which little is
yet known.</p>


<h4>Cost of Living</h4>

<p>Every animal must expend a basic amount
of energy on normal maintenance, excluding
activities normally allocated to a relatively
narrow time span, such as reproduction. This
"existence energy" expenditure consists of
basal metabolism, thermoregulation, and the
costs of gathering and processing food, and
could also be referred to as the animal's basic
"cost of living." In discussing the components
of the cost of living, energy use is emphasized
and time largely ignored&mdash;partly because
metabolism occurs irrespective of time
(it is not something the animal can turn off for
a period) and partly because time use in normal
maintenance and foraging has been little
studied.</p>


<h5><em>Metabolism</em></h5>

<p>Basal metabolic rate (BMR) depends
greatly on body size (Lasiewski and Dawson
1967; Zar 1968), and the costs per unit size are
higher for a small bird than for a large one
(Fig. 1). The BMR is somewhat lower in seabirds
and other nonpasserines than in passerines
of similar size (Dawson and Hudson
1970).</p>

<div class="figcenter" style="width: 564px;">
<img src="images/p_142.jpg" width="564" height="700" alt="" />
<div class="caption"><p>Fig. 1. Energy cost of various metabolic functions
in relation to body size in birds. "0&deg; Existence"
refers to total metabolic costs of caged birds held
at 0&deg;C. From Calder (1974).</p></div>
</div>

<p>The relationship between BMR and body
size is paralleled by that between size and
other metabolic costs, such as for thermoregulation
at a given temperature and for activity
(Fig. 1; Kendeigh 1970; Tucker 1970;
Schmidt-Nielsen 1972; Berger and Hart 1974;
Calder 1974). Basal metabolic rate can therefore
be used as an index of the overall cost of
living as far as metabolic functions are concerned.
Small birds must allocate a greater
proportion of their energy resources than
larger ones to merely staying alive, and have a
higher cost of living.</p>

<p>The suggestion in Fig. 1 that it is easy to
measure activity costs in a straightforward
manner is misleading, because the figure represents
measures taken under standard conditions.
Factors known to affect the cost of
flight, for example, include anatomical adaptations
(such as wing loading and wing shape),
the type of flight (ascending, descending, glid<span class="pagenum"><a name="Page_143" id="Page_143">[Pg 143]</a></span>ing),
and the speed of flight (Fig. 2; Tucker
1969, 1974; Hainsworth and Wolf 1975). The
cost of a series of short flights may be higher
than that for a long one because of the extra
energy required for takeoff and landing. A few
estimates have been made for the cost of
flight, mostly in birds moving almost constantly
(Lasiewski 1963; Nisbet 1963; Tucker
1972, 1974; Utter and LeFebvre 1970; Berger
and Hart 1974), but the methods may be inadequate
for birds that fly short distances
frequently.</p>

<div class="figcenter" style="width: 692px;">
<img src="images/p_143.jpg" width="692" height="700" alt="" />
<div class="caption"><p>Fig. 2. Energy cost of flying at different speeds and
angles as compared with basal metabolic rate
(BMR). Solid lines and solid circle refer to flight
cost and BMR for budgerigar <em>(Melopsitticus
undulatus)</em>. Dashed line and open circle refer to
flight cost and BMR of the laughing gull. From
Tucker (1969).</p></div>
</div>

<p>Little work has been done on the cost of
locomotion in seabirds: that of Eliassen (1963)
on great black-backed gulls <em>(Larus marinus)</em>,
Berger et al. (1970) on ring-billed gulls
<em>(L. delawarensis)</em>, and Tucker (1972) on the
laughing gull <em>(L. atricilla)</em>, and indirect calculations
of soaring flight characteristics in
albatrosses, <em>Diomedea</em> spp. (Cone 1964), and
the fulmar, <em>Fulmarus glacialis</em> (Pennycuick
1960). Swimming has been shown to be more
costly than flying in ducks (Schmidt-Nielsen
1972) and may be for seabirds as well. More
energy is also probably used in underwater
swimming than in flying.</p>

<p>The energetic costs of thermoregulation
under natural conditions are not easy to estimate.
Thermal energy is gained from and lost
to the environment, and the degree of exchange
depends not only on air temperature
but also on metabolic rate, insulation, body
temperature, posture, humidity, convection,
and radiation. Radiation, in turn, depends on
cloud cover, shade, and reflective and absorptive
characteristics of the organism and of the
environment (Porter and Gates 1969; Calder
and King 1974). Most of these quantities are
changing constantly, and insulation and
metabolic rate may vary on a seasonal basis
with acclimation (Dawson and Hudson 1970).</p>

<p>At present, no direct measurement technique
exists for determining natural thermoregulatory
costs, although a few estimates
have been made (King 1974), including several
for seabird nestlings (Dunn 1976<em>a</em>, 1976<em>b</em> for
double-crested cormorants, <em>Phalacrocorax
auritus</em>, and for herring gulls, <em>Larus argentatus</em>).
For most birds, the temperature environment
actually faced over a year's time has
never been measured, and for no bird has a
full description of the complete thermal environment
been made. It is clear that climate
and degree of exposure are important elements
in the basic cost of living, and that
thermoregulatory costs average higher in
small birds than in larger ones, but beyond
that little information is available. Work on
thermoregulatory costs of free-living chicks of
two species of seabirds suggests that insulative
properties can lead to marked differences
in the metabolic costs of different species in
an essentially identical environment (Dunn
1976<em>a</em>, 1976<em>b</em>).</p>


<h5><em>Food Procurement and Processing</em></h5>

<p>Gathering and processing food is another
major component of the cost of living. Both
the nutritional value of food and its availability
(a rather vague term covering both
abundance and ease of capture) are extremely
diverse and variable, making estimations of
foraging cost and benefit difficult (Ashmole
1971; Fisher 1972; Sealy 1975<em>a</em>).</p>

<p>Availability of food varies throughout the
year, particularly in marine invertebrates
that form the diet of many seabirds (e.g.,
Spaans 1971; B&eacute;dard 1969<em>a</em>). High arctic<span class="pagenum"><a name="Page_144" id="Page_144">[Pg 144]</a></span>
oceans have a very high peak of productivity
in the summer, whereas the low arctic has a
lower, but longer-lasting, peak (Ashmole
1971). Fish stocks increase in summer as well
(Snow 1960; Pearson 1968; Sealy 1975<em>a</em>), and
decline or disperse in autumn (Potts 1968).
Catch-ability may also differ widely from year
to year (e.g., E. K. Dunn 1973).</p>

<p>Marine foods are likely to have a patchy distribution,
which may make food stocks difficult
to locate, even in times of abundance
(Ashmole 1971; Sealy 1975<em>a</em>). Birds in localities
with low food abundance frequently show
alterations in time and pattern of foraging,
sometimes even changing diets (Cramp 1972;
Henderson 1972; Hunt 1972; Lemmetyinen
1972). The time and energy expended in finding
and capturing food by different seabird
species must vary widely according to the
form of foraging used: plunge-diving, beach
scavenging, aerial robbing, underwater pursuit,
and so on. Even when different species
have traveled the same distance to an identical
food stock, therefore, the costs of procurement
differ.</p>

<p>Time and energy spent foraging depends
not only on abundance and ease of capture,
but also on nutritional return, and on the age
and size of the bird. Fig. 3 shows that the
smaller species in a seabird community may
spend the most time foraging. Even though
this illustration is taken from the breeding
season when food demands of the young must
be taken into account, it suggests a difference
based on cost of living according to size.</p>

<div class="figcenter" style="width: 638px;">
<img src="images/p_144.jpg" width="638" height="700" alt="" />
<div class="caption"><p>Fig. 3. Time spent foraging in the breeding season
as a function of body size. From Pearson (1968).
AT = arctic tern <em>(Sterna paradisaea)</em>, CT = common
tern <em>(S. hirundo)</em>, ST = sandwich tern
<em>(Thalasseus sandvicensis)</em>, K = black-legged
kittiwake, P = common puffin, M = common
murre, LBB = lesser black-backed gull, S =
shag.</p></div>
</div>

<p>Age of the bird affects time and energy commitment
to foraging because younger birds
are often less skilled at capturing food. This
has been noted particularly in long-lived seabird
species (Orians 1969; Dunn 1972; LeCroy
1972; Buckley and Buckley 1974; Barash
et al. 1975). Older juveniles may be excluded
from feeding areas by more experienced, territorial
adults (Moyle 1966), whereas immatures
are not (Drury and Smith 1968; Ingolfsson
1969).</p>

<p>Nutritional and energetic return obtained
from food is a very important factor in foraging
strategy that has not received the attention
it deserves. Table 1 lists the caloric value
of various foodstuffs and illustrates how little
is known about foods eaten by seabirds. Although
caloric content and abundance of food
have often been accepted as the most important
determinants of foraging strategies
(Bookhout 1958; Emlen 1966; West 1967;
Bryant 1973), they may frequently be less important
than nutritional value and digestibility,
also shown in Table 1 (Pulliam 1974).</p>

<p>Since fish seem to be highly digestible, most
of the energy contained in them is available to
the consumer. There are, unfortunately, no
data on the digestibility of marine invertebrates,
but those for insects suggest that digestibility,
at least of crustaceans with exoskeletons,
is somewhat lower than that for
fish. A bird would therefore have to eat a
larger biomass of invertebrates than of fish to
satisfy the same energetic needs (although
cost of procurement might not be as high as
for fish).</p>

<p><span class="pagenum"><a name="Page_145" id="Page_145">[Pg 145]</a></span></p>

<div class="center">
<table border="0" cellpadding="4" cellspacing="0" summary="Table 1. Nutritional value of foods eaten by birds.">
    <caption>Table 1. <em>Nutritional value of foods eaten by birds.</em> After data in Hunt (1972) and E. H. Dunn (1973).</caption>
<col></col>
<col span="6" width="12.5%"></col>

  <tr>
    <th rowspan="2">Food type</th>
    <th rowspan="2">Kcal/g fresh wt.</th>
    <th colspan="3">Percent fresh wt. composed of:</th>
    <th rowspan="2">Digestive efficiency</th>
    <th rowspan="2">Kcal metabolizable energy/g fresh wt.</th>
  </tr>
  <tr>
    <th>H<sub>2</sub>O</th>
    <th>Fat</th>
    <th>Protein</th>
  </tr>
  <tr>
    <td>Vegetable</td>
    <td>1.2-5.2</td>
    <td>59-86</td>
    <td>0.4-3</td>
    <td>3-5</td>
    <td>30-32</td>
    <td>0.3-2.3</td>
  </tr>
  <tr>
    <td>Tropical fruits</td>
    <td>1.2</td>
    <td>75</td>
    <td>8</td>
    <td>1</td>
  </tr>
  <tr>
    <td>Various seeds</td>
    <td>4.0-7.3</td>
    <td>3-13</td>
    <td>1-40</td>
    <td>10-29</td>
    <td>76-80</td>
    <td>3.0-5.2</td>
  </tr>
  <tr>
    <td>Various insects</td>
    <td>1.4-5.2</td>
    <td>65-75</td>
    <td>1-3</td>
    <td>9-18</td>
    <td>66-69</td>
    <td>0.9-3.5</td>
  </tr>
  <tr>
    <td>Whiting (fish)</td>
    <td>1.1</td>
    <td>81</td>
    <td></td>
    <td></td>
    <td>79</td>
    <td>0.9</td>
  </tr>
  <tr>
    <td>Various freshwater fishes</td>
    <td>1.2</td>
    <td>75</td>
    <td>5</td>
    <td>18</td>
    <td>81</td>
  </tr>
  <tr>
    <td>Mix of fish eaten by double-crested cormorants on NE coast</td>
    <td>1.1</td>
    <td>74</td>
    <td>1</td>
    <td>16</td>
    <td>82</td>
    <td>0.9</td>
  </tr>
  <tr>
    <td>Fresh herring and mackerel</td>
    <td>1.9</td>
    <td>67</td>
    <td>13</td>
    <td>19</td>
  </tr>
  <tr>
    <td>Garbage ("average" mix)</td>
    <td>1.5</td>
    <td>67</td>
    <td>8</td>
    <td>19</td>
  </tr>
  <tr>
    <td>Crab with eggs</td>
    <td>1.0</td>
    <td>68</td>
    <td>5</td>
    <td>1</td>
  </tr>
  <tr>
    <td>Euphausid shrimp</td>
    <td>0.8</td>
    <td>80</td>
    <td>2</td>
    <td>1</td>
  </tr>
  <tr>
    <td>Clam (edible part only)</td>
    <td>0.8</td>
    <td>80</td>
    <td>1</td>
    <td>13</td>
  </tr>
</table></div>

<p>A bird must satisfy not only energetic
needs, but also nutritional requirements. Fish
are high in protein (Table 1), but what little is
known of marine invertebrates suggests a low
proportion of protein in relation to total bulk.
Protein is vital to growth of nestlings (Fisher
1972; Lemmetyinen 1972), and 4-8% protein
in the diet seems to be required for minimal
maintenance of adults (Martin 1968; Fisher
1972). Some seabirds (such as puffins, <em>Fratercula</em>)
that eat a varied diet raise their young
exclusively on fish (B&eacute;dard 1969<em>b</em>; Nettleship
1972; Sealy 1973<em>a</em>).</p>

<p>Other aspects of nutrition, such as vitamins
and minerals, are also important to avian
health (Brisbin 1965; Fisher 1972). To further
complicate matters, nutritional values vary
with season, as do birds' requirements for
them (Myton and Ficken 1967; Moss 1972).
Adults must adjust their time and energy
allocation to foraging to optimize not only
energetic, but also nutritional return.</p>

<p>Optimal time and energy allocation has
been studied in theory (Orians 1971; Schoener
1971; Pulliam 1974; Katz 1974) and some direct
observations have been carried out,
largely on seedeaters (Bookhout 1958; Myton
and Ficken 1967; Royama 1970; Moss 1972;
Willson 1971; Willson and Harmeson 1973).
The direct studies, in particular, point out the
basic importance of studying cost-benefit
ratios by interrelating complex factors of food
availability, searching patterns, and type,
size, and caloric and nutritional value of
foods.</p>


<h4>Time-energy Use Beyond
the Cost of Living</h4>

<p>This section concerns variation in time and
energy allocated by seabirds to activities
above and beyond the cost of living&mdash;particularly
to migration, molt, and reproduction.
Allocation to such items as avoidance of predation
and competition is not considered here,
because they are not readily analyzed as activities
to which time and energy are devoted
in a specific part of the annual cycle.</p>

<p>The previous discussion dwelt on energy
considerations and could have referred to almost
any group of birds. The following treatment
centers on time use of northern seabirds.
Little is known of energetics beyond the cost
of living, although estimates have been made
for certain aspects of migration, molt, and reproduction
(Nisbet 1963; Hussell 1969; Hart
and Berger 1972; Payne 1972; Ricklefs 1974).
Essentially nothing is known, however, of the
relationship of such costs to the amount of
energy available to the bird once basic maintenance
costs have been met (productive
energy). Because such complete data are not
available, the following account dwells largely
on variation in timing and total time devoted
to activities beyond basic maintenance.</p>

<p><span class="pagenum"><a name="Page_146" id="Page_146">[Pg 146]</a></span></p>

<div class="figcenter" style="width: 700px;">
<img src="images/p_146.jpg" width="700" height="466" alt="" />
<div class="caption"><p>Fig. 4. Typical patterns of generalized annual cycles in reproduction, migration, and wing molt in northern
seabirds. Solid line shows reproductive season, dotted line the period of migration or dispersion, and
dashed line the period of annual primary molt. Data from Dorst (1961), Stresemann and Stresemann
(1966), and Ashmole (1971).</p></div>
</div>


<h5><em>Migration</em></h5>

<p>Among northwestern North American seabirds,
most coastal feeders, such as gulls, cormorants,
and many alcids and petrels, have
only a short southward migratory movement,
and many others are more or less resident
(Dorst 1961; Ashmole 1971). Terns, on the
other hand, migrate long distances in a short
time to places where small fish are available
near shore in the winter. Other long-distance
migrants&mdash;Sabine's gull <em>(Xema sabini)</em>,
jaegers (<em>Stercorarius</em> spp.), pelagic phalaropes
(<em>Phalaropus</em>, <em>Lobipes</em>), and kittiwakes (<em>Rissa</em>
spp.)&mdash;tend to scatter widely over the southern
ocean, concentrating near areas of upwelling
(Dorst 1961; Ashmole 1971). Groups such
as murres (<em>Uria</em> spp.), eiders (<em>Somateria</em> spp.),
and grebes (<em>Podiceps</em> spp.) may move considerable
distances by swimming (Dorst 1961;
Tuck 1960). True migration tends to take
place directly before and after reproduction,
whereas dispersal or nomadism takes place
over a long period of the winter (Fig. 4).</p>

<p>Among species remaining in the northern
hemisphere, younger birds frequently disperse
greater distances than do breeding
adults (Coulson 1961; Kadlec and Drury 1968;
Southern 1967), and the degree of dispersal
can vary among colonies of the same species
(Coulson and Brazendale 1968).</p>

<p>Energy costs of migration must vary according
to distances covered and amount of
time allocated to migration. Aside from the
references to cost of flight mentioned earlier,
however, migratory costs have scarcely been
studied. Dolnik (1971) has estimated that
chaffinches <em>(Fringilla coelebs)</em> expend about
as much energy migrating south as they
would on thermoregulation if they overwintered
on their breeding grounds. Long-distance
migration is presumably selected because
the birds are able to collect food more
efficiently, because the risks of death or injury
in migrating are less than in residency,
and so on. Interspecific and intraspecific competition
may also be involved (Cox 1968). In
other words, migratory patterns are selected
to optimize survival and reproduction in alternating
environments (Cohen 1967; Drury and
Nisbet 1972).</p>

<p><span class="pagenum"><a name="Page_147" id="Page_147">[Pg 147]</a></span></p>

<div class="center">
<table border="0" cellpadding="4" cellspacing="0" summary="Table 2. Wing molt in alcids.">
    <caption>Table 2. <em>Wing molt in alcids.</em> After Stresemann and Stresemann (1966).</caption>
  <tr>
    <th>Rapidity of wing molt and (indented) flight capability in molt</th>
    <th>Timing of start of molt</th>
    <th>Species</th>
  </tr>
  <tr>
    <td>Slow<br /><span style="margin-left:2em">Retained</span></td>
    <td>During care of young</td>
    <td>Cassin's auklet, parakeet auklet <em>(Cyclorhynchus psittacula)</em>, whiskered auklet <em>(Aethia pygmaea)</em></td>
  </tr>
  <tr>
    <td><span style="margin-left:2em">Retained</span></td>
    <td>After young become independent</td>
    <td>Least auklet <em>(A. pusilla)</em>, crested auklet <em>(A. cristatella)</em></td>
  </tr>
  <tr>
    <td>Rapid<br /><span style="margin-left:2em">Poor</span></td>
    <td>After arrival in winter quarters</td>
    <td>Marbled murrelet, Kittlitz's murrelet <em>(Brachyramphus brevirostris)</em></td>
  </tr>
  <tr>
    <td>Almost synchronous<br /><span style="margin-left:2em">None</span></td>
    <td>After end of breeding</td>
    <td>Xantus' murrelet <em>(Endomychura hypoleuca)</em></td>
  </tr>
  <tr>
    <td>Synchronous<br /><span style="margin-left:2em">None</span></td>
    <td>As soon as young go to sea</td>
    <td>Guillemots (<em>Cepphus</em> spp.), murres, razorbill <em>(Alca torda)</em>, dovekie <em>(Alle alle)</em></td>
  </tr>
  <tr>
    <td><span style="margin-left:2em">None</span></td>
    <td>In winter, after body molt</td>
    <td>Puffins</td>
  </tr>
</table></div>

<p>Although one may suspect that location of
winter food supply is the main environmental
factor affecting migratory patterns, there is
little direct evidence on the reasons for, or the
benefits accruing from, the different patterns
seen in seabirds. Study of cost-benefit ratios
of foraging in different stages of migration
might help clarify the question.</p>


<h5><em>Molt</em></h5>

<p>Patterns of molt vary widely among seabirds.
The commonest pattern is for a prenuptial
body molt to occur in spring, and for an
extended wing molt to begin after the breeding
season and continue well into the winter
(Fig. 4). In short-distance migrants, molt may
overlap slightly with the end of breeding and
can last up to 6 months, as in most gulls,
terns, alcids, nonmigratory jaegers, and cormorants
(Stresemann and Stresemann 1966).</p>

<p>Long-distance migrants frequently delay
molt until in the winter quarters (lesser black-backed
gull, <em>Larus fuscus</em>; Sabine's gull;
jaegers; arctic tern, <em>Sterna paradisaea</em>; and
marbled murrelet, <em>Brachyramphus marmoratus</em>)
and molt there may occur rapidly
(3.5 months in the arctic tern). Certain other
long-distance migrants begin molt before
leaving the breeding grounds (herring gull;
skua, <em>Catharacta skua</em>; Leach's petrel,
<em>Oceanodroma leucorhoa</em>; and fulmar), although
molt may be interrupted during migration,
as in <em>Larus argentatus heuglini</em>
(Stresemann and Stresemann 1966). Duration,
timing, and rapidity of molt are particularly
varied among the alcids (Table 2).</p>

<p>A few unusual molt patterns are found in
northern seabirds. The ivory gull <em>(Pagophila
eburnea)</em> has its major annual wing and body
molt immediately before it breeds. In several
other species such as the glaucous gull <em>(Larus
hyperboreus)</em> and Cassin's auklet <em>(Ptychoramphus
aleuticus)</em> the molt almost completely
overlaps the reproductive cycle (Johnston
1961; Payne 1965). Potts (1971) documented a
molt pattern in shags <em>(Phalacrocorax aristotelis)</em>
which is more typical of tropical seabirds.
Several cycles of wing molt take place
simultaneously, each lasting more than a
year, and molt ceases in winter. By the time
breeding age is reached, each flight feather is
replaced once a year.</p>

<p>Within these broad categories of molt pattern
there are sometimes variations according
to age, sex, and even subspecies (Stresemann
and Stresemann 1966). Male common eiders
<em>(Somateria mollissima)</em> molt directly after
mating, when their reproductive role is completed,
whereas females molt only after they<span class="pagenum"><a name="Page_148" id="Page_148">[Pg 148]</a></span>
have taken their young to sea. Nonbreeders
and failed breeders frequently begin molt
while other adults are still raising young and
not molting&mdash;e.g., many alcids, gulls, storm-petrels,
and fulmars (Stresemann and Stresemann
1966; Ingolfsson 1970; Harris 1971;
Harris 1974; Sealy 1975<em>b</em>). In ivory gulls,
which molt just before reproduction, and in
Sabine's gulls, which complete molt just before
breeding, nonbreeders may extend wing
feather growth into the breeding season
(Stresemann and Stresemann 1966).</p>

<p>There is little information on the energetic
cost of molt, although there are indications of
at least some expense. Belopol'skii (1961)
showed that nonmolting seabird species
tended to gain weight after reproduction,
whereas those that immediately started molt
tended to lose weight. Among other birds,
however, it is common for individuals to gain
weight just before, and even during molt
(Payne 1972). The BMR is known to rise in
molting birds (Blackmore 1969; Lustick 1970;
Payne 1972), from as little as 5% to as much
as 34% above nonmolting levels. In one
study, about 35-40% of the increased BMR
represented extra thermoregulatory costs incurred
by lessening of insulation and increase
in heat loss from well-vascularized new
feathers; the rest of the increase represented
the energetic cost of growing feathers (Gavrilov
1974). The fact that molt rarely overlaps
with breeding suggests that the energetic
cost, even if slight, may be incompatible with
the already high costs of reproduction (Payne
1972). Cassin's auklets, which do molt while
breeding, may cease molt while feeding large
young (Payne 1965), and certain species interrupt
molt during migration (Stresemann and
Stresemann 1966). Doubtless a rapid simultaneous
molt is more costly than a long
gradual one.</p>

<p>Rapid molt appears to occur at a time in the
annual cycle when food resources are abundant
(spring or late summer), whereas extended
molt generally occurs over winter (e.g.,
B&eacute;dard 1969<em>a</em>). If one speculates that energy
availability is the main determinant of molt
patterns, one can also speculate on the cause
behind some of the more unusual patterns.
Possibly birds in which molt and breeding
overlap either have extraordinary available
energy at that time or else face shortages in
other periods. For example, ivory gulls, which
breed in the high Arctic, molt when food resources
have become abundant in the low Arctic
but before the high Arctic breeding
grounds have thawed sufficiently for reoccupation.</p>

<p>Speed of molt may also reflect availability
of energy resources or of nutrients needed for
feather growth (Payne 1972), but must also be
influenced by the need for full flight capabilities
to obtain food. The eider duck and many
alcids that shed wing feathers almost simultaneously
do not need their wings for flight
after the young have left the breeding colony.
Hydrodynamic considerations suggest that
their fishing capabilities may even be improved
(Storer 1960). This is not true for the
smaller species&mdash;e.g., <em>Aethia</em> molts only one
feather at a time and retains full flight capabilities
(Table 2). Climate may also influence
simultaneity of molt if heat loss in rapid molt
is particularly severe.</p>


<h5><em>Reproduction</em></h5>

<p>Time use of seabirds is best known for the
reproductive period, when the birds are on
relatively accessible breeding grounds, the
weather is most suitable for observation, and
academic researchers are freed from their
jobs. Even so, the details of timing are known
for only a few of the species and localities on
the northwest North American coast (e.g.,
Drent and Guiguet 1961; Drent et al. 1964;
Cody 1973; Sealy 1973<em>a</em>, 1975<em>b</em>, 1975<em>c</em>). The
following discussion emphasizes the multitude
of environmental factors known to influence
timing and total length of time devoted
to various aspects of the reproductive cycle.</p>


<h6>Timing of the Season</h6>

<p>Each species of seabird returns to the
colony site when weather conditions have
ameliorated sufficiently to meet its particular
needs. For example, the early arrivals to islands
in the Barents Sea are murres, kittiwakes,
and herring gulls, which need only
small cracks in the sea ice to meet their feeding
requirements (Belopol'skii 1961). Eiders in
North America also return early, when a few
ice leads have formed (Schamel 1974). Common
puffins <em>(Fratercula arctica)</em> and mew<span class="pagenum"><a name="Page_149" id="Page_149">[Pg 149]</a></span>
gulls <em>(Larus canus)</em> are somewhat later
arrivals, and terns and a few parasitic jaegers
<em>(Stercorarius parasiticus)</em> are the latecomers
to Barents Sea colonies (Belopol'skii 1961).</p>

<p>The timing of the season (as illustrated in
Fig. 5) varies widely among localities, and because
of local weather patterns and ocean currents,
this variation can be unrelated to latitude
(Belopol'skii 1961). Examples of such
variation are also known in North America:
for instance, Leach's storm-petrels in Alaska
lay eggs 2 to 3 weeks later than do those in
California (Harris 1974); however, the details
of timing are largely unknown for many
species in this region. Progression of thaw,
which also varies from year to year, causes
variation in the timing of the breeding season
(Belopol'skii 1961; Evans and McNicholl
1972). Fig. 6 shows the diversity in start of
the breeding season for different species on
the same island in the Barents Sea as well as
variation in time devoted to various components
of the reproductive cycle.</p>

<div class="figcenter" style="width: 700px;">
<img src="images/p_149a.jpg" width="700" height="409" alt="" />
<div class="caption"><p>Fig. 5. Differential average arrival on breeding
grounds and average duration of prenesting
period of thick-billed murres <em>(Uria lomvia)</em> and
black-legged kittiwakes on various colonies in the
Barents Sea. From Belopol'skii (1961). Length of
prenesting period in days (shaded bars) indicated
on right. Letters represent locations as follows: A
= Novaya Zemlya, Kara Straits; B = Novaya
Zemlya, Karmakuly Bay; C = Franz Josef Land;
and D = East Murman.</p></div>
</div>


<h6>Prenesting Activities</h6>

<p>Some species are apparently able to delay
maturity of sexual organs until environmental
conditions are suitable for nesting&mdash;e.g.,
burrow and crevice nesters in the
Barents Sea do not become sexually mature
until snowmelt (Belopol'skii 1961). Many
others, however, reach sexual maturity soon
after arrival on the breeding grounds, and a
few (such as jaegers and kittiwakes) mature in
migration or on the wintering grounds (Belopol'skii
1961). Northern phalaropes <em>(Lobipes
lobatus)</em> sometimes lay eggs as early as
1 week after arrival (Hilden and Vuolanto
1972). This factor, in combination with timing
of arrival, affects the amount of time spent in
prenesting activities (Fig. 6). Most species
gain weight during this period (Belopol'skii
1961), and the time required for each species
to reach full breeding condition must also depend
on feeding conditions and the state of
the bird on its arrival at the nesting site.
These factors help explain why early arriving
species are not necessarily early nesters
(Fig. 6).</p>

<div class="figcenter" style="width: 700px;">
<img src="images/p_149b.jpg" width="700" height="289" alt="" />
<div class="caption"><p>Fig. 6. Variation in timing of events in the reproductive
cycle of Barents Sea seabirds nesting on the
same island. Data from Belopol'skii (1961).
Shaded bars at left indicate the prelaying
periods, open bars the incubation periods, and
shaded bars at right the portion of the growth
period in which the chick remains at the nest site.
Total length of time indicated is about 6 months.</p></div>
</div>

<p>Aside from nest building, most prenesting
activity consists of courtship and territorial
behavior. These activities have been well described
for representative seabird species, but
because assessments of time and energy devoted
to them have been almost completely
neglected, they are not discussed further here.
For examples, see accounts in Gross (1935) for
Leach's storm-petrel; Tinbergen (1935),
Bengtson (1968), H&ouml;hn (1971), and Howe
(1975) for phalaropes; Storer (1952) for common
murre, <em>Uria aalge</em>, and black guillemot,
<em>Cepphus grylle</em>; Tschanz (1959) for common
murre; Brown et al. (1967) for Sabine's gull;
Tinbergen (1960) for herring gull; McKinney<span class="pagenum"><a name="Page_150" id="Page_150">[Pg 150]</a></span>
(1961) for eiders; Snow (1963) for shag; Thoresen
(1964) for Cassin's auklet; Vermeer (1963)
and James-Veitch and Booth (1974) for glaucous-winged
gull <em>(Larus glaucescens)</em>; and
Andersson (1973) for jaegers.</p>


<h6>Nest Building</h6>

<p>Although many northern seabirds have essentially
no nest, they may spend considerable
time working or displaying at the site
(Belopol'skii 1961). Black-legged kittiwakes
<em>(Rissa tridactyla)</em> have substantial nests, but
they are built in a comparatively short time
(about a week) soon after the birds arrive
(Fig. 6). Shags also have substantial nests,
but they are not completed until about 1 or 2
weeks before the first egg is laid (Snow 1963).
Herring gulls build smaller nests, 5 to 10 days
before laying, although in the Far North they
and glaucous gulls may not start building the
nest until the first egg is laid (Belopol'skii
1961). The eider always begins preparing the
nest when the first egg is laid (Belopol'skii
1961; Schamel 1974), and terns and skuas,
which build no nests, choose their sites at that
time. Murres, which frequently lay their eggs
directly on snow, choose a site somewhat
earlier and spend considerable time protecting
it (Belopol'skii 1961). Burrows may be dug
within a period as short as 3 days for Leach's
storm-petrel (Gross 1935) to one as long as
several weeks in Cassin's auklets (Manuwal
1974<em>a</em>). Overall, the prelaying period is longer
for burrow nesters than for those using
crevices (Sealy 1973<em>a</em>).</p>

<p>The amount of time and energy spent by the
male and female in nest building differs
among species. In Leach's storm-petrel, the
male digs the burrow (Gross 1935), whereas in
eiders, the nest is built entirely by the female.
In most seabird species, the sexes share in
nest construction, but roles may still be separated.
For example, in shags the male collects
the nest material and the female builds the
nest (Snow 1963).</p>


<h6>Egg Laying</h6>

<p>Timing of egg laying is influenced not only
by weather (Erskine 1972; Sealy 1975<em>c</em>), but
also by numerous biotic factors. Smith (1966)
showed that where glaucous gulls, herring
gulls, and Thayer's gulls <em>(Larus thayeri)</em> breed
in mixed colonies, the peak of sexual activity
and egg laying in Thayer's gull is about midway
between the peaks for the other two
species (Fig. 7). In nearby colonies where
herring gulls are absent, however, the peak of
sexual activity in Thayer's gulls is delayed
about a week, and activity continues for a significantly
longer period (Fig. 7).</p>

<div class="figcenter" style="width: 700px;">
<img src="images/p_150.jpg" width="700" height="470" alt="" />
<div class="caption"><p>Fig. 7. Timing of peak sexual activity (a combined
measure of egg laying and testes size) in colonies
of arctic gulls of different species composition.
From Smith (1966).</p></div>
</div>

<p>Annual variations in food supply also will
affect the start of the egg-laying season. Belopol'skii
(1961) cited an example from the
Barents Sea in 1940 when a series of storms
made it difficult for certain seabirds to find
food. Murres and kittiwakes, which were able
to catch fish, started reproductive activities
on schedule. Gull breeding was delayed, however,
and egg laying began in force only after
fishing boats arrived and started discarding
offal. Onset of egg laying in great cormorants
<em>(Phalacrocorax carbo)</em> is correlated to April air
temperatures (Erskine 1972), and this may
also be related to variations in spring increase
of food availability. In certain birds the breeding
season has been shown to start particularly
early when food supplies are unusually
abundant (H&ouml;gstedt 1974; K&auml;llender 1974),
but this has not yet been demonstrated in
seabirds.</p>

<p>Lastly, age and sex of seabirds are known to
affect the timing of egg laying (e.g., Coulson
and White 1960; Lack 1966); older, more experienced
birds tend to lay earlier than do
younger ones. In shags, males tend to breed
progressively earlier as they increase in age,
but females do not (Snow 1963).</p>

<p><span class="pagenum"><a name="Page_151" id="Page_151">[Pg 151]</a></span></p>

<p>Contrary to the situation in passerines, seabirds
tend to lay their clutches with relatively
large time intervals between eggs. Eggs may
be laid every 2nd or 3rd day in alcids, larids,
sternids, stercorariids, and phalacrocoracids
(Lack 1968), but every day in phalaropes
(Howe 1975). Inasmuch as clutch size in
northern seabirds varies from one to five or
six, the length of the laying period varies
widely among species.</p>

<p>Energetic costs of egg laying depend on the
actual caloric content of the egg and the speed
with which the ova are developed (Ricklefs
1974). The energy in the egg is contained
mainly in the yolk, and yolk size depends
largely on the developmental pattern shown
by the young after hatching (Table 3). Precocial
chicks are hatched at a relatively advanced
stage, are covered with down, and
have open eyes, can maintain reasonably
homeothermic body temperature, and leave
the nest site to feed themselves after a few
hours or days. At hatching, semiprecocial
chicks appear similar to precocial chicks, although
they are slightly less well developed
(Ricklefs 1974; Dunn 1975<em>a</em>). In contrast to
precocial chicks, they remain at the nest site
for some time, are fed by their parents, and
tend to grow rather rapidly (Ricklefs 1968).
Altricial nestlings hatch at a much less advanced
stage of development. They are naked,
blind, helpless, essentially poikilothermic, and
depend completely on their parents for food
and shelter. They usually remain at the nest
until full grown. Semialtricial chicks show
somewhat intermediate characteristics (Nice
1962).</p>

<div class="center">
<table border="0" class="tdc" cellpadding="4" cellspacing="0" summary="Table 3. Amount of yolk in eggs of different types of birds.">
    <caption>Table 3. <em>Amount of yolk in eggs of different types of birds.</em> After Ricklefs (1974).</caption>
  <tr>
    <th>Developmental type</th>
    <th>Percent yolk<br />(by weight)</th>
  </tr>
  <tr>
    <td>Precocial</td>
    <td> 30-60</td>
  </tr>
  <tr>
    <td>Semiprecocial</td>
    <td> 25-30</td>
  </tr>
  <tr>
    <td>Altricial</td>
    <td> 15-25</td>
  </tr>
</table></div>

<p>The amount of yolk (and therefore energy)
in an egg is positively correlated to the degree
of development at hatching (Table 3). The
same is true for egg size: altricial and semialtricial
birds have smaller eggs relative to
adult body weight than do semiprecocial and
precocial birds (Fig. 8). Clutch size, however,
is unrelated to energy content of the eggs. For
example, shags (which are altricial) and eiders
(precocial) have among the largest clutches of
northern seabirds (four to six eggs).</p>

<div class="figcenter" style="width: 593px;">
<img src="images/p_151.jpg" width="593" height="700" alt="" />
<div class="caption"><p>Fig. 8. Egg weight as a function of body weight in
various northern seabirds. Solid symbols represent
precocial and semiprecocial species, and
open symbols altricial and semialtricial species:
solid circles, alcids; solid triangles, gulls, terns,
and jaegers; solid squares, eiders; open squares,
cormorants and <em>Morus bassanus</em>; and open circles,
petrels. Data from Belopol'skii (1961); Drent
(1965); Sch&ouml;nwetter (1967); Lack (1968); B&eacute;dard
(1969<em>a</em>); Cody (1973); Sealy (1973<em>b</em>); Harris (1974);
and Manuwal (1974<em>a</em>).</p></div>
</div>

<p>The energetic cost of egg laying depends
not only on caloric content of the egg and
clutch size, but also on speed of development.
Ricklefs (1974) has shown that the energetic
cost per day of egg laying can be calculated
from the energy content of the yolk and white,
clutch size, the amount of follicular growth
per day, and the laying interval between eggs.
The energy content of a single egg (expressed
as percentage of BMR) has been estimated as
follows: 45 (altricial passerines), 103 (semial<span class="pagenum"><a name="Page_152" id="Page_152">[Pg 152]</a></span>tricial
raptors), 126 (precocial galliformes),
180 (precocial ducks), 226 (precocial shorebirds),
and 320 (semiprecocial gulls and terns).
Gulls and terns thus have very costly eggs, as
well as a moderately high clutch size (three).
However, the development time for a single
ovum in the herring gull is unusually long&mdash;9
to 10 days (King 1973). Ricklefs (1974), who
calculated the energetic cost per day (expressed
as percentage BMR), estimated the
cost of a clutch in gulls and terns (120% BMR
per day) to be similar to that for various
groups of precocial birds (about 125-180%
BMR per day). Unfortunately, the data required
for calculation of the average energetic
cost of a clutch are not available for other
northern seabirds.</p>

<p>For no species have all the additional factors
influencing the energetic cost of a clutch
been taken into account. For example, eggs in
a clutch may vary in size (and caloric content)
according to sequence in the clutch (Preston
and Preston 1953; Snow 1960; Coulson 1963;
Coulson et al. 1969). Age has a definite effect
on laying energetics, as older birds lay larger
eggs (Coulson and White 1958; Snow 1960;
Coulson 1963; Coulson et al. 1969) and lay
larger clutches (Coulson and White 1960;
Snow 1960). They also lay, on average, earlier
in the season (Coulson and White 1958; Snow
1960; Coulson et al. 1969), and eggs laid late
in the season (whether by young birds or older
ones in re-nesting attempts) tend to be smaller
and contain less energy. In addition, egg
quality can vary with food supply: Snow
(1960) found eggs to have more yolk in years
when food was abundant than in years when
food was scarce.</p>

<p>Egg-laying costs are, of course, borne entirely
by the female, although males may contribute
some time and energy toward egg laying
through courtship feeding (Ashmole 1971;
Henderson 1972; Nisbet 1973). Courtship
feeding takes place in most lariforms but not
in eiders, phalaropes, or cormorants.</p>

<p>The time and energy expended on egg laying
can be profoundly influenced by the degree
of nest destruction, since females usually
lay a replacement clutch if the loss of the first
does not occur too late in the season. Factors
causing egg destruction are numerous, but
among the most important in the north is predation.
As is shown in Table 4, the degree of
egg predation in common murres is correlated
to degree of exposure of the nest&mdash;so even
such an unlikely sounding factor as physical
characteristics of the nest site can affect the
average time and energy expended on egg laying
by a given species or population. Genuine
second clutches are occasionally laid by phalaropes
(Hilden and Vuolanto 1972) and
Cassin's auklets (Manuwal 1974<em>a</em>).</p>

<div class="center">
<table border="0" cellpadding="4" cellspacing="0" summary="Table 4. Predation on nests of common murres according to degree of exposure.">
    <caption>Table 4. <em>Predation on nests of common murres according to degree of exposure.</em> From Belopol'skii (1961).</caption>
  <tr>
    <th>Nest exposure</th>
    <th>Nests destroyed by predators<br />(%)</th>
  </tr>
  <tr>
    <td>Completely hidden</td>
    <td class="tdr"> 3.2</td>
  </tr>
  <tr>
    <td>Partly exposed</td>
    <td class="tdr"> 5.8</td>
  </tr>
  <tr>
    <td>Largely exposed</td>
    <td class="tdr"> 13.6</td>
  </tr>
  <tr>
    <td>Completely exposed</td>
    <td class="tdr"> 18.2</td>
  </tr>
</table></div>

<p>In short, time and energy devoted to egg
laying depend not only on the species, but also
on a multitude of other biotic and abiotic factors,
such as age, sex, degree of nest destruction,
weather, other species present, and feeding
conditions.</p>


<h6>Incubation</h6>

<p>The total time devoted to incubation does
not depend directly on developmental type or
egg size but differs markedly among families
(Lack 1968). Since incubation period seems to
be closely linked to fledging period, factors affecting
growth rate (discussed later) apparently
affect incubation period as well.</p>

<p>Each species has a different incubation
rhythm. In birds in which the sexes share in
incubation, the sexes exchange places at intervals
that differ widely among different birds:
several hours in lariforms and some alcids
(Drent 1965; Lack 1968; Preston 1968; Drent
1970); about 4 h in shags (Snow 1963); up to
11 h in the ivory gull (Bateson and Plowright
1959); up to 24 h in certain other alcids (Manuwal
1974<em>a</em>; Sealy 1975<em>a</em>); 33 h (on the average)
in common puffins (Myrberget 1962); 72 h in
ancient murrelets, <em>Synthliboramphus antiquus</em>
(Sealy 1975<em>a</em>); and 96 h in Leach's
storm-petrel (Gross 1935). Degree of attentiveness
once a bird is on the nest also varies.
Petrels may leave the egg for several days<span class="pagenum"><a name="Page_153" id="Page_153">[Pg 153]</a></span>
(Gross 1935), whereas herring gulls cover
their eggs 98% of the time (Drent 1970).</p>

<p>The sexes share in incubation in most seabirds
(Snow 1960; Drent 1965, 1970; B&eacute;dard
1969<em>a</em>), although females frequently take on
the greater role (Belopol'skii 1961). Only male
phalaropes incubate the eggs, and only female
eiders. Eider hens do not feed during the entire
incubation period (25 days) and leave the
nest only for short periods of about 10 min
(Belopol'skii 1961; Schamel 1974).</p>

<p>Several methods exist for calculating the
amount of heat input necessary for normal development
of a clutch of eggs (Ricklefs 1974).
There is controversy, however, as to whether
an adult can provide this warmth from excess
body heat lost during the course of normal
metabolism or whether the adult must raise
its metabolic level to produce extra heat (Kendeigh
1973; King 1973; Ricklefs 1974). Several
studies of incubating birds suggest that, in at
least some situations, adults need not raise
metabolic levels, but in others (large clutch,
severe weather), they probably do (Ricklefs
1974). Drent (1972) estimated that herring
gulls raise metabolic levels to a significant degree
during incubation.</p>

<p>In spite of the lack of quantitative data, one
can surmise that the cost of incubation varies
among seabirds. Precocial and semiprecocial
birds tend to have a larger clutch weight relative
to body weight than do altricial birds
(Fig. 8; Lack 1968), and therefore require
greater heat input to the eggs. These costs
may be reduced by heavily insulating the nest
(e.g., eiders), or by nesting in burrows, which
have much more moderate and even climates
than do external nests (Richardson 1961;
Manuwal 1974<em>a</em>). Other semiprecocial species,
however, such as the murre, may sometimes
lay eggs directly on snow or ice (Belopol'skii
1961)&mdash;presumably at increased incubation
costs. Lastly, certain species incubate eggs
with their feet (e.g., cormorants), rather than
develop featherless brood patches. There are
no measurements of comparative heat flow
from feet versus brood patches.</p>


<h6>Raising Nestlings</h6>

<p>The length of the nestling period (hatching
until departure from the nest) varies greatly
among northern seabirds (Fig. 6). Nestling
period depends on the stage of growth at
which the young leave the nest and the rate at
which they attain that stage. Growth rate in
turn depends largely on body size and developmental
type.</p>

<p>The stage of growth attained when birds
leave the nest varies considerably (Fig. 9).
Precocial eiders leave the nest within a day of
hatching, whereas altricial shags remain until
completely grown. The young of semiprecocial
species, on the other hand, leave the nest at all
stages between these extremes. Larids normally
remain at the nest until 75-90% grown,
but certain alcids leave much sooner&mdash;well before
the young can fly.</p>

<div class="figcenter" style="width: 700px;">
<img src="images/p_153.jpg" width="700" height="515" alt="" />
<div class="caption"><p>Fig. 9. Percentage of total growth completed in the
egg (shaded bar at left), at the nest site (open bar),
and after nest-leaving (shaded bar at right) in
various northern seabirds. From Belopol'skii
(1961).</p></div>
</div>

<p>Growth rate depends both on body size and
developmental type (Fig. 10). The length of
stay at the nest for precocial young is unaffected
by growth rate (which is typically very
slow), since they leave soon after hatching.
The nestling period of semiprecocial and altricial
seabirds is, however, affected by the rate
at which the young grow to the nest-leaving
stage. This depends mainly on body size
(Fig. 10) and to a certain degree on developmental
type, as some semiprecocial species
grow rather slowly. Certain seabirds with
clutches of one egg grow particularly slowly
(petrels, some alcids, sulids). Several other alcids
with single-egg clutches, however, grow<span class="pagenum"><a name="Page_154" id="Page_154">[Pg 154]</a></span>
at rates normal for semiprecocial chicks
(Fig. 10). Very slow growth may be related to
food stress (Lack 1968; Ricklefs 1968) or to reduction
of reproductive effort in the adults
(discussed later). Contrary to Cody (1973),
slow growth in alcids does not correlate to the
distance adults must commute for food. (Cody
tried to directly compare growth in birds of
different sizes.) Chicks in nocturnal species,
however, tend to have slow growth rates
(Sealy 1973<em>b</em>).</p>

<div class="figcenter" style="width: 700px;">
<img src="images/p_154.jpg" width="700" height="393" alt="" />
<div class="caption"><p><strong>Fig. 10.</strong> Growth rate as a function of body weight. Growth rate <sup>T</sup>10-90 represents the number of days to
grow from 10% to 90% of asymptotic weight (Ricklefs 1968). Data from Ricklefs (1968, 1973), E. H.
Dunn (1973), and Sealy (1973<em>b</em>). Solid circles and regression line, altricial birds; solid triangles, semiprecocial
birds except for seabirds with one-egg clutches; open circles, precocial shorebirds; open triangles,
precocial ducks, rails, and gallinaceous birds; solid squares, alcids with one-egg clutches; and open
squares, northern petrels, gannet, and Manx shearwater <em>(Puffinus puffinus)</em>.</p></div>
</div>

<p>Daily time budgets of adults raising nestlings
also vary widely, depending on the
amount of brooding required, food requirements
of the young, and foraging costs (which
differ in the breeding season from those at
other times of the year).</p>

<p>Nestlings have imperfect control of body
temperature at hatching (Fig. 11) and develop
this capacity only gradually. Altricial birds
are hatched at a particularly undeveloped
stage; e.g., double-crested cormorants attain
reasonable control of body temperature in
moderate ambient temperatures only after
about 14 days (Fig. 11; Table 5). Semiprecocial
seabirds, which are more fully developed
physically at hatching, attain control
of body temperature much sooner, in a matter
of several days, and precocial eiders can thermoregulate
within a few hours after hatching
(Table 5).</p>

<p>Until the age of temperature control, nestlings
must be brooded almost constantly, and
occasional brooding takes place for some time
afterward, especially in severe weather, in all
species studied (Tinbergen 1960; Belopol'skii
1961; Weaver 1970; Dunn 1976<em>a</em>, 1976<em>b</em>).
Thermoregulatory capabilities in cold
weather are better in ducklings of species
nesting at high latitudes than at lower ones
(Koskimies and Lahti 1964), and the same
may be true of gull species (Dawson et al.
1972). The cooling mechanisms of double-crested
cormorants are better than in the
more northerly distributed pelagic cormorant,
<em>Phalacrocorax pelagicus</em> (Lasiewski and
Snyder 1969). Thus, variation in cost of thermoregulation
due to different environments
may be reduced through adaptation.</p>

<p>Food requirements of the chick depend on<span class="pagenum"><a name="Page_155" id="Page_155">[Pg 155]</a></span>
growth rate, amount of fat deposition, cost of
thermoregulation, degree of activity and
other factors (E. H. Dunn 1973). Estimated
energy budgets for nestling double-crested
cormorants and herring gulls in the same year
and locality (Fig. 12) indicate that these factors
vary according to developmental type,
and comparison with budgets for nonseabird
species suggests wide variation within developmental
types according to the particular
adaptations of each species to its own environment
(E. H. Dunn 1973).</p>

<div class="figcenter" style="width: 543px;">
<img src="images/p_155a.jpg" width="543" height="700" alt="" />
<div class="caption"><p>Fig. 11. Development of thermoregulatory capabilities
in nestling double-crested cormorants. From
Dunn (1976<em>a</em>). Ages at right refer also to corresponding
oxygen consumption data on the left.
Thin diagonal lines show equality between body
and air temperature. All data taken after 2 h of
exposure.</p></div>
</div>

<p>Thus, the energy demands of nestlings are
not easy to predict. Brood size differences
multiply variation in food demand on adults
(except in precocial birds whose young feed
themselves). Energy demands are labile, however,
particularly in requirements for activity
and growth, and adults can frequently raise
young successfully without providing optimum
amounts of food (Spaans 1971; Kadlec
et al. 1969; LeCroy and Collins 1972; Lemmetyinen
1972; Cody 1973; E. H. Dunn and I. L.
Brisbin, manuscript in preparation). Studies
of double-crested cormorants by Dunn (1975<em>b</em>)
and pigeon guillemots <em>(Cepphus columba)</em> by
Koelink (1972) have suggested that each adult
providing optimum amounts of food to a
normal-sized brood would have to approximately
double the amount of food gathered
each day over the amount gathered by nonbreeders.
This relation does not imply, however,
that the time and energy allocation of
the adults would be the same for the two
species.</p>

<div class="figcenter" style="width: 604px;">
<img src="images/p_155b.jpg" width="604" height="700" alt="" />
<div class="caption"><p>Fig. 12. Energy budgets of nestling double-crested
cormorants and herring gulls. Data from E. H.
Dunn (1973) and Brisbin (1965).</p></div>
</div>

<p>Cost-benefit ratios of food gathering in the
nestling period differ from those at other
times. Besides facing increased food demands,
costs of delivery to the nest, and changes in
food availability, the parents' choice of foods
is constrained by the need to forage within
reasonable commuting distance of the nest
and perhaps by concentrated competition
with conspecifics and other seabird species. In
addition, small nestlings are frequently unable
to eat foods normally eaten by adults
(Drent 1965; personal observation). In the
face of these constraints, adults often shift<span class="pagenum"><a name="Page_156" id="Page_156">[Pg 156]</a></span>
food preferences while raising nestlings (Belopol'skii
1961). For example, female mew gulls
in the Barents Sea forage in the tidal zone,
eating more small invertebrates than at other
times of the year, while males continue to forage
at sea and consume larger quantities of
fish (Fig. 13).</p>

<div class="center">
<table border="0" cellpadding="4" cellspacing="0" summary="Table 5. Age of thermoregulatory control in various species of northern seabirds.">
    <caption>Table 5. <em>Age of thermoregulatory control in various species of northern seabirds.</em></caption>
  <tr>
    <th>Species</th>
    <th>Age when moderate temperature control is attained<br />(days)</th>
    <th>Source</th>
  </tr>
  <tr>
    <td>Common eider</td>
    <td class="tdc">0.1-0.3<a name="FNanchor_41_41" id="FNanchor_41_41"></a><a href="#Footnote_41_41" class="fnanchor">[41]</a> </td>
    <td>V. V. Rolnik, in Belopol'skii (1961)</td>
  </tr>
  <tr>
    <td>Herring gull </td>
    <td class="tdc">1.5-2 </td>
    <td>V. V. Rolnik, in Belopol'skii (1961)</td>
  </tr>
  <tr>
    <td>Herring gull </td>
    <td class="tdc">2-3 </td>
    <td>E. H. Dunn (1976<em>b</em>)</td>
  </tr>
  <tr>
    <td>Leach's storm-petrel</td>
    <td class="tdc">[2]</td>
    <td>Ricklefs (1974)</td>
  </tr>
  <tr>
    <td>Mew gull </td>
    <td class="tdc">2-3 </td>
    <td>V. V. Rolnik, in Belopol'skii (1961)</td>
  </tr>
  <tr>
    <td>Lesser black-backed gull </td>
    <td class="tdc">2-3 </td>
    <td>E. K. Barth (in Farner and Serventy 1959)</td>
  </tr>
  <tr>
    <td>Greater black-backed gull </td>
    <td class="tdc">2-3 </td>
    <td>E. K. Barth (in Farner and Serventy 1959)</td>
  </tr>
  <tr>
    <td>Pigeon guillemot </td>
    <td class="tdc">2-4 </td>
    <td>Drent (1965)</td>
  </tr>
  <tr>
    <td>Common tern </td>
    <td class="tdc">3 </td>
    <td>LeCroy and Collins (1972)</td>
  </tr>
  <tr>
    <td>Roseate tern <em>(Sterna dougallii)</em> </td>
    <td class="tdc">3 </td>
    <td>LeCroy and Collins (1972)</td>
  </tr>
  <tr>
    <td>Common murre </td>
    <td class="tdc">3 </td>
    <td>V. V. Rolnik and Yu. M. Kaftonowski<br />(in Sealy 1973<em>b</em>)</td>
  </tr>
  <tr>
    <td>Razorbill <em>(Alca torda)</em> </td>
    <td class="tdc">3 </td>
    <td>V. V. Rolnik and Yu. M. Kaftonowski<br />(in Sealy 1973<em>b</em>)</td>
  </tr>
  <tr>
    <td>Black guillemot </td>
    <td class="tdc">3-4 </td>
    <td>V. V. Rolnik, in Belopol'skii (1961)</td>
  </tr>
  <tr>
    <td>Tufted puffin </td>
    <td class="tdc">3.5<a name="FNanchor_42_42" id="FNanchor_42_42"></a><a href="#Footnote_42_42" class="fnanchor">[42]</a> </td>
    <td>Cody (1973)</td>
  </tr>
  <tr>
    <td>Northern phalarope </td>
    <td class="tdc">4-5<a name="FNanchor_43_43" id="FNanchor_43_43"></a><a href="#Footnote_43_43" class="fnanchor">[43]</a> </td>
    <td>Hilden and Vuolanto (1972)</td>
  </tr>
  <tr>
    <td>Cassin's auklet </td>
    <td class="tdc">5-6 </td>
    <td>Manuwal (1974<em>a</em>)</td>
  </tr>
  <tr>
    <td>Horned puffin<br /><em>(Fratercula corniculata)</em></td>
    <td class="tdc">2-6 </td>
    <td>Sealy (1973<em>a</em>)</td>
  </tr>
  <tr>
    <td>Common puffin </td>
    <td class="tdc">6-7 </td>
    <td>V. V. Rolnik and Yu. M. Kaftonowski<br />(in Sealy 1973<em>b</em>)</td>
  </tr>
  <tr>
    <td>Black-legged kittiwake </td>
    <td class="tdc">6-7 </td>
    <td>V. V. Rolnik, in Belopol'skii (1961)</td>
  </tr>
  <tr>
    <td>Double-crested cormorant </td>
    <td class="tdc">14 </td>
    <td>Dunn (1976<em>a</em>)</td>
  </tr>
  <tr>
    <td>Shag </td>
    <td class="tdc">12-15 </td>
    <td>V. V. Rolnik, in Belopol'skii (1961)</td>
  </tr>
</table></div>

<p>Commuting distances vary tremendously
among species (Fig. 14), but the number of feeding
trips to the nest per day does not correlate
with foraging distance (Cody 1973; Sealy
1973<em>a</em>, 1973<em>b</em>). There is not, therefore, a
simple relationship between time and energy
expenditures of the adults and foraging distances.
Nocturnality, on the other hand, correlates
with reduced feeding rates (usually one
visit to the nest each night). Seabirds feeding
far from the colony tend to show adaptations
for bringing larger amounts of food per visit,
such as carrying more than one fish at a time,
as in tufted puffins, <em>Lunda cirrhata</em>, and rhinoceros
auklets, <em>Cerorhinca monocerata</em>, vs.
guillemots and murres (Richardson 1961;
Cody 1973; Sealy 1973<em>a</em>, 1973<em>b</em>); developing a
sublingual storage pouch, as in Cassin's auklets
(Speich and Manuwal 1974); or concentration
of food into stomach oil, as in petrels and
albatrosses (Ashmole 1971). Commuting
costs are largely eliminated when the young
leave the nest, but only in the alcids does nest
leaving occur long before attainment of full
growth. Early nest leaving may allow adults
and young to disperse to better feeding areas
than are exploitable from the colony site
(Sealy 1973<em>b</em>) and probably involves a major
change in optimal food size and type as well
(Lind 1965).</p>

<p>Patterning of adult time budgets may differ<span class="pagenum"><a name="Page_157" id="Page_157">[Pg 157]</a></span>
between geographical regions. For example,
rhinoceros auklets are nocturnal in the far
north (where the summer night is particularly
short), crepuscular in the Olympic Peninsula,
and mainly diurnal in the Farallon Islands
(Manuwal 1974<em>a</em>).</p>

<div class="figcenter" style="width: 551px;">
<img src="images/p_157a.jpg" width="551" height="700" alt="" />
<div class="caption"><p>Fig. 13. Foraging ranges of a pair of mew gulls during
the breeding season, on a Barents Sea colony.
From Belopol'skii (1961).</p></div>
</div>

<p>Food demands of nestlings have a great influence
on the time and energy allocation of
breeding over nonbreeding seabirds. Because
food is particularly abundant in the reproductive
season, however, one cannot ascertain
whether the vulnerability of breeding birds to
time or energy crises is far different from that
at other times of the year.</p>


<h6>Post-fledging Care</h6>

<p>Little is known about the amount of care
provided by adults to young after they are
fully grown. At least some species, such as
gannets and procellariiformes (Ashmole 1971),
are known to desert their young, whereas
others are known to feed their young, at least
occasionally, for some weeks or months after
they can fly&mdash;e.g., terns and gulls, many alcids,
and shags (Snow 1963; Vermeer 1963;
Drury and Smith 1968; Ashmole and Tovar S.
1968; Potts 1968; Ashmole 1971; LeCroy
1972).</p>

<div class="figcenter" style="width: 700px;">
<img src="images/p_157b.jpg" width="700" height="418" alt="" />
<div class="caption"><p>Fig. 14. Percentage observations of foraging seabirds
at different distances from the nest site.
After Cody (1973). Each vertical bar represents
5% of total observations. Note nonlinear horizontal
scale.</p></div>
</div>


<h4>Annual Time and
Energy Budgets</h4>

<p>The discussion of time and energy allocation
during reproduction was complex and detailed
because so much more is known about
the influences altering budgeting during this
period than during other times of the year. It
is likely that influences on molt and migration
will prove to be equally complicated, once
more is learned about them.</p>

<p>If all data on time and energy allocation for
a single species were known, it would be possible
to make up detailed budgets for birds of
different age, sex, and experience throughout
the year. However, such detailed data have
not been collected for any species. An annual
time budget for male and female yellow-billed
magpie, <em>Pica nuttalli</em> (Verbeek 1972), points
out the great amount of difference between
the sexes (Fig. 15). A time and energy budget
for the reproductive season only (Fig. 16)
shows large differences between two closely
related species, as well as between sexes; it
also indicates the wide difference between the
budgeting of energy as opposed to budgeting
of time. All other time-energy budgets to date
are for nonseabird species and for only a portion
of the annual cycle (Verbeek 1964; Verner<span class="pagenum"><a name="Page_158" id="Page_158">[Pg 158]</a></span>
1965; Schartz and Zimmerman 1971; Stiles
1971; Wolf and Hainsworth 1971; Smith 1973;
Utter and LeFebvre 1973).</p>

<div class="figcenter" style="width: 458px;">
<img src="images/p_158.jpg" width="458" height="700" alt="" />
<div class="caption"><p>Fig. 15. Time budget of male (upper panel) and female
(lower panel) yellow-billed magpies throughout
the year. From Verbeek (1972). Non-labeled
portions in each graph correspond to labeled sections
in the other.</p></div>
</div>

<p>Time-energy budget analysis can be useful
in determining the leeway a bird has in surviving
unusual stress at different times of the
year. For example, a study by Feare et al.
(1974) showed that rooks <em>(Corvus frugilegus)</em>
in the dry part of the summer spent 90% of
15 h of daylight to collect 150 kcal of food
energy. In winter, foraging in snow, the same
birds were able to collect 240 kcal of food in
only 30% of a 10-h day. This suggests that
rooks would be far more vulnerable to unexpected
periods of stress in late summer than
in winter. Such information would clearly be
useful in making management decisions.</p>

<p>A more precise measure of vulnerability, although
much more difficult to determine, is
that of productive energy&mdash;the amount of
caloric intake left over after the birds' cost of
living (metabolic functions and procurement
and processing of food) have been accounted
for. Costs are highest when temperatures are
extremely hot or cold or when food is most difficult
to obtain. Productive energy is highest
in summer (Kendeigh 1972), and that is presumably
why reproduction normally takes
place then. It is unknown whether birds are
more vulnerable to time and energy shortages
in the harder nonbreeding season or in the
breeding season after the extra demands of reproduction
have been accounted for. Vulnerability
may also differ between sexes and
among age groups.</p>

<p>Time-energy studies, although useful in
comparing ecology, determining vulnerability,
and cataloging location of birds, do
have limitations. Careful studies are time-consuming
and are not the best approach to determining
key factors influencing population
increase or decrease. Even when different
kinds of data are being sought, however, it is
worthwhile keeping the time-energy framework
in mind as a "big picture" into which
other facts can be fitted and their significance
considered.</p>


<h4>Life History Strategies</h4>

<p>The study of life history strategies is
largely theoretical, and in the following discussion
I do not comment on current theoretical
arguments. On the other hand, life history
strategies can be regarded as time and energy
allocation on a grand scale, and it therefore
seems appropriate to look briefly at their implications
for seabird management.</p>

<p>Annual reproduction evidently has a negative
effect on resources remaining for other
functions, and may reduce the chances for an
organism to reproduce again in a later season
(Cody 1966, 1971; Williams 1966; Gadgil and
Bossert 1970; Gadgil and Solbrig 1972;
Hussell 1972; Trivers 1972; Calow 1973). If
the chances of survival to another breeding
season are small, the selective advantage lies
with the bird putting the most effort into
early reproduction, in spite of its negative effects
on survival, because future chances of
reproduction are small. If chances of survival<span class="pagenum"><a name="Page_159" id="Page_159">[Pg 159]</a></span>
are good, however, it may be more advantageous
to reduce annual reproductive effort
and allocate resources to other functions.</p>

<div class="figcenter" style="width: 700px;">
<img src="images/p_159.jpg" width="700" height="650" alt="" />
<div class="caption"><p>Fig. 16. Time and energy budgets of male and female red-winged <em>(Agelaius phoeniceus)</em> and tricolored <em>(A. tricolor)</em>
blackbirds in the breeding season. From Orians (1961). Dotted lines show male (M) activity, dashed
lines show female (F) activity, and solid lines show shared activities.</p></div>
</div>

<p>Seabirds are generally long-lived, have
small clutches, and generally delay first
breeding until at least the 2nd year, and
usually longer (Table 6). Phalaropes seem to
differ from this pattern (Hilden and Vuolanto
1972; Howe 1975). Several ecological factors
(not entirely independent) are believed to contribute
to the evolution of the long life and
low reproductive effort pattern favored by
seabirds.</p>

<p>First, if population size is determined
largely by density-dependent mortality, individuals
may be favored that allocate resources
to attaining longer life (and more
chances to reproduce) or insuring greater
chances of survival of their offspring (Murphy
1968; Hairston et al. 1970). Density-independent
mortality, on the other hand, is so unpredictable
that there is no advantage in allocating
resources toward protection against it
(Gadgil and Solbrig 1972).</p>

<p>Two factors closely linked with density-dependence
are high levels of competition, and
perennial difficulties in obtaining food. In
adapting to these difficulties, a bird may be
selected which develops more efficient foraging
techniques, wider dispersal, or better abili<span class="pagenum"><a name="Page_160" id="Page_160">[Pg 160]</a></span>ties
to defend nesting territory&mdash;all of which
may reduce resources available for reproduction.
As mentioned earlier, marine foods tend
to be patchily distributed, and a long learning
period seems to be necessary before seabirds
become proficient at foraging. In addition,
there is evidence that food availability is low,
at least in the tropics, and perhaps in the winter
in other regions (Ashmole 1971). If nesting
places are in short supply, long life may be
favored so that the bird can live long enough
for a place to become vacant. Several authors
feel that competition is a serious factor in the
life of seabirds, both for food (Lack 1966;
Cody 1973) and for nesting space (Snow 1960;
Belopol'skii 1961; Lack 1966; Manuwal
1974<em>b</em>). Others, however, disagree, at least for
the breeding season (e.g., Pearson 1968).</p>

<div class="center">
<table border="0" cellpadding="4" cellspacing="0" summary="Table 6. Life history data for certain northern seabirds.">
    <caption>Table 6. <em>Life history data for certain northern seabirds.</em><a name="FNanchor_44_44" id="FNanchor_44_44"></a><a href="#Footnote_44_44" class="fnanchor">[44]</a></caption>
  <tr>
    <th>Species</th>
    <th>Annual adult survival<br />(%)</th>
    <th>Age at first breeding<br />(years)</th>
    <th>Clutch size</th>
  </tr>
  <tr>
    <td>Fulmar</td>
    <td>94</td>
    <td>7+</td>
    <td>1</td>
  </tr>
  <tr>
    <td>Gannet <em>(Morus bassanus)</em></td>
    <td>94</td>
    <td>(4)-5+</td>
    <td>1</td>
  </tr>
  <tr>
    <td>Manx shearwater</td>
    <td>93-96</td>
    <td>(4)-5+</td>
    <td>1</td>
  </tr>
  <tr>
    <td rowspan="2">Shag</td>
    <td>85 (&#9794;)</td>
    <td rowspan="2">(2)-3</td>
    <td rowspan="2">3-4</td>
  </tr>
  <tr>
    <td>80 (&#9792;)</td>
  </tr>
  <tr>
    <td rowspan="2">Herring gull</td>
    <td rowspan="2">91-96</td>
    <td>3.5 (&#9794;)</td>
    <td rowspan="2">(2)-3</td>
  </tr>
  <tr>
    <td>5 (&#9792;)</td>
  </tr>
  <tr>
    <td rowspan="2">Black-legged kittiwake</td>
    <td rowspan="2">88</td>
    <td>4-5 (&#9794;)</td>
    <td rowspan="2">3</td>
  </tr>
  <tr>
    <td>3-4 (&#9792;)</td>
  </tr>
  <tr>
    <td rowspan="3">Arctic tern</td>
    <td>89-91</td>
    <td rowspan="3">(2)-3+</td>
    <td rowspan="3">2</td>
  </tr>
  <tr>
    <td>75</td>
  </tr>
  <tr>
    <td>82<a name="FNanchor_45_45" id="FNanchor_45_45"></a><a href="#Footnote_45_45" class="fnanchor">[45]</a></td>
  </tr>
  <tr>
    <td>Common murre</td>
    <td>87</td>
    <td>3+?</td>
    <td>1</td>
  </tr>
  <tr>
    <td>Black guillemot</td>
    <td>88+<a name="FNanchor_46_46" id="FNanchor_46_46"></a><a href="#Footnote_46_46" class="fnanchor">[46]</a></td>
    <td>3?<a href="#Footnote_46_46" class="fnanchor">[46]</a></td>
    <td>2<a href="#Footnote_46_46" class="fnanchor">[46]</a></td>
  </tr>
  <tr>
    <td>Cassin's auklet</td>
    <td>83<a name="FNanchor_47_47" id="FNanchor_47_47"></a><a href="#Footnote_47_47" class="fnanchor">[47]</a></td>
    <td>3<a href="#Footnote_47_47" class="fnanchor">[47]</a></td>
    <td>1</td>
  </tr>
</table></div>

<p>There is some evidence of density-dependent
population size control in seabirds,
although much of it is circumstantial. For
example, there are large nonbreeding populations
in such diverse species as shags, herring
gulls, and Cassin's auklets, which move into a
breeding area when established adults are removed
or colonize new breeding areas (J. C.
Coulson, personal communication; Kadlec and
Drury 1968; Drury and Nisbet 1972; Manuwal
1974<em>b</em>). Lack (1966) and Ashmole (1971) presented
other arguments for density-dependence.
Density-dependent mortality is difficult
to demonstrate, at best, and may be obscured
by interpopulation movements (Drury
and Nisbet 1972).</p>

<p>If long life is a life history option, a low
annual reproductive effort could be favored in
several ways. First, it may be necessary for insuring
long life, if breeding has a serious negative
feedback on life expectancy (Calow 1973).
Second, if survival of offspring is more unpredictable
than that of adults, low annual effort
may be selected so that reproductive effort
will not be wasted in years when young have
poor chances of survival. Unpredictable and
variable first-year survival in seabirds has
been documented (Potts 1968; Drury and Nisbet
1972). In addition, some seabirds show
adaptations that allow high reproductive success
in any given year but which do not drain
off resources if the season turns out to be poor
(e.g., small last eggs in the clutch or asynchronous
hatching, both of which lead to elimination
of the smallest chicks when conditions
are poor [Parsons 1970; E. H. Dunn 1973]).</p>

<p>It should be emphasized that the factors involved
in the evolution of life histories are<span class="pagenum"><a name="Page_161" id="Page_161">[Pg 161]</a></span>
complex and poorly understood, and simple
formulas should not be expected to apply to
all situations (Wilbur et al. 1974).</p>

<p>In the framework of life-history strategies,
small clutch sizes and slow growth rates exhibited
by some seabirds can be explained as
adaptive reductions in annual reproductive effort,
rather than as responses to immediate
food shortages. Arguments for this view are
presented on theoretical grounds (Dunn 1973)
and by the fact that many seabirds are able to
raise larger than normal broods in certain
situations (Vermeer 1963; Nelson 1964;
Harris 1970; Hussell 1972; Ward 1972; Corkhill
1973). In addition, seabirds with particularly
slow growth rates all grow at about the
same rate, regardless of body size (contrary to
the situation in other birds). This suggests
that low growth rates do not reflect variations
in feeding abilities among species (Ricklefs
1968).</p>

<p>Several conclusions relating to management
of seabird populations can be drawn
from the above discussion. First, if population
size is determined largely by density-dependent
factors, the birds are not adapted to
precipitous and unexpected declines in population
levels. Because there is low annual reproductive
effort geared to a world in which
there is slow turnover in population, seabirds
are not able to rebound quickly from
disasters. Provision of excess food should not
be expected to improve breeding performance,
at least in experienced birds.</p>

<p>Second, because seabirds are able to reproduce
in many different seasons and are
adapted to a low reproductive effort within a
given season, one should expect them to be
easily disturbed and to fail to complete the reproductive
cycle during any given breeding
attempt. A few indications of such failures
have already been observed (Erskine 1972;
Manuwal 1974<em>a</em>; Nettleship 1975).</p>

<p>Again, the tentative nature of this discussion
should be emphasized, and conclusions
drawn from it may not apply equally to all
seabird species.</p>


<h4>Conclusions</h4>

<p>In this discussion I have tried to emphasize
the variety of factors affecting seabird life
cycles and the diverse responses among different
species to their environment. The main
conclusion I stress is that each species (and
age group and sex within that species) has a
different vulnerability to stress, which may be
most severe at different times of the year for
each group. To determine these periods of
stress, researchers may find a time-energy approach
to be useful.</p>

<p>As for northwestern North American seabirds
in particular, ignorance is vast. Twelve
years ago, Bourne (1963:846) noted the following
needs in seabird research (among others):
"The investigation of seabird biology has
been reduced to a routine, but there is a great
need for more study of some other aspects of
the life or annual cycle, including events in the
period immediately after fledging, and behaviour
and survival in the immature period
and outside the breeding season. Much more
accurate information is needed about breeding
distribution and seasons in many parts of
the world, about molting seasons and ranges
in most parts, and the distribution of birds of
different age groups during these periods in
practically all areas."</p>

<p>Since the time of Bourne's remarks, a number
of excellent studies have provided data on
the breeding biology of certain northwestern
seabird species. Scientists remain largely
ignorant, however, about where birds of different
age groups are located throughout the
year. Such knowledge is necessary for effective
protection and is basic to understanding
population dynamics, even if it does not elucidate
causes. Studies of timing of annual
cycles and movements should be carried out
hand in hand with resource analysis&mdash;not just
finding what birds eat, but discovering where
the food is at what times, how hard it is to
catch, and what the nutritional return is.
Much careful field work must be done before
effective management of most of our northwestern
seabirds can become a reality.</p>


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<div class="footnotes"><h4>FOOTNOTES:</h4>

<div class="footnote">

<p><a name="Footnote_41_41" id="Footnote_41_41"></a><a href="#FNanchor_41_41"><span class="label">[41]</span></a> Common eider, 2 to 7 h.</p></div>

<div class="footnote">

<p><a name="Footnote_42_42" id="Footnote_42_42"></a><a href="#FNanchor_42_42"><span class="label">[42]</span></a> No data given.</p></div>

<div class="footnote">

<p><a name="Footnote_43_43" id="Footnote_43_43"></a><a href="#FNanchor_43_43"><span class="label">[43]</span></a> Indirect evidence that young are brooded this long.</p></div>

<div class="footnote">

<p><a name="Footnote_44_44" id="Footnote_44_44"></a><a href="#FNanchor_44_44"><span class="label">[44]</span></a> Data from Lack (1968) and Ashmole (1971) unless otherwise noted.</p></div>

<div class="footnote">

<p><a name="Footnote_45_45" id="Footnote_45_45"></a><a href="#FNanchor_45_45"><span class="label">[45]</span></a> Cullen (1957).</p></div>

<div class="footnote">

<p><a name="Footnote_46_46" id="Footnote_46_46"></a><a href="#FNanchor_46_46"><span class="label">[46]</span></a> Birkhead (1974).</p></div>

<div class="footnote">

<p><a name="Footnote_47_47" id="Footnote_47_47"></a><a href="#FNanchor_47_47"><span class="label">[47]</span></a> Speich and Manuwal (1974).</p>

<p><span class="pagenum"><a name="Page_167" id="Page_167">[Pg 167]</a></span></p></div></div>
<div class="chapter"></div>
<hr class="chap" />




<h3><a name="Zoogeography_and_Taxonomic_Relationships_of" id="Zoogeography_and_Taxonomic_Relationships_of">Zoogeography and Taxonomic Relationships of
Seabirds in Northern North America</a></h3>


<p class="center">by</p>

<p class="center">M. D. F. Udvardy</p>

<p class="center">California State University<br />
Sacramento, California 95819
</p>


<h4>Abstract</h4>

<div class="blockquot">

<p>The zoogeography and taxonomic relationships among 42 living and 1 extinct
species of marine birds from the northern and northwestern coasts of North
America are described. Seventeen species are circumpolar in distribution; 17 are
endemic to Beringia, and 8 have origins in the North Pacific.</p></div>

<p>This discussion concerns the northern and
western coasts of the continent, from about
the Mackenzie Delta westward and southward
to the mouth of the Columbia River. Besides
bona fide seabirds, I include marine
birds that predominantly breed and feed on or
around the marine littoral, but exclude two
groups: shorebirds, jaegers, and phalaropes,
which breed inland and move out from the
Arctic after an undetermined postbreeding
period; and Anseriformes which become
"marine birds" in their southern winter quarters.
What remains is 42 living species
(Table 1).</p>

<p>The Procellariiformes, or tube-nosed seabirds,
have a predominantly southern hemispheric,
Gondwanan distribution. The North
Pacific basin is an important feeding ground
of several shearwaters (<em>Puffinus</em> spp.) that
breed in the South Pacific and subantarctic.
Only three species breed in the area under consideration:
the fulmar <em>(Fulmarus glacialis)</em>
and two storm-petrels (<em>Oceanodroma</em> spp.), all
of which are still relatively widespread.</p>

<p>Of the Pelecaniformes, the very successful,
worldwide cormorants (<em>Phalacrocorax</em> spp.)&mdash;inland
water as well as coastal and "amphibious"
species are on every continent&mdash;are
ancient Pacific dwellers, with a high grade of
endemism here: Of the two subarctic species,
one <em>(P. perspicillatus)</em> became extinct long
ago, and the other, the red-faced cormorant
<em>(P. urile)</em>, is very restricted, and deserves our
greatest attention. The pelagic cormorant
<em>(P. pelagicus)</em>, Brandt's cormorant <em>(P. penicillatus)</em>,
and the double-crested cormorant
<em>(P. auritus)</em> are widespread and successful, extending
south of the area here considered;
double-crested cormorants also breed inland
and across toward the North Atlantic coast.
As fish-eaters they are often persecuted where
coastal fishermen possess firearms, and thus
are sensitive to increasing human influence on
the coasts.</p>

<p>Two species of arctic geese need special attention.
The emperor goose <em>(Philacte canagica)</em>
is a Beringean endemic and lives in a
very restricted area of both sides of this sea;
its status (endangered?) is unknown to me.
Since the black brant <em>(Branta bernicla)</em> is a
long-range migrant, it is hunted as a game
bird at its winter grounds, and subject to
management measures. Whereas the emperor
goose is a unique offshoot of the genus Anser,
the Pacific brant is considered a subspecies;
its general distribution is circumpolar.</p>

<p>Five arctic ducks, and one other, constitute
the sea ducks of the area. The common eider
<em>(Somateria mollissima)</em>, king eider <em>(S. spectabilis)</em>,
and the oldsquaw <em>(Clangula hyemalis)</em>
are widespread, and circumpolar or nearly so;
hunting and down-robbing in other parts of
the Arctic may provide clues as to their relative
tolerance of primitive or advanced civilization.
The spectacled eider <em>(S. fischeri)</em> and
Steller's eider <em>(Polysticta stelleri)</em> are restricted
to the Bering Sea coasts and neighboring
High Arctic coasts, respectively; their
status is precarious.</p>

<p><span class="pagenum"><a name="Page_168" id="Page_168">[Pg 168]</a></span></p>

<div class="center">
<table border="0" cellpadding="4" cellspacing="0" summary="Table 1. Seabirds in northwestern North America.">
    <caption>Table 1. <em>Seabirds in northwestern North America.</em> (x = breeding, w = wintering or transient, () = either scarce or restricted distribution, * = stragglers only, nesting status unclear)</caption>
  <tr>
    <th rowspan="2">Species</th>
    <th colspan="0">Distribution</th>
  </tr>
  <tr>
    <th>Circumpolar</th>
    <th>Widespread in North Pacific</th>
    <th>North coast of Alaska</th>
    <th>Beringia<a name="FNanchor_48_48" id="FNanchor_48_48"></a><a href="#Footnote_48_48" class="fnanchor">[48]</a></th>
    <th>Aleutian Islands</th>
    <th>South coast of Alaska<a name="FNanchor_49_49" id="FNanchor_49_49"></a><a href="#Footnote_49_49" class="fnanchor">[49]</a></th>
    <th>Temperate northeast Pacific coast<a name="FNanchor_50_50" id="FNanchor_50_50"></a><a href="#Footnote_50_50" class="fnanchor">[50]</a></th>
  </tr>
  <tr>
    <td><em>Fulmarus glacialis</em></td>
    <td class="tdc"> x</td>
    <td class="tdc"> w</td>
    <td></td>
    <td class="tdc"> x</td>
    <td class="tdc"> x</td>
    <td class="tdc"> x</td>
    <td class="tdc"> w</td>
  </tr>
  <tr>
    <td><em>Oceanodroma furcata</em></td>
    <td></td>
    <td class="tdc"> x</td>
    <td></td>
    <td></td>
    <td class="tdc"> x</td>
    <td class="tdc"> x</td>
    <td class="tdc"> x</td>
  </tr>
  <tr>
    <td><em>O. leucorhoa</em></td>
    <td class="tdc"> x</td>
    <td class="tdc"> x</td>
    <td></td>
    <td></td>
    <td class="tdc"> x</td>
    <td class="tdc"> x</td>
    <td class="tdc"> x</td>
  </tr>
  <tr>
    <td><em>Phalacrocorax auritus</em></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc"> x</td>
    <td class="tdc"> x</td>
    <td class="tdc"> x</td>
  </tr>
  <tr>
    <td><em>P. penicillatus</em></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">(x)</td>
    <td class="tdc"> x</td>
  </tr>
  <tr>
    <td><em>P. pelagicus</em></td>
    <td></td>
    <td class="tdc"> x</td>
    <td></td>
    <td class="tdc"> x</td>
    <td class="tdc"> x</td>
    <td class="tdc"> x</td>
    <td class="tdc"> x</td>
  </tr>
  <tr>
    <td><em>P. urile</em></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc"> x</td>
    <td class="tdc"> x</td>
    <td class="tdc"> x</td>
  </tr>
  <tr>
    <td><em>Branta bernicla</em></td>
    <td class="tdc"> x</td>
    <td></td>
    <td class="tdc"> x</td>
    <td class="tdc"> x</td>
    <td class="tdc">(w)</td>
    <td class="tdc">(w)</td>
    <td class="tdc"> w</td>
  </tr>
  <tr>
    <td><em>Anser canagicus</em></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc"> x</td>
    <td class="tdc"> w</td>
    <td class="tdc"> w</td>
  </tr>
  <tr>
    <td><em>Clangula hyemalis</em></td>
    <td class="tdc"> x</td>
    <td class="tdc"> w</td>
    <td class="tdc"> x</td>
    <td class="tdc"> x</td>
    <td class="tdc"> w</td>
    <td class="tdc"> w</td>
    <td class="tdc"> w</td>
  </tr>
  <tr>
    <td><em>Histrionicus histrionicus</em></td>
    <td class="tdc"> x</td>
    <td class="tdc"> w</td>
    <td></td>
    <td class="tdc"> w</td>
    <td class="tdc"> w</td>
    <td class="tdc"> w</td>
    <td class="tdc"> w</td>
  </tr>
  <tr>
    <td><em>Polysticta stelleri</em></td>
    <td></td>
    <td></td>
    <td class="tdc"> x</td>
    <td class="tdc"> x</td>
    <td></td>
    <td class="tdc">(w)</td>
  </tr>
  <tr>
    <td><em>Somateria mollissima</em></td>
    <td class="tdc"> x</td>
    <td></td>
    <td class="tdc"> x</td>
    <td class="tdc"> x</td>
    <td class="tdc"> x</td>
    <td class="tdc"> x</td>
  </tr>
  <tr>
    <td><em>S. spectabilis</em></td>
    <td class="tdc"> x</td>
    <td></td>
    <td class="tdc"> x</td>
    <td class="tdc"> x</td>
    <td class="tdc"> w</td>
    <td class="tdc">(w)</td>
  </tr>
  <tr>
    <td><em>S. fischeri</em></td>
    <td></td>
    <td></td>
    <td class="tdc"> x</td>
    <td class="tdc"> x</td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td><em>Larus hyperboreus</em></td>
    <td class="tdc"> x</td>
    <td class="tdc"> w</td>
    <td class="tdc"> x</td>
    <td class="tdc"> x</td>
    <td class="tdc"> w</td>
    <td class="tdc"> w</td>
    <td class="tdc"> w</td>
  </tr>
  <tr>
    <td><em>L. glaucescens</em></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc"> x</td>
    <td class="tdc"> x</td>
    <td class="tdc"> x</td>
    <td class="tdc"> x</td>
  </tr>
  <tr>
    <td><em>L. occidentalis</em></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">(x)</td>
  </tr>
  <tr>
    <td><em>L. argentatus</em></td>
    <td class="tdc"> x</td>
    <td class="tdc"> w</td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc"> x</td>
    <td class="tdc">(x)w</td>
  </tr>
  <tr>
    <td><em>L. thayeri</em></td>
    <td></td>
    <td class="tdc"> w</td>
    <td class="tdc"> x</td>
    <td class="tdc"> w</td>
    <td class="tdc"> w</td>
    <td class="tdc"> w</td>
    <td class="tdc"> w</td>
  </tr>
  <tr>
    <td><em>L. canus</em></td>
    <td class="tdc"> x</td>
    <td class="tdc"> w</td>
    <td class="tdc"> x</td>
    <td></td>
    <td></td>
    <td class="tdc"> x</td>
    <td class="tdc">(x)w</td>
  </tr>
  <tr>
    <td><em>Rissa tridactyla</em></td>
    <td class="tdc"> x</td>
    <td class="tdc"> w</td>
    <td class="tdc"> x</td>
    <td class="tdc"> x</td>
    <td class="tdc"> x</td>
    <td class="tdc"> x</td>
    <td class="tdc"> w</td>
  </tr>
  <tr>
    <td><em>R. brevirostris</em></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc"> x</td>
    <td class="tdc"> x</td>
    <td class="tdc">(x)</td>
  </tr>
  <tr>
    <td><em>Xema sabini</em></td>
    <td class="tdc"> x</td>
    <td></td>
    <td class="tdc"> x</td>
    <td class="tdc"> x</td>
    <td></td>
    <td></td>
    <td class="tdc"> w</td>
  </tr>
  <tr>
    <td><em>Sterna paradisaea</em></td>
    <td class="tdc"> x</td>
    <td class="tdc"> w</td>
    <td class="tdc"> x</td>
    <td class="tdc"> x</td>
    <td class="tdc"> x</td>
    <td></td>
    <td class="tdc"> w</td>
  </tr>
  <tr>
    <td><em>S. aleutica</em></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc"> x</td>
    <td></td>
    <td class="tdc"> x</td>
  </tr>
  <tr>
    <td><em>Uria aalge</em></td>
    <td class="tdc"> x</td>
    <td class="tdc"> x</td>
    <td class="tdc">(x)</td>
    <td class="tdc"> x</td>
    <td class="tdc"> x</td>
    <td class="tdc"> x</td>
    <td class="tdc"> x</td>
  </tr>
  <tr>
    <td><em>U. lomvia</em></td>
    <td class="tdc"> x</td>
    <td class="tdc"> x</td>
    <td class="tdc"> x</td>
    <td class="tdc"> x</td>
    <td class="tdc"> x</td>
    <td class="tdc"> x</td>
  </tr>
  <tr>
    <td><em>Alle alle</em></td>
    <td class="tdc"> x</td>
    <td></td>
    <td></td>
    <td class="tdc"> *</td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td><em>Cepphus grylle</em></td>
    <td class="tdc"> x</td>
    <td></td>
    <td class="tdc"> x</td>
    <td class="tdc"> w</td>
    <td></td>
    <td></td>
  </tr>
  <tr>
    <td><em>C. columba</em></td>
    <td></td>
    <td class="tdc"> x</td>
    <td></td>
    <td class="tdc"> x</td>
    <td class="tdc"> x</td>
    <td class="tdc"> x</td>
    <td class="tdc"> x</td>
  </tr>
  <tr>
    <td><em>Brachyramphus marmoratus</em></td>
    <td></td>
    <td class="tdc"> x</td>
    <td></td>
    <td></td>
    <td class="tdc">(x)</td>
    <td class="tdc"> x</td>
    <td class="tdc"> x</td>
  </tr>
  <tr>
    <td><em>B. brevirostris</em></td>
    <td></td>
    <td></td>
    <td class="tdc"> x</td>
    <td class="tdc"> x</td>
    <td class="tdc"> x</td>
    <td></td>
  </tr>
  <tr>
    <td><em>Synthliboramphus antiquus</em></td>
    <td></td>
    <td class="tdc"> x</td>
    <td></td>
    <td class="tdc"> x</td>
    <td class="tdc"> x</td>
    <td class="tdc"> x</td>
    <td class="tdc"> x</td>
  </tr>
  <tr>
    <td><em>Ptychoramphus aleuticus</em></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc"> x</td>
    <td class="tdc"> x</td>
    <td class="tdc"> x</td>
  </tr>
  <tr>
    <td><em>Cyclorrhynchus psittacula</em></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc"> x</td>
    <td class="tdc"> x</td>
    <td></td>
  </tr>
  <tr>
    <td><em>Aethia cristatella</em></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc"> x</td>
    <td class="tdc"> x</td>
    <td></td>
  </tr>
  <tr>
    <td><em>A. pusilla</em></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc"> x</td>
    <td class="tdc"> x</td>
    <td></td>
  </tr>
  <tr>
    <td><em>A. pygmaea</em></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc"> x</td>
    <td></td>
  </tr>
  <tr>
    <td><em>Cerorhinca monocerata</em></td>
    <td></td>
    <td class="tdc"> x</td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc"> x</td>
    <td class="tdc"> x</td>
  </tr>
  <tr>
    <td><em>Fratercula corniculata</em></td>
    <td></td>
    <td class="tdc"> x</td>
    <td></td>
    <td class="tdc"> x</td>
    <td class="tdc"> x</td>
    <td class="tdc"> x</td>
    <td class="tdc"> x</td>
  </tr>
  <tr>
    <td><em>Lunda cirrhata</em></td>
    <td></td>
    <td class="tdc"> x</td>
    <td></td>
    <td class="tdc"> x</td>
    <td class="tdc"> x</td>
    <td class="tdc"> x</td>
    <td class="tdc"> x</td>
  </tr>
  <tr>
    <td>Total number of nesting species</td>
    <td class="tdc"> 17</td>
    <td class="tdc"> 11</td>
    <td class="tdc"> 15</td>
    <td class="tdc"> 27</td>
    <td class="tdc"> 25</td>
    <td class="tdc"> 24</td>
    <td class="tdc"> 17</td>
  </tr>
  <tr>
    <td>Total number of wintering species</td>
    <td></td>
    <td class="tdc"> 9</td>
    <td></td>
    <td class="tdc"> 4</td>
    <td class="tdc"> 7</td>
    <td class="tdc"> 9</td>
    <td class="tdc"> 9</td>
  </tr>
  <tr>
    <td>Grand total </td>
    <td class="tdc">17</td>
    <td class="tdc"> 20</td>
    <td class="tdc"> 15</td>
    <td class="tdc"> 31</td>
    <td class="tdc"> 32</td>
    <td class="tdc"> 33</td>
    <td class="tdc"> 26</td>
  </tr>
</table></div>

<div class="pagenum"><a name="Page_169" id="Page_169">[Pg 169]</a></div>

<p>The harlequin duck <em>(Histrionicus histrionicus)</em>
stands alone without close relatives. It
often breeds far from the sea, but spends the
shortest time&mdash;only a few weeks&mdash;away from
the rocky coast. There is a year-round population
of yearlings in the sea. The drakes of the
nearest breeding pairs at lower latitudes are
back to the sea, abandoning their mates at the
breeding stream when the alpine stream-dwellers
are still at sea awaiting the thawing
of their breeding grounds. Harlequin ducks
live in large parts of Siberia, from arctic
Alaska to central California and Colorado,
and also in the eastern Arctic. They do not
seem to me to be in immediate danger
globally, though perhaps they are locally.</p>

<p>Gulls are a highly successful group of seabirds,
and of the eight species on our coasts
the four more southern ones&mdash;the western gull
<em>(Larus occidentalis)</em>, glaucous-winged gull
<em>(L. glaucescens)</em>, common gull <em>(L. canus)</em>, and
herring gull <em>(L. argentatus)</em>&mdash;are expanding
wherever civilization creates new scavenging
opportunities. Nothing is said about the populations
of the kittiwake <em>(Rissa tridactyla)</em>,
black-legged kittiwake <em>(R. brevirostris)</em> and
Sabine's gull <em>(Xema sabini)</em>, or of the other
two high arctic species (<em>Pagophila eburnea</em>,
<em>Rhodostethia rosea</em>) which do not nest regularly
in the area considered here.</p>

<p>The arctic tern <em>(Sterna paradisaea)</em> is circumpolarly
widespread and successful, whereas
the Aleutian tern <em>(S. aleutica)</em> is a very restricted
Beringean endemic, and its status
needs to be exactly known.</p>

<p>Almost one-third of the seabirds in this area
are alcids, a family centered in the North
Pacific and, more specifically, in the Bering
Sea. Most species breed in enormous
rookeries. Any impact of civilization is highly
detrimental under such circumstances. Of the
four circumpolar species the two <em>Uria</em> guillemots
(murres) are important. The dovekie
<em>(Alle alle)</em> is a sparse pioneer of Bering Strait,
as is the black guillemot <em>(Cepphus grylle)</em> on
our side of the Arctic Sea. Its congener, the
pigeon guillemot <em>(C. columba)</em>, is common and
successful all the way to coastal central California.
Of the remaining 11 species, special attention
should be paid to the whiskered auklet
<em>(Aethia pygmaea)</em> of the Aleutian chain; the
Kittlitz's murrelet <em>(Brachyramphus brevirostris)</em>
of the eastern Beringean and southern
Alaska coast; and to the widespread, but very
sporadic rhinoceros auklet, or puffin <em>(Cerorhinca
monocerata)</em>.</p>

<p>To sum up, I have tabulated these 42
species, and indicated whether modern life-history
and population studies are extant:</p>

<div class="center">
<table border="0" cellpadding="4" cellspacing="0" summary="modern life-history
and population studies">
  <tr>
    <th></th>
    <th>No. species</th>
    <th>No. studied</th>
  </tr>
  <tr>
    <td>Procellariiformes</td>
    <td class="tdr">3</td>
    <td class="tdr">2</td>
  </tr>
  <tr>
    <td><em>Phalacrocorax</em></td>
    <td class="tdr">4</td>
    <td class="tdr">2</td>
  </tr>
  <tr>
    <td>Anseres</td>
    <td class="tdr">2</td>
    <td class="tdr">1</td>
  </tr>
  <tr>
    <td>Anates</td>
    <td class="tdr">6</td>
    <td class="tdc">&mdash;</td>
  </tr>
  <tr>
    <td>Lari</td>
    <td class="tdr">9</td>
    <td class="tdr">2</td>
  </tr>
  <tr>
    <td>Sterni</td>
    <td class="tdr">2</td>
    <td class="tdc">&mdash;</td>
  </tr>
  <tr>
    <td>Alcidae</td>
    <td class="tdr">16</td>
    <td class="tdr">7</td>
  </tr>
  <tr>
    <td class="tdp">Total</td>
    <td class="tdr">42</td>
    <td class="tdr">14</td>
  </tr>
</table></div>

<p>Thus, 28 species await studies preliminary to,
and highly necessary for, conservation
measures.</p>

<p>Seventeen species of marine birds are
spread either circumpolarly around the northern
perimeter or along the north-south coasts
of the Laurasian continents. Four of these are
of the High Arctic (<em>Branta bernicla</em>, <em>Somateria
spectabilis</em>, <em>Xema sabini</em>, <em>Alle alle</em>); another
seven penetrate the Bering Sea as well (<em>Fulmarus
glacialis</em>, <em>Somateria mollissima</em>, <em>Clangula
hyemalis</em>, <em>Larus hyperboreus</em>, <em>Rissa tridactyla</em>,
<em>Sterna paradisaea</em>, <em>Uria lomvia</em>); and
six are panboreal-subboreal, widespread in
their distribution&mdash;<em>Oceanodroma leucorhoa</em>
(extends far south), <em>Histrionicus histrionicus</em>,
<em>Larus argentatus</em> (widespread latitudinally),
<em>L. canus</em> (also inland), <em>Uria aalge</em>, and <em>Cepphus
grylle</em>.</p>

<p>Seventeen species of marine birds are endemic
to Beringia: <em>Anser canagicus</em>, <em>Polysticta
stelleri</em>, <em>Somateria fischeri</em>, <em>Rissa brevirostris</em>,
and <em>Aethia pusilla</em> (and the extinct
<em>Phalacrocorax perspicillatus</em>); <em>P. urile</em>, <em>Sterna
aleutica</em>, <em>Aethia pygmaea</em>, <em>A. cristatella</em>, and
<em>Cyclorrhynchus</em> extend westward to the Sea
of Okhotsk, as do <em>Brachyramphus brevirostris</em>
and <em>Larus glaucescens</em>, which also extend
eastward; and <em>Phalacrocorax pelagicus</em>,
<em>Cepphus columba</em>, <em>Fratercula corniculata</em>, and<span class="pagenum"><a name="Page_170" id="Page_170">[Pg 170]</a></span>
<em>Lunda cirrhata</em> are amphipacific species in
Beringia.</p>

<p>Eight species of marine birds are associated
with the North Pacific. Four are found on
both sides of the ocean&mdash;<em>Oceanodroma furcata</em>,
<em>Brachyramphus marmoratus</em>, <em>Synthliboramphus
antiquus</em>, and <em>Cerorhinca monocerata</em>
(very disjunct). The four others occur
on only the North American side&mdash;<em>Phalacrocorax
auritus</em> (also inland), <em>P. penicillatus</em>,
<em>Larus occidentalis</em> (albeit barely), and <em>Ptychoramphus
aleuticus</em>.</p>

<p>Finally, one species, <em>Larus thayeri</em>, is endemic
at the central Canadian Arctic, extending
westward into the area here considered.</p>

<div class="footnotes"><h4>FOOTNOTES:</h4>

<div class="footnote">

<p><a name="Footnote_48_48" id="Footnote_48_48"></a><a href="#FNanchor_48_48"><span class="label">[48]</span></a> Beringia comprises the islands and coasts of the Bering Sea.</p></div>

<div class="footnote">

<p><a name="Footnote_49_49" id="Footnote_49_49"></a><a href="#FNanchor_49_49"><span class="label">[49]</span></a> South coast of Alaska extends from the tip of the Alaska Peninsula to Glacier Bay.</p></div>

<div class="footnote">

<p><a name="Footnote_50_50" id="Footnote_50_50"></a><a href="#FNanchor_50_50"><span class="label">[50]</span></a> Temperate northeast Pacific coast extends from Glacier Bay south to the mouth of the Columbia River.</p>

<p><span class="pagenum"><a name="Page_171" id="Page_171">[Pg 171]</a></span></p></div></div>

<p><span class="pagenum"><a name="Page_172" id="Page_172">[Pg 172]</a><br /><a name="Page_173" id="Page_173">[Pg 173]</a></span></p>
<div class="chapter"></div>
<hr class="chap" />




<h2><a name="CONFLICTS_BETWEEN_THE_CONSERVATION_OF" id="CONFLICTS_BETWEEN_THE_CONSERVATION_OF">CONFLICTS BETWEEN THE CONSERVATION OF
MARINE BIRDS AND USES OF OTHER RESOURCES</a></h2>
<div class="chapter"></div>
<hr class="chap" />




<h3>Social and Economic Values of Marine Birds</h3>


<p class="center">by</p>

<p class="center">David R. Cline<a name="FNanchor_51_51" id="FNanchor_51_51"></a><a href="#Footnote_51_51" class="fnanchor">[51]</a> and Cynthia Wentworth</p>

<p class="center">U.S. Fish and Wildlife Service<br />
Anchorage, Alaska</p>

<p class="center">and</p>

<p class="center">Thomas W. Barry</p>

<p class="center">Canadian Wildlife Service<br />
Edmonton, Alberta, Canada
</p>


<h4>Abstract</h4>

<div class="blockquot">

<p>Throughout history, marine birds have provided tangible and intangible benefits
to human societies. Unregulated exploitation of some species by explorers,
mariners, and colonists led to the extinction of the great auk <em>(Pinguinus impennis)</em>
and near extinction of others, including the Bermuda petrel <em>(Pterodroma
cahow)</em> and the North Pacific albatrosses (<em>Diomedea</em> spp.). Marine birds continue
to provide commercial, subsistence, recreational, scientific, and educational
values to people of many nations, while playing critical roles in the economies of
the world's oceans.</p>

<p>Annual harvest of slender-billed shearwaters <em>(Puffinus tenuirostris)</em> known as
"muttonbirds" in Australia, sooty tern <em>(Sterna fuscata)</em> eggs in the Caribbean,
murres (<em>Uria</em> spp.) and eiders (<em>Somateria</em> spp.) in Greenland and the Soviet
Union, and guano in Peru and Africa represent the principal commercial uses of
marine birds and their products. Residents of the Faeroes Islands and thousands
of native people in Greenland and arctic Canada and Alaska use various species
for subsistence. The annual rituals of bird hunting and egg gathering are deeply
ingrained in the sociocultural traditions of these peoples and continue to be
important to their social welfare.</p>

<p>Most countries of the world are currently providing at least some protection to
their marine bird resources. However, the destruction of bird habitats by man's
developments and the contamination of marine environments by industrial pollutants
are posing increasingly serious threats to many species. If managed and
used in accordance with scientific principles of sustained yield, some of the more
abundant species of marine birds can continue to provide long-term social and
economic benefits to man.</p>

<p>Increasing numbers of people are expending considerable sums of money to
reach marine bird viewing areas off the coasts of North American States and
Provinces. Preliminary evidence indicates such nonconsumptive pursuits are
contributing significant amounts of money to regional economies and helping
businessmen earn a living. An accurate evaluation of both biological and economic
impacts resulting from these nonconsumptive activities is urgently
needed.</p>

<p>The possibility of establishing an excise tax on designated outdoor recreational
equipment appears to hold considerable potential for more adequately funding
marine bird programs, as well as those for other nongame wildlife.</p>

<p>Greater citizen involvement in sociopolitical processes will, to a large extent,
determine the success of marine bird conservation programs. Sound conservation
legislation that insures adequate protection of habitat and provides for enlightened
and innovative thrusts in conservation, education, research, management,
and law enforcement will help insure the survival of all species of marine
birds and, in turn, provide social and economic benefits to people across
generations.</p></div>
<p><span class="pagenum"><a name="Page_174" id="Page_174">[Pg 174]</a></span></p>
<p>In its 17 March 1975 issue, <cite>Time</cite> magazine
reported battalions of observers from all over
the country flocking to Salisbury, Massachusetts,
armed with telescopes, cameras
dwarfed by huge telephoto lenses, sketch
pads, and binoculars. There, 1,500 strong the
first weekend alone, they took up vigil along
the seawall of the Merrimack River. A local
businessman circulated among the chilly bird-watchers
with free coffee and hot chocolate,
while handing out a pamphlet advertising his
restaurant.</p>

<p>The cause of the commotion was the appearance
of a single, unassuming, pigeon-like seabird
called a Ross' gull <em>(Rhodostethia rosea)</em>,
almost never seen south of the Arctic Circle
and never before in the contiguous 48 States.
Time stated that "for those who care about
such matters the event was as electrifying as
the descent of a Martian spaceship."</p>

<p>Meanwhile, far above the Arctic Circle at
Point Barrow on the Arctic Ocean, Eskimo
hunters probably puzzled at the strange ways
of the white "birdmen," as they recalled the
savory dishes Ross' gulls provided many of
them during the previous fall hunting season.
This particular gull is considered a delicacy by
the Eskimos, and the birds are actively
sought each year as they fly near shore during
their fall wanderings from Asian breeding
grounds.</p>

<p>Perhaps this dichotomy of people's interests
in a single species is indicative of the
broad spectrum of social and economic values
man derives from marine birds. Perhaps, too,
it represents the challenge that wildlife professionals,
administrators, and citizen conservation
leaders face in today's complex world
in striving to sort out priorities in allocation
of such common property (amenity) resources
among beneficial users.</p>

<p>As with the Ross' gull, socioeconomic
values of marine birds involve both consumptive
and nonconsumptive uses. Consumptive
uses may provide socioeconomic values in the
form of meat, eggs, oil, feathers, down, and
guano. Cultural and recreational benefits may
also be involved. Nonconsumptive uses benefit
the tourist and recreation industries as
well as providing less tangible social values,
such as esthetic appreciation and environmental
education and scientific study opportunities.</p>

<p>In this paper we examine some social and
economic indicators that are believed to
demonstrate people's growing awareness and
interest in marine birds. These indicators involve
a broad spectrum of values and illustrate
the critical need for adoption of a strong
North American marine bird conservation
program.</p>


<h4>Historical Perspective</h4>

<p>Since earliest times, marine birds have accompanied
the evolution of human societies in
coastal and insular environments of the
world. Their social value is in part recorded in
kitchen middens of ancient campsites and
villages. From the time man first inhabited
the seacoasts and ventured out in ships, the
company of seabirds has added life and inspiration
to what otherwise would be a bleak
and desolate landscape. Fishermen long ago
learned to use seabirds to show them where
the rich fishing grounds were located, and the
cries of birds were often used to guide
mariners away from dangerous cliffs during
foggy weather.</p>

<p>At the time of the first contact with Europeans,
native peoples of arctic Canada and
Alaska reportedly took birds with bolas,
snares, spears, arrows, and nets; they herded
flightless waterfowl and gathered eggs as
well. Brandt (1943) said that Alaskan
Eskimos would have been destitute if eiders
(<em>Somateria</em> spp.) had not been available for
food and clothing, and Ekblaw (1928) believed
the dovekie <em>(Plautus alle)</em> saved the polar
Eskimo from extinction.</p>

<p>Marine birds have often served as an emergency
food supply for explorers, sailors, and
others: according to Tuck (1960) "The accounts
of early arctic explorers and marooned
whalers describe many instances in which
starvation was averted by eating murres"
(<em>Uria</em> spp.). One burrowing petrel of Australia
was given the title "the bird of providence"
because it saved the lives of shipwrecked
mariners and convicts when supply ships
from Sydney failed to reach them between
March and August of 1790 (Serventy 1958).</p>

<p>Marine birds have also been taken because
of the economic values of their feathers and
oil. When economic overutilization has oc<span class="pagenum"><a name="Page_175" id="Page_175">[Pg 175]</a></span>curred,
entire species were sometimes totally
destroyed. This in fact happened to the great
auk <em>(Pinguinus impennis)</em>. When Jacques Cartier
visited the Funk Islands off Newfoundland
in May 1534, he and his crew filled
several barrels with great auks and salted
them down for future consumption. So severe
was the slaughter in the next 3 centuries that
the species became extinct in its known breeding
haunts, which originally extended from
Newfoundland through Greenland and Iceland,
to the Hebrides. The last one was killed
at a stack rock off Iceland in 1884 (Lockley
1973).</p>

<p>Other species have been almost totally destroyed.
Colonization of Bermuda by Spain in
the 17th century resulted in the near annihilation
of the Bermuda petrel <em>(Pterodroma
cahow)</em> there. Ships' crews found the birds to
be fat and delicious, and they dried and salted
those that could not be eaten fresh. Today,
only about 20 breeding pairs remain, and are
under strict protection by the Bermudan government
(Lockley 1973).</p>

<p>The North Pacific albatrosses (<em>Diomedea</em>
spp.) were nearly exterminated by Japanese
feather hunters near the end of the 18th century.
The short-tailed albatross <em>(D. albatrus)</em>
was also nearly wiped out at its breeding colonies
west of the Hawaiian Islands (Bourne
1972).</p>

<p>Other species that were carelessly exploited
for their meat and plumage in the past, but
which have since regained their numbers, include
the fulmar <em>(Fulmarus glacialis)</em> on St.
Kilda Island in the North Atlantic; and the
North Atlantic, South African, and Australian
gannets (<em>Morus bassanus</em>, <em>M. capensis</em>,
and <em>M. serrator</em>) (Bourne 1972; Lockley
1973). In some instances entire breeding colonies
of a species have been destroyed while
others have survived. On the Abrothos Islands
in western Australia, for example, large
nesting colonies of sooty terns <em>(Sterna fuscata)</em>
and common noddies <em>(Anous stolidus)</em>
appear to have been wiped out on Rat Island
by indiscriminate "egging" for food, whereas
similar-sized colonies survive on other islands,
where they are now controlled by the
Fisheries and Fauna Department (Serventy
et al. 1971).</p>

<p>Historically, it has probably been man's unregulated
harvest of marine birds that has
been the primary cause of their destruction.
Generally, the loss of a species because of unregulated
harvest is no longer a matter of major
concern, because most countries of the
world are providing at least some protection
for their marine birds. However, other factors
such as habitat destruction and contamination
of the marine environment by industrial
pollutants are posing increasingly serious
threats to many.</p>


<h4>Social and Economic Indicators</h4>

<p>Economic indicators concerning consumptive
uses of wildlife, including marine birds,
are frequently misunderstood. In a dollar-oriented
and over-consumptive society like
ours, economic values are usually seen as
being in conflict with esthetic values. "Economic
use" usually conjures up images of
man's overutilization and, hence, long-term
depletion of wildlife resources. However,
when speaking of economic use, it is important
to distinguish between such overuse and
sustained-yield management.</p>

<p>Although both types of use have provided
economic benefits over the years, overharvest
that results in long-term resource depletion is
not usually the most or best economic use in
the long run; obviously a "harvest" cannot be
sustained at a given level when the resource
base is constantly being depleted. On the
other hand, when certain species of marine
birds are used in accordance with principles of
sustained yield, they can provide long-term
economic values to society in conjunction
with the social, esthetic, and intangible values
that their preservation insures. Of course, for
many species esthetic values far outweigh
economic ones derived through commercialization.</p>


<h5><em>Commercial Uses</em></h5>


<h6>Muttonbirds</h6>

<p>The muttonbird industry of Australia is an
excellent example of the commercial use of
marine birds on a sustained-yield basis. Fledgling
Tasmanian muttonbirds, or slender-billed
shearwaters <em>(Puffinus tenuirostris)</em>, are commercially
harvested each year from their colonies
on islands of Bass Strait, mainly in the<span class="pagenum"><a name="Page_176" id="Page_176">[Pg 176]</a></span>
Flinders Island group.</p>

<p>These muttonbirds are marketed as fresh or
salted "Tasmanian squab." Various by-products,
including oil, body fat, and feathers, are
also sold. In 1968, a total of just under one-half
million young birds were taken. Prices to
the producers varied from $12 to $14 (Australian
dollars) per hundred salted birds and
$16 per hundred fresh birds. Stomach oil
brought 75&cent; per gallon. Assuming the average
price per hundred birds to be $14, the
meat alone was worth about $70,000 per year
to the producers. The retail value was of
course much higher. Although the muttonbird
harvest is no longer the mainstay of the
Flinders Island economy, according to Serventy
(1969) it is still a picturesque and important
annual social event.</p>

<p>Serventy et al. (1971) believed the commercialization
of the muttonbird preserved its
numbers: "Had there been no vested interests
to preserve the 'birding islands' as such,
many of them would in the course of time
have been 'improved' as sheep stations and
the shearwater populations would have declined
and vanished."</p>


<h6>Sooty Terns</h6>

<p>The Caribbean is the home of the world's
most important wild egg producer&mdash;the sooty
tern. In some years about 2 million sooty tern
eggs from the Seychelles and 0.6 million from
Morant and Pedro bays have reached Caribbean
markets (Tuck 1960).</p>


<h6>Eiders and Murres</h6>

<p>Although the shooting of birds is not as important
economically to Greenland's approximately
50,000 residents as are sealing, whaling,
and fox hunting, the harvest of seabirds
is an ancient tradition that still means production
of an important food source that the
many Greenlanders could not exist without.
About 30 species of marine birds are harvested
for human consumption, eider ducks
and murres being by far the most important.
In west Greenland about 750,000 birds
(equivalent to about 825 tons of meat) and
10,000 eggs are harvested annually. Murres
constitute the main dish in summer at small
coastal outposts with access to rookeries.
Great quantities are also dried and salted for
use in winter. Murre canneries at Upernavik
have supplied southern cities with the frozen
meat of about 25,000 to 30,000 murres annually.
However, this commercial activity
would be prohibited by a proposed new Greenland
game law (Salomonsen 1970).</p>

<p>Banding has shown that about 22% of
Greenland's eider population, or about
150,000 birds, is shot annually. Collecting of
eider eggs is now prohibited except in the
Thule District, where 10,000 are taken annually.
Eider down is still collected from nests
for sale to a trading company for the manufacture
of much demanded eider-down coverlets
(Salomonsen 1970).</p>

<p>A growing human population, the widespread
use of modern firearms, and the increasing
use of speedboats in hunting have resulted
in serious declines in many of Greenland's
marine bird populations. The Greenland
government has demonstrated its concern
by instituting protective measures in response
to Danish expert advice. For example,
the common puffin <em>(Fratercula arctica)</em> was
given 10 years of total protection in 1961 after
bird numbers had seriously declined as a result
of over-harvesting of the birds and their
eggs (Lockley 1973). This protection was extended
in 1970. Also, it is now illegal to discharge
firearms at most marine bird rookeries
in Greenland.</p>

<p>With protection of bird habitats from human
intrusion and toxic environmental pollutants,
adequate enforcement of sound conservation
laws, greater efforts in conservation
education, and scientific regulation of harvests,
Greenland's valuable marine bird resource
could probably withstand intensive
utilization indefinitely (F. Salomonsen, personal
communication). Salomonsen has been
quick to point out, however, that people
should not be encouraged to believe that the
value of seabirds for food is the only reason
they should be saved.</p>

<p>Although several species of marine birds
serve as sources of food in the Soviet Union,
down of eider ducks and eggs of murres are
considered to be the most important to the
economy. These birds are referred to as trade
birds due to their commercial importance
(Belopol'skii 1961).</p>

<p><span class="pagenum"><a name="Page_177" id="Page_177">[Pg 177]</a></span></p>


<h6>Guano</h6>

<p>Peruvian guano beds are currently being
managed on a sustained-yield basis; the harvest,
as in the days of the Incas, depends entirely
on the amount of guano deposited each
year. Conservation and management policies
have resulted in a steady increase in the
amount extracted, from around 20,000 tons in
1900 to over 200,000 tons in 1971 (Lockley
1973).</p>

<p>The islands off south and southwest Africa
are also commercial producers of guano. The
annual yield from these breeding colonies
averaged 3,971 tons in the 12-year period,
1961-72. In 1969, guano brought 4.75 Rands
(equivalent to $7.11) per 200-pound bag.
South African gannets are apparently depositing
guano that is worth twice as much as
the fish they consume to produce it (Jarvis
1971).</p>


<h5><em>Indirect Commercial Benefits</em></h5>

<p>Marine birds also play significant roles in
the economies of the world's oceans, where
algae, invertebrates, fish, seabirds, mammals,
and man interact in complex ways. The bioenergetics
and nutrient cycling in ocean ecosystems
is even less well understood than the
contributions seabirds make to man's dollar
economies.</p>

<p>Sanger (1972) has conservatively estimated
that in the subarctic Pacific region alone,
birds consume from 0.6 to 1.2 million tons of
food and return from 0.12 million to 0.24 million
tons of feces each year.</p>

<p>Marine bird excrement is especially rich in
nitrates and phosphates, which phytoplankton,
the basis of ocean food pyramids, requires.
Marine birds then, at least to some extent,
help to sustain the northern commercial,
recreational, and subsistence fishing industries.
The fisheries in turn sustain seals and
certain other mammals which are also essential
elements of northern subsistence and recreational
economies. Thus, marine birds contribute
economic benefits indirectly as well as
directly by serving as critical links in ecosystem
food chains (Tuck 1960).</p>


<h5><em>Subsistence Uses</em></h5>

<p>The use of marine birds and their products
does not have to be commercial to be economic.
Economics is the science of the allocation
of scarce resources. Any resource, regardless
of whether it is bought or sold, has value
to people and is therefore an economic commodity.
Thus, any society has an economy
whether or not it uses cash, and when the
meat, feathers, or oil of marine birds are used,
the birds have economic value. The problem,
of course, is that of trying to determine just
what this value is when a cash medium does
not exist.</p>

<p>One of the ways to estimate this value is to
assign implicit gross dollar values to seabirds,
based on what it would cost to replace products
derived from them with store-bought
items of a similar, or substitutable, nature
(this is a gross rather than a net value because
it does not include the cost of guns, ammunition,
transportation, etc., required to harvest
and process the resource).</p>

<p>There have been many occasions in the past
when it would have been physically impossible
to find substitutes for seabird products.
In such cases, and where seabirds may well
have meant the difference between life and
death, the economic value of the resource
could be considered a plus infinity.</p>

<p>There are probably few, if any, places in the
world today where people would starve if they
could not obtain marine birds. However, there
are still many situations where available substitutes
are poor, or very expensive. And
there are others where, even though the birds
are no longer necessary for economic survival,
they are still very important in terms of sociocultural
traditions. According to Tuck (1960),
"Wherever a wild animal is important to the
economy of a people, its capture and use become
part of the tradition of that people."
Thus, while economic values can be measured
in terms of substitutable store-bought foods,
social and cultural values cannot be. To force
complete dependence on a people by flying in
foods from "Outside" is often socially intolerable
because it tends to remove pride, a sense
of worth, and therefore the reasons for living.</p>

<p>Marine birds have served as important
sources of food in the Faeroes Islands for cen<span class="pagenum"><a name="Page_178" id="Page_178">[Pg 178]</a></span>turies,
the puffin being unquestionably the
most valuable. Williamson (1945) reported
that in a good year the total puffin catch may
be between 400,000 and 500,000. In addition,
as many as 120,000 murres are snared or shot
annually by the Faeroese, and at least twice
that many eggs are taken and Tuck (1960)
stated, "The economic necessity of 'fowling'
in the Faeroes has by virtue of long centuries
of usage become part of the national life, affecting
folklore and customs, and providing
outlets for the sporting instinct inherent in
the people." A Faeroese guidebook even suggests
that its importance to the Faeroese culture
has been in no way diminished by the influence
of modern civilization. Current
Faroese game laws appear to be effective in
assuring a sustained yield of marine birds
while guaranteeing their long-term survival.</p>

<p>Seabirds and their eggs constitute a small,
but still very important, part of the total diet
of the Eskimos and Indians living along the
Arctic coast of the Northwest Territories and
Alaska. In spite of the many changes occurring
in the North, there is, even for the wage
earner, a strong psychological attachment to
the land and sea and the free life it represents.
In spring, the release from the long monotonous
winter is marked by the rites of ratting,
fishing, sealing, whaling, or marine bird hunting
and egg gathering, according to village
tradition.</p>

<p>For those living off the land in such remote
coastal outposts as Sachs Harbor on Banks
Island, Holman Island on the Mackenzie
Delta, Point Hope and Point Barrow in northern
Alaska, Inalik on Diomede Island in the
Bering Strait, or Hooper Bay on the Yukon-Kuskokwim
Delta, the spring marine bird
hunt represents a change of diet and activity.
It offers opportunity to renew age-old traditions
and continues a cultural bond among
those confined to jobs in the settlements&mdash;vacationing
and absenteeism from jobs and
schools are always highest during late May
and early June.</p>

<p>Marine birds yield between a few grams and
2 kg of meat, depending on the species.
Usually the birds are either consumed soon
after they are taken or stored in an icehouse
for use throughout the summer. Most often
the meat is cooked into a soup or stew with
rice, noodles, and onions. A few birds may be
dried or salted so that they can be used for
special holiday feasts during the winter.
Sometimes feathers are saved for the manufacture
of parkas, ceremonial fans, and
masks. In some areas of the Yukon Delta,
goose and duck down is still saved and used in
quilts that can be found in nearly every home.
In the spring 1975 issue of the catalog of a
Seattle, Washington, outfitter, down quilts
for single beds were listed at $95. Thus, there
is a substantial cash savings by home manufacture
of such items.</p>

<p>The Yukon Delta in western Alaska is the
area where the use of marine birds is most extensive
and significant. Klein (1966) provided
harvest data by village for the entire area and
showed that, in general, geese were more important
than ducks, representing about two
thirds of the take in both the spring and the
fall. The average numbers of ducks (mostly
pintails, <em>Anas acutus</em>) and geese (primarily
white-fronted geese, <em>Anser albifrons</em>); emperor
geese, <em>Philacta canagica</em>; cackling
Canada geese, <em>Branta canadensis minima</em>;
and black brant, <em>Branta nigricans</em>, taken per
household were 77 by the Yukon River
villages, 69 by the Kuskokwim River and tundra
villages, and 94 by the Bering Sea coastal
villages. Although eggs gathered by Yukon
River villagers averaged less than a dozen per
household, Kuskokwim people took about 3
dozen and coastal people about 6.5 dozen on
the average. Eggs of black brant and cackling
Canada geese were especially favored, but
even those of small passerines were acceptable.
The average size of households for all
areas was believed to be between 5.5 and 6.5
persons.</p>

<p>A 1968 survey of waterfowl taken in the
Mackenzie Delta region, made by the Canadian
Wildlife Service, showed an average take
per household of about 70 birds, a figure comparable
to that for the Yukon Delta. In the
Mackenzie region, however, ducks were more
important than geese, representing about
60% of the harvest.</p>

<p>More recent data on Alaska waterfowl harvest
per household is available for other
coastal regions. Data provided by two regional
native corporations for the Joint Federal-State
Land Use Planning Commission
for Alaska in 1973 showed an average per-household
waterfowl harvest of 33 ducks and<span class="pagenum"><a name="Page_179" id="Page_179">[Pg 179]</a></span>
geese for Kotzebue area villages, 68 for
Norton Sound villages, 24 for northwest
Seward Peninsula villages, and 37 for St.
Lawrence, Diomede, and King Island villages.</p>

<p>A 1974 subsistence survey carried out
jointly by the University of Alaska and the
Bristol Bay Native Corporation showed that,
in 20 Bristol Bay villages, 57% of the households
harvested waterfowl. The average kill
was 32 birds per household.</p>

<p>Eider ducks are the most important marine
birds taken by residents of Barrow, Alaska.
Johnson (1971) interviewed 31 adult hunters
with average kills of 88 birds per hunter. Barrow
people also take substantial numbers of
geese at Atkasook, a summer camp on the
Meade River 80 miles southeast of Barrow.</p>

<p>Point Hope, Alaska, villagers also favor
eider ducks above all others. Pederson (1971)
indicated that each household that hunted
took about 150 eiders in the summer of 1971.
Each summer, Point Hope and Kivalina residents
travel to the Cape Thompson and Cape
Lisburne cliffs to gather murre eggs. Both
Pederson (1971) and Kessel and Saario (1966)
showed an average harvest of 5 to 10 dozen
eggs per household (equivalent in weight to 10
to 20 dozen chicken eggs).</p>

<p>To our knowledge, there is no available evidence
to indicate that the number of migratory
birds taken in the North in spring and
fall is a significant factor in the survival of a
particular species. The birds are, however, a
significant factor in the economy and culture
of the people of the Mackenzie Delta region
and much of coastal Alaska. This may not always
be true, for their social and economic
conditions are changing rapidly.</p>

<p>With the native birthrate twice the national
average and with hunting technology improving
yearly, the day will undoubtedly come
when marine birds and other wildlife resources
are not able to withstand intensified
harvest pressures without more regulation
and control. An obvious need exists for government
conservation agencies to work more
closely with the native people of northern regions
in conservation education and development
of sound harvest regulations.</p>


<h5><em>Recreational Uses</em></h5>

<p>No attempt was made in this evaluation to
affix dollar values to every marine bird enjoyed
by recreationists. Goldstein (1971), in
his economic study of wetlands, found it impossible
to fix the value of the production and
harvest of migratory waterfowl in Minnesota.</p>

<p>The amount of money spent by recreationists
in seeking enjoyment from marine
birds does not measure the values they derive;
it measures only their costs to participate in
such ventures. The analogy that could be
made is that the value of a diamond is equal to
the cost of mining it. Nevertheless, expenditure
data for services and goods provided by
air-taxi and charter boat operators and merchants
selling bird guides, binoculars, and
other outdoor recreational equipment are useful
indicators in establishing the secondary or
indirect benefits of recreational activities associated
with marine birds.</p>

<p>The normal economic concept of net benefits
from marine bird recreation would include
only those accruing to individuals who provide
goods and services to the recreationists,
gross revenues minus the costs (Wollman
1962; Pearse and Bowden 1969). This economic
return, however, in no way measures direct
benefits of marine bird resources to the
recreationists.</p>

<p>Another important consideration in evaluating
recreational use of marine birds is to
recognize that many of the nonparticipants
either value the option of being able to take
advantage of them in the future, or simply believe
that the availability of such resources
benefits society (Stegner 1968). Such benefits
are difficult, if not impossible, to quantify yet
may be exceedingly important due to the
uniqueness of the marine bird resource and because
many decisions affecting it may prove
irreversible.</p>

<p>Increasing numbers of bird enthusiasts
throughout North America are discovering
the excitement and pleasures derived from
visiting marine bird rookeries. As pointed out
by Sowl and Bartonek (1974), and as anyone
can attest who has ever had the privilege of
watching the antics of tufted puffins <em>(Lunda
cirrhata)</em> near their colonies on a day when the
sun is obscured and the air buoyant, watching
seabirds is fun.</p>

<p>We have found that organizations and businesses
in practically every North American
coastal State and Province, from Nova Scotia<span class="pagenum"><a name="Page_180" id="Page_180">[Pg 180]</a></span>
to Florida and Alaska to California, are busy
scheduling boat or airplane excursions to
marine-bird viewing areas off their shores.
The Alaska and Washington State ferry systems
have for years been providing passengers
opportunity to enjoy seabirds of the
North Pacific coast. Audubon chapters in San
Diego, Los Angeles, Monterey, Seattle, Anchorage,
and other cities sponsor annual excursions
to seabird colonies.</p>

<p>In 1975 a charter airline service in Anchorage,
Alaska, booked 530 people in 51 tours to
fly to the Pribilof Islands in the Bering Sea to
view the outstanding seabird and fur seal
colonies there. Included in the bookings were
three National Audubon Society International
Ecology Workshops, the Massachusetts
Audubon Society, the National Wildlife
Federation, and Canadian Nature Federation.
Participants paid from $1,500 to $2,000 for
these tour packages to Alaska. At $300 to
$380 per person, depending on the length of
the excursion, the air charter service grossed
about $160,000 from these tours (Reeve Aleutian
Airways, personal communication).</p>

<p>Fairweather Outings, a small cruise business
based in Sitka, Alaska, takes people on
wilderness excursions in the west Chichagof-Glacier
Bay area of the southeastern part of
the State. The seabird rookeries are one of the
principal attractions for the 90 people taking
these trips each year. Over one-third of the
clientele has been from outside Alaska; thus
their dollars are new dollars to the State's
economy. Fairweather Outings grossed about
$11,000 in 1974 (Charles Johnstone, personal
communication).</p>

<p>These examples illustrate how seabirds,
both directly and indirectly, help small
coastal businessmen earn a living. It is also
important to recognize that the multiplier effects
generated by the expenditures in all of
the above examples ripple through the regional
and State economies.</p>

<p>Despite the great social and economic significance
of such activities along our coasts,
apparently no attempt is being made to determine
the number of people involved in such
pursuits and how much they are spending. A
study of the phenomenon would undoubtedly
produce startling results.</p>

<p>The Wildlife Management Institute (1975)
revealed that the national estimated value of
manufacturers' shipments in 1972 was $157
million for camping equipment, $5 million for
binoculars, and $19.9 million for bird feed.
Sales of wild bird feed have been increasing 5
to 10% per year recently. These are all economic
indicators of recreation trends of which
enjoyment of marine birds is a part.</p>

<p>A major use of photographic equipment and
related products and services is in the natural
and scenic areas of the nation. Manufacturers'
shipments of photographic equipment, and
photofinishing, were valued at $2.3 billion in
1972. A 5% excise tax on these items would
have generated nearly $118 million (Wildlife
Management Institute 1975).</p>

<p>Since inadequate funding plagues most nongame
management initiatives, the Wildlife
Management Institute (1975) recommended
that Congress authorize a matching grant-in-aid
program to benefit nongame fish and wildlife.
Funds would be obtained from new manufacturers'
excise taxes on designated outdoor
recreational equipment to initially yield at
least $40 million annually.</p>

<p>The Executive Committee of the International
Association of Game, Fish and Conservation
Commissioners and the Council of the
Wildlife Society have already endorsed model
legislation for a State program for nongame
wildlife conservation (Madson and Kozicky
1972). We urge that these proposals be given
serious consideration in terms of future funding
of marine bird conservation programs in
North America.</p>

<p>It is encouraging to note that several
States, including Washington, Oregon, and
California, have recently initiated nongame
wildlife programs that have resulted in substantial
benefits to their citizens. The California
legislature, for example, enacted a law
in 1974 to provide a means for individuals and
organizations to donate funds for supporting
nongame species management. The California
Department of Fish and Game has increased
its nongame staff and appointed a citizen
Nongame Advisory Committee to help develop
and implement nongame programs.</p>

<p>Because most species of marine birds are
not hunted by sportsmen in North America,
this increased emphasis on nongame species
may eventually benefit research and management
programs for seabirds substantially.</p>

<p><span class="pagenum"><a name="Page_181" id="Page_181">[Pg 181]</a></span></p>


<h5><em>Scientific Research</em></h5>

<p>Even now, marine-bird research studies and
inventories require the expenditure of several
million dollars annually along our coasts. In
Alaska a multimillion dollar Federal effort
has been initiated to assess the environmental
risks of developing offshore petroleum potential
in the Gulf of Alaska and five other key
areas of the State. These areas represent 60%
of the nation's total continental shelf and support
some of the largest marine-bird populations
in the world. The program to examine
life-forms and the physical environment of the
petroleum lease areas will require 4 to 5 years
to complete. Approximately $1.5 million has
been allocated to conduct an environmental
assessment of marine bird resources in the
first 18 months alone.</p>

<p>The U.S. Fish and Wildlife Service is spending
about $40,000 to determine the seasonal
occurrence, density, and distribution of
marine birds in coastal waters adjacent to
new national wildlife refuges in Alaska being
proposed pursuant to the Alaska Native
Claims Settlement Act of 1971, and almost
$200,000 to study and manage migratory
birds&mdash;including marine birds&mdash;on existing
refuges.</p>

<p>Although generated by external events (including
requirements pursuant to the National
Environmental Policy Act of 1969)
rather than by the resources themselves,
these expenditures at least indirectly reflect a
social concern for the welfare of marine birds.</p>


<h5><em>Citizen Involvement
(Social Indicator)</em></h5>

<p>Another encouraging aspect of seabird conservation
and its meaning to society is the increasing
involvement of citizens in the issue.
Although agencies have not been as responsive
as many would like, administration of
government at all levels has been shaken and
stimulated by citizen participation. As
Russell W. Peterson, Chairman of the Council
on Environmental Quality, has stated, "Citizen
action is the essence of democracy. Citizen
movements should be encouraged and expanded.
The involvement of people is necessary
to counterbalance the disproportionate
influence of the professional lobbyists and
public relations operators hired to further the
special interests of their clients." Mr. Peterson
further emphasized that government
thrives much better on citizen concern and attention
than on indifference and neglect.</p>

<p>Therefore, it is highly significant that the
Pacific Seabird Group has many nonprofessional,
as well as professional, members and
that the 1975 International Symposium on
Conservation of Marine Birds of Northern
North America had strong citizen involvement
and participation. As everyone recognizes,
nothing works in government unless
people, be they doctors, lawyers, college professors,
students, environmentalists, or Indian
chiefs, make it work.</p>

<p>Educators must upgrade training in environmental
sciences so that an environmental
awareness (conservation ethic) is instilled in
young people. In this regard, an Alaskan bird
study program proposed for Alaska schools
by J. G. King, Jr., of the U.S. Fish and Wildlife
Service in 1962 deserves close scrutiny.
This highly innovative and practical environmental
education proposal apparently arrived
before its time, for nothing was ever done to
institute it. Possibly, now would be a good
time to give it a closer look.</p>


<h4>Conclusions</h4>

<p>Success in more adequately recognizing and
using social and economic indicators to
strengthen and broaden seabird programs will
depend on the ability of the resource management
agencies to blend the old with the new.
It is obvious to most that new alignments,
programs, authorities, and sources of funds
are needed, but by themselves, they will not
be enough to overcome the continuing massive
losses of wildlife habitat due to population
growth and technological impacts resulting
from various developmental programs.</p>

<p>No marine bird programs will be successful
without a strong political base. If this is to be
assured, resource agencies must be more responsive
to the needs of both consumptive
and nonconsumptive users and involve them
in their programs from early in the planning
process. Because marine birds and the natural
environments they inhabit are jointly valued<span class="pagenum"><a name="Page_182" id="Page_182">[Pg 182]</a></span>
over time and are jointly owned, it is important
to ask not only what is efficient from the
point of view of the present generation but
also what is equitable across generations.</p>


<h4>References</h4>

<div class="hangindent">

<p>Belopol'skii, L. O. 1961. Ecology of sea colony birds
of the Barents Sea. (Transl. from Russian.) Israel
Program for Scientific Translations, Jerusalem.
346 pp.</p>

<p>Bourne, W. R. P. 1972. Threats to seabirds. Int.
Counc. Bird Preserv. Bull. II:200-218.</p>

<p>Brandt, H. 1943. Alaskan bird trails: adventures of
an expedition by dogsled to the delta of the
Yukon River at Hooper Bay. Bird Research
Foundation, Cleveland, Ohio. 464 pp.</p>

<p>Ekblaw, W. E. 1928. The material response of the
polar Eskimo to their far arctic environment.
Ann. Assoc. Am. Geog. 17(4):147-195.</p>

<p>Goldstein, J. H. 1971. Competition for wetlands in
the Midwest: an economic analysis. Resources for
the Future, Inc., Washington, D.C. 105 pp.</p>

<p>Jarvis, M. J. F. 1971. Interactions between man
and the South African gannet. Biol. Conserv.
3(4):269-273.</p>

<p>Johnson, L. L. 1971. The migration, harvest and importance
of waterfowl at Barrow, Alaska. M.S.
Thesis. Univ. of Alaska, Fairbanks. 87 pp.</p>

<p>Kessel, B., and D. Saario. 1966. Human ecological
investigations at Kivalena. Pages 969-1039 <em>in</em>
N. J. Wilimovsky and J. N. Wolfe, eds. Environment
of the Cape Thompson region, Alaska.
U.S.A.E.C., Div. Tech. Inf., Oak Ridge, Tennessee.</p>

<p>Klein, D. R. 1966. Waterfowl in the economy of the
Eskimos on the Yukon-Kuskokwim Delta,
Alaska. Arctic 19(4):319-335.</p>

<p>Lockley, R. M. 1973. Man and seabirds. Pages 74-90
<em>in</em> R. M. Lockley. Ocean wanders: the migratory
seabirds of the world. Stackpole Books, Harrisburg,
Pennsylvania.</p>

<p>Madson, J., and E. Kozicky. 1972. A law for wildlife:
model legislation for a State nongame wildlife
conservation program. Winchester-Western
Division, Conservation Department, East Alton,
Illinois. 20 pp.</p>

<p>Pearse, P. H., and G. K. Bowden. 1969. Economic
evaluation of recreational resources: problems
and prospects. Trans. N. Am. Wildl. Nat. Resour.
Conf. 34:283-293.</p>

<p>Pederson, S. 1971. Status and trends of subsistence
resource use at Point Hope. Pages 37-89 <em>in</em> B.
MacLean, ed. Point Hope project report. Univ. of
Alaska, Fairbanks.</p>

<p>Salomonsen, F. 1970. Birds useful to man in Greenland.
Pages 169-175 <em>in</em> Productivity and conservation
in northern circumpolar lands. International
Union for Conservation of Nature and
Natural Resources, Morges, Switzerland.</p>

<p>Sanger, G. A. 1972. Preliminary standing stock and
biomass estimates of seabirds on the subarctic
Pacific region. Pages 589-611 <em>in</em> A. Y. Yakenouti
et al., eds. Biological oceanography of the North
Pacific. Idemitsy Shoten, Tokyo.</p>

<p>Serventy, D. L. 1958. Mutton-birding. Pages 233-234
<em>in</em> A. H. Chisholm, ed. The Australian Encyclopedia,
Vol. 6. Angus and Robertson, Sydney.</p>

<p>Serventy, D. L. 1969. Mutton-birding. Pages 53-60
<em>in</em> K. Taylor, ed. Bass Strait Australia's last frontier.
Australian Broadcasting Company, Sydney.</p>

<p>Serventy, D. L., U. Serventy, and J. Warham. 1971.
Seabird conservation problems in Australia.
Pages 40-44 <em>in</em> D. L. Serventy, U. Serventy, and
J. Warham. The handbook of Australian birds.
A. H. and A. W. Reed, Sydney.</p>

<p>Sowl, L. W., and J. C. Bartonek. 1974. Seabirds:
Alaska's most neglected resource. Trans. N. Am.
Wildl. Nat. Resour. Conf. 39:117-126.</p>

<p>Stegner, W. 1968. The meaning of wilderness in
American civilization. Pages 192-197 <em>in</em> R. Nash,
ed. The American environment: readings in the
history of conservation. Addison-Wesley Publishing
Co., Reading, Massachusetts.</p>

<p>Tuck, L. M. 1960. The murres. Ottawa, Can. Wildl.
Ser. 1. 260 pp.</p>

<p>Wildlife Management Institute. 1975. Current investments,
projected needs, and potential new
sources of income for nongame fish and wildlife
programs in the United States. Wildlife Management
Institute, Washington, D.C. 55 pp.</p>

<p>Williamson, K. 1945. The economic importance of
seafowl in the Faeroes Islands. Ibis 87:249-269.</p>

<p>Wollman, N., chairman. 1962. The value of water in
alternative uses, with special application to water
use in the San Juan and Rio Grande basins of
New Mexico. Univ. of New Mexico Press, Albuquerque.
426 pp.</p></div>

<div class="footnotes"><h4>FOOTNOTES:</h4>

<div class="footnote">

<p><a name="Footnote_51_51" id="Footnote_51_51"></a><a href="#FNanchor_51_51"><span class="label">[51]</span></a> Present address: National Audubon Society, 2 Marine Way, Juneau, Alaska
99801.</p>

<p><span class="pagenum"><a name="Page_183" id="Page_183">[Pg 183]</a></span></p></div></div>
<div class="chapter"></div>
<hr class="chap" />




<h3><a name="Resource_Development_Along_Coasts_and_on_the_Ocean_Floor" id="Resource_Development_Along_Coasts_and_on_the_Ocean_Floor">Resource Development Along Coasts and on the Ocean Floor:
Potential Conflicts with Marine Bird Conservation</a></h3>


<p class="center">by</p>

<p class="center">Donald E. McKnight</p>

<p class="center">Alaska Department of Fish and Game<br />
Subport Building<br />
Juneau, Alaska 99801</p>

<p class="center">and</p>

<p class="center">C. Eugene Knoder</p>

<p class="center">National Audubon Society<br />
Lakewood, Colorado
</p>


<h4>Abstract</h4>

<div class="blockquot">

<p>Although development of hard mineral resources, expansion of the timber
industry, and resultant increases in human pressures along the North Pacific and
Arctic coasts will ultimately adversely affect northern marine bird populations,
current and proposed activities of the petroleum industry are the most immediate
threat to marine birds. The Federal Government's recently announced
plans for oil and gas leasing on the Pacific outer continental shelf eclipse the significance
of North Slope and Cook Inlet oil developments. Within a few years,
onshore storage facilities and supertankers plying these waters will undoubtedly
result in widespread chronic and localized catastrophic contamination of northern
marine ecosystems.</p>

<p>Coastal and offshore waters south of the reaches of the seasonal ice pack are
tremendously productive, supporting a diverse wealth of bird life throughout the
year. Because these ecosystems are relatively stable and the impact of temporal
oscillations on the physical environment is not as great as in the Arctic, birds in
these areas are probably least susceptible to man's influence on a long-term
basis.</p>

<p>Avifaunal associations of the Arctic are less diverse and have shorter food
chains than more southerly ones; consequently they are more susceptible to environmental
perturbations. Slow growth and maturation rates of arctic species and
resultant prolonged population recovery periods further aggravate this
situation.</p>

<p>Available knowledge of northern seabirds and their environmental requirements
is in inverse relation to the latitude at which they are found and to the ecological
stability of the ecosystems involved. Arctic bird associations and their
fragile environments are least understood, but are doubtless the most vulnerable
to the detrimental effects of man-caused environmental degradation. The
paucity of knowledge about them limits the possibility of predicting the consequences
of petrochemical exploitation and thereby safeguarding against potential
problems. Existing technology and support system capabilities of the oil
industry are more poorly defined for arctic areas, further compounding this problem.
Regardless of information amassed in the future and precautionary
measures taken during exploitation of arctic petroleum reserves, the potential
for disastrous and perhaps irrecoverable losses to northern marine bird species
and populations is great. Losses of major magnitude could appreciably alter the
productivity of northern marine ecosystems.</p></div>

<p><span class="pagenum"><a name="Page_184" id="Page_184">[Pg 184]</a></span></p>

<p>Although the coastal waters of the northwestern
United States and western Canada
support a plenitude of marine life, including
marine birds, relatively little is known about
these ecosystems. Sustained interest in quantitative
aspects of this area's marine bird
populations has developed only within the
past few years. As Sowl and Bartonek (1974)
indicated, seabirds are the most visible component
of a marine ecosystem and, at the
same time, they are the least understood.
Management information has been haphazardly
gathered, and because seabirds occur
in incredibly large numbers in north
Pacific and arctic waters, it has been convenient
to assume that, in the absence of
problems, systematized data gathering and
analysis were unnecessary.</p>

<p>The sudden emergence in the late 1960's of
Alaska and portions of northwest Canada as
potential major oil production areas has
changed this situation dramatically. Ongoing
and planned petroleum development in the
North and the concurrent expansion of hard
mineral extraction and logging activities now
threaten to adversely affect these marine bird
resources. Alaska's human population, which
numbered only slightly over 400,000 in 1975,
will probably double within the present
decade. Doubtless, increased numbers of
people, oriented toward mineral and other resource
exploitation rather than toward traditional
wildland values, will compound these
problems. Pressures on State and local governments
for increased services necessitated
by increasing populations will require additional
expenditures. In Alaska, at least, these
demands are being imposed before revenues
from minerals become available. This necessitates
additional oil leases, timber sales, and
other means for obtaining immediate funding,
thereby adding to the acceleration and irreversibility
of industrial expansion into the
North.</p>

<p>This atmosphere of change has spawned
major government-and industry-supported
programs to broaden knowledge of northern
marine ecosystems, including their avifauna.
There has been a recent flurry of publications
on seabird populations and biology and a proliferation
of papers stressing the need to learn
more about the biota of this area. Nevertheless,
environmental impact statements on
proposed developmental programs in the
North still raise more questions than are
being answered. Attempts are being made to
apply available information on oil spills, human
disturbance, and other aspects of environmental
degradation gathered from experiences
in other areas to expected problems in
northern environments, but one must realize
that much of the information gained from experience
elsewhere is not applicable to these
areas. It is realistic to assume that, until development-related
problems occur in the
North, biologists cannot estimate the magnitude
or ecological dimensions of their effects.
However, existing knowledge of ecological
"laws" and of the biology of some species provides
the base for limited predictive efforts.</p>

<p>It is the purpose of this paper to describe
significant current and proposed resource development
along the coasts and the ocean
floors, to summarize existing knowledge of
the ecology of marine birds in these areas, and
to identify potential conflicts with marine
bird conservation. We hope that identification
of these problems will provide impetus to data
gathering and management programs necessary
for conservation of these valuable
resources.</p>


<h4>The Region and its Avifauna</h4>

<p>The region discussed here encompasses
nearly half of the United States and Canadian
coastlines, extending from Washington to the
eastern edge of the Northwest Territories.
Alaska alone has two-thirds of the United
States' continental shelf (Bartonek et al.
1971). This region's marine and estuarine
waters are some of the most productive in the
world and support a diverse wealth of bird life
throughout the year. Sanger (1972), for
example, estimated total summer standing
stocks of some 21 million birds in an area approximating
the outer continental shelf from
the Bering Strait south along the coasts of
the Aleutian Islands and North America to
central California. Sanger and King (this
volume), to whom more data were available,
revised this estimate upward to 45 million.
Bartonek et al. (1974) provided estimates of
year-round standing stocks of 27 million birds<span class="pagenum"><a name="Page_185" id="Page_185">[Pg 185]</a></span>
in the Bering Sea alone.</p>

<p>North and east of the Bering Strait, population
estimates of the bird fauna are less complete.
Swartz (1966) estimated, however, that
seabird populations of five colonies in the vicinity
of Cape Thompson in the Chukchi Sea
exceeded a total of 420,000 breeding birds in
1960. Information provided by Bartonek and
Sealy (this volume) indicates that large colony
complexes at Cape Lisburne and Little Diomede
Island each number, in aggregate, over
1 million breeding birds, mainly alcids, kittiwakes
(<em>Rissa</em> spp.), gulls (<em>Larus</em> spp.), fulmars
<em>(Fulmarus glacialis)</em>, and cormorants (<em>Phalacrocorax</em>
spp.). Although the Chukchi Sea
coast north of Cape Lisburne has no rocks
suitable for cliff-nesting seabirds, large numbers
of tundra-nesting species use the inshore
waters as a migratory pathway, and many
nonbreeding cliff nesters summer in these
waters (J. M. Scott, comments by Pacific Seabird
Group on U.S. Department of the Interior
Draft Environmental Statement 74-90).
According to Scott, sea ducks and gulls are
the most numerous birds in the Beaufort Sea.
Observations by Thompson and Person (1963)
of an estimated 1 million eiders, mostly king
eiders <em>(Somateria spectabilis)</em> and Pacific
eiders <em>(S. mollissima)</em>, passing over Point Barrow
en route to molting areas, reflect the numbers
involved. Oldsquaws <em>(Clangula hyemalis)</em>
use coastal waters of the Beaufort Sea for
postbreeding wing molts; Bartels (1973) estimated
their numbers at nearly 400,000 in the
fall and perhaps more during the molting
period. Shorebirds, jaegers (<em>Stercorarius</em>
spp.), gulls, and terns, most of which use
coastal waters at some time during the summer
season, swell bird numbers by several millions
in this area (Arctic Institute of North
America 1974).</p>

<p>As indicated by Sanger (1972), the seabirds
inhabiting coastal areas south of Bering
Strait are mainly members of the Procellariidae
in summer and Alcidae in winter. Sooty
shearwaters <em>(Puffinus griseus)</em> are the prevalent
summer species and ancient murrelets
<em>(Synthliboramphus antiquus)</em> and marbled
murrelets <em>(Brachyramphus marmoratus)</em> are
the most abundant winter species. Sanger's
central subarctic domain (offshore waters including
the Gulf of Alaska) had a different
species composition. During the summer, procellariids&mdash;mostly
slender-billed shearwaters
<em>(Puffinus tenuirostris)</em> and sooty shearwaters&mdash;made
up 94% of the biomass. Procellariids,
including fulmars, larids (largely glaucous-winged
gulls, <em>Larus glaucescens</em>), black-legged
kittiwakes <em>(Rissa tridactyla)</em>, and large
alcids, including the tufted puffin <em>(Lunda cirrhata)</em>,
made up 87% of the winter biomass in
this domain (Sanger 1972).</p>

<p>Although most of the arctic waters, including
the Bering, Chukchi, and Beaufort seas,
are unavailable to birds during the winter because
of pack ice, they seasonally host an avifauna
dominated by colony nesters, such as
common and thick-billed murres (<em>Uria aalge</em>
and <em>U. lomvia</em>), and tundra nesters, such as
oldsquaws and eiders. In far northern waters,
sea ducks (mainly eiders and oldsquaws), red
phalaropes <em>(Phalaropus fulicarius)</em>, and gulls
are the predominant species.</p>

<p>Intertidal areas throughout the Alaska,
British Columbia, and Washington coasts
support characteristic assemblages of shorebirds,
including the black oystercatcher
<em>(Haematopus bachmani)</em>, rock sandpiper
<em>(Erolia ptilocnemis)</em>, wandering tattler
<em>(Heteroscelus incanum)</em>, surfbird <em>(Aphriza virgata)</em>,
and black turnstone <em>(Arenaria melanocephala)</em>
as reported by J. M. Scott (comments
by Pacific Seabird Group to U.S. Department
of the Interior Draft Environmental
Statement 74-90). Perhaps the greatest concentrations
of shorebirds in this whole region
occur during spring and fall migrations in
Prince William Sound. The tremendous numbers
of migrating birds using these tidal and
marsh areas are hard to imagine, but densities
of up to 250,000 shorebirds per 259 hectares
(ha) on portions of the more than 51,820-ha
tidal flats of the Copper River Delta have
been recorded (Isleib and Kessel 1973).</p>

<p>Although this region's avifauna is remarkable
from the numerical standpoint, it is important
to remember also that some of its
species are limited in distribution to this area.
According to Bartonek et al. (1971), Alaska is
the only known breeding area for black turnstones,
bristle-thighed curlews <em>(Numenius
tahitiensis)</em>, surfbirds, western sandpipers
<em>(Ereunetes mauri)</em>, and Kittlitz's murrelets
<em>(Brachyramphus brevirostris)</em>. Several waterfowl
species, including the dusky Canada
goose <em>(Branta canadensis occidentalis)</em>, cack<span class="pagenum"><a name="Page_186" id="Page_186">[Pg 186]</a></span>ling
Canada goose <em>(B. c. minima)</em>, Aleutian
Canada goose <em>(B. c. leucopareia)</em>, and Aleutian
common teal <em>(Anas crecca nimia)</em> nest only in
Alaska coastal areas (Bartonek et al. 1971).
Izembek Lagoon on the Alaska Peninsula annually
hosts the entire population of black
brant, <em>Branta nigricans</em> (Hansen and Nelson
1957), and many other waterfowl, seabird, and
shorebird species nest or live in this region in
numbers important to their worldwide
welfare.</p>


<h4>Current and Planned
Resource Development</h4>

<p>The immense nonrenewable resource wealth
of Alaska and other arctic regions has remained
virtually unrecognized or unexploited
until recently because of the availability of
these resources in more accessible locations.
As supplies have diminished or been exhausted
elsewhere and demands have increased,
however, it has become economically
feasible or necessary to tap supplies in less-accessible
regions. For this reason, the petroleum
industry has recently expanded its exploratory
efforts in the far North with well-known
success. Deposits of metallic ores,
coal, and other raw materials to feed industry
have likewise been discovered and plans devised
for their extraction and sale. Pressed
with decreased availability of commercial timber
elsewhere, the logging industry has similarly
begun to broaden its efforts into Alaska.
Expansion of industrial activities into the
North is proceeding at a rapidly accelerating
pace, and these industries, their associated
support industries, and expanded human
populations are having and will continue to
have unprecedented impact on these marine
ecosystems, including their avifauna.</p>


<h5><em>Petroleum Development</em></h5>

<p>The existence of potentially marketable oil
and gas deposits in Alaska has been recognized
since the early 1900's, but it was not
until the Swanson River, Alaska, oil field was
discovered in 1957 and later developed that
the Arctic entered the modern era of oil development
(McKnight and Hiliker 1970). This
field and offshore fields in the Upper Cook
Inlet basin have been producing oil for nearly
a decade. The discovery of petroleum reserves
on Alaska's North Slope and Canada's Mackenzie
River Delta is common knowledge, and
a pipeline has been constructed to transport
Alaska oil to a tanker facility at Valdez in
Prince William Sound. Alternative proposals
to pipe North Slope natural gas along the
existing corridor to a facility in Prince William
Sound or to build a new pipeline to take
this gas to existing fields, and a planned pipeline
on the Mackenzie River Delta and south
through Canada, are being considered. Construction
of a gas liquefaction facility in
Prince William Sound and tanker traffic
through the Sound and the Gulf of Alaska are
potential ramifications of an Alaska gas pipeline.</p>

<p>As McKnight and Hiliker (1970) and Bartonek
et al. (1971) pointed out, the greatest
potential problem for marine bird populations
from North Slope oil will be associated with
the operations of the Alyeska Pipeline system's
terminal at Valdez. Oil storage and
ship-loading facilities at this port and heavy
tanker traffic through Prince William Sound
represent a pollution source that could result
in significant seabird and waterfowl mortalities.
Certainly, development of gas liquefaction
facilities in the Sound, with inherent increases
in human populations and tanker
traffic, would compound this potential
problem.</p>

<p>Although future impacts from existing
petrochemical developments are cause for
concern, the Federal Government's recently
announced plans for oil and gas leasing on the
Pacific outer continental shelf (Fig. I) eclipse
the significance of North Slope and Cook Inlet
oil developments. It now appears the Gulf of
Alaska is the most favorable area of the outer
continental shelf for oil and gas production
(Council on Environmental Quality 1974).
This area, covering more than 10.3 million ha,
has already been subjected to extensive seismic
investigations, and estimates of its undiscovered,
economically recoverable crude oil
and natural gas resources range from 3 to 25
billion barrels and 15 to 30 trillion cubic feet,
respectively (Council on Environmental
Quality 1974).</p>

<p><span class="pagenum"><a name="Page_187" id="Page_187">[Pg 187]</a></span></p>

<div class="figcenter" style="width: 700px;">
<img src="images/p_187.jpg" width="700" height="428" alt="" />
<div class="caption"><p>Fig. 1. North Pacific, showing portions of the outer continental shelf being considered for gas and oil leasing
by the Federal Government (vertical hatching) and areas leased or proposed for leasing by the State
of Alaska (cross hatching).</p></div>
</div>

<p>Kinney et al. (1970) reported that in Cook
Inlet, Alaska, an estimated 0.3% of the oil
produced and handled in offshore platform
wells is spilled. Several routine offshore operations
result in discharges of oil and other
materials into water, and, unlike accidental
spills, the probability of their occurrence is
100% (Council on Environmental Quality
1974). During drilling operations, cleaned
drilling mud and drill cuttings are discharged
overboard. Drilling mud may consist of such
substances as bentonite clay, caustic soda, organic
polymer, proprietary defoamer, and
ferrochrome lignosulfate. Waters from geological
formations are often produced and discharged
into the sea while the wells are in production.
These waters may be fresh or saline,
and often contain small amounts of oil. All of
these pollutants increase the adverse effects
of offshore oil production, and when potential
spills are also considered, the ultimate impact
on the marine ecosystem may be substantial.</p>

<p>The State of Alaska has already leased offshore
sites in Kachemak Bay, and present
considerations for future leases in the lower
Cook Inlet and Beaufort Sea further reflect
the widespread and massive nature of petrochemical
developments in the Arctic planned
for the next 2 decades (Fig. 1). Proved crude
oil reserves are less than 1 billion barrels and
natural gas reserves are less than 2 trillion
cubic feet in Cook Inlet, but it appears that
undiscovered recoverable oil and gas resources
may be much greater (Council on Environmental
Quality 1974). There are also indications
that known onshore oil reserves
along Alaska's northwest coast will soon be
opened for development by the Arctic Slope
Regional Corporation, landowners in the area
as a result of the Native Land Claims Act of
1971. This group is at least considering the
transportation of these petroleum products to
market in tankers, from an open-water port in
the Chukchi Sea&mdash;thereby adding to the
tanker traffic in northern waters.</p>


<h5><em>Hard Mineral Resource Development</em></h5>

<p>As indicated by Bartonek et al. (1971), there
has been renewed interest in opening up
Alaska's hard mineral resources to economic
development as new transportation routes<span class="pagenum"><a name="Page_188" id="Page_188">[Pg 188]</a></span>
and modes have been developed. Plans are
being completed to develop the Bering River
coal field, with the eventual goal of exporting
coking coal to Japan. Although mining operations
might ultimately affect freshwater environments
to the detriment of several waterfowl
species, including the trumpeter swan
<em>(Olor buccinator)</em>, the chief cause for concern
will be additional freighter traffic through
Prince William Sound. Similar plans to develop
Klukwan and Snettisham iron deposits
in southeastern Alaska for the use of Japanese
industry (Bartonek et al. 1971) may result
in the imposition of further traffic in
Alaska shipping lanes.</p>

<p>Plans are under way to strip-mine coal deposits
in the Beluga field near the west side of
Cook Inlet and transport a coal slurry via
pipeline to a thermal electric generation plant
opposite Anchorage on the Inlet. Impact on
tidal areas may be minor, but thermal pollution
of the waters is a possibility.</p>

<p>Development plans for tin and tungsten deposits
in the Lost River area of Alaska's
Seward Peninsula are under way after several
years of faltering starts and stops. These activities
and possible extraction of gold lying
offshore from Nome may ultimately have
some effect on these coastal areas. Methods
for recovering gold, regardless of the type,
would disrupt marine and estuarine environments
used by marine birds (Bartonek et al.
1971), and transportation of ores would also
increase freighter traffic in the Bering Sea.</p>


<h5><em>Timber Resource Development</em></h5>

<p>Although the timber industry has long been
established along the coast from Washington
north through southeastern Alaska, timber
harvests are rapidly expanding on U.S. Forest
Service lands in Alaska. The impact of this industry
is principally on terrestrial ecosystems,
but certainly log rafting in estuarine
areas, disposal of wastes from pulp mills, and
freighter traffic transporting wood pulp or
logs to Japan and west coast markets contribute
to the chronic degradation of marine
bird environments. Recent meager studies on
the Vancouver Canada goose <em>(Branta canadensis
fulva)</em> in southeastern Alaska have
pointed out the importance to this species of
coastal timber stands for nesting and
estuarine environments for brood rearing and
wintering. This essentially nonmigratory
goose (Hansen 1962) may be particularly vulnerable
to logging activities in these areas.
Similarly, recent evidence indicates that
marbled murrelets may nest in large conifer
trees adjacent to the coast, from northwestern
California to northern southeastern
Alaska (Harris 1971; Savile 1972). If this is
true, logging may eventually greatly restrict
the breeding of this numerically important inhabitant
of northern coastal waters.</p>


<h4>Assessment of Resource
Development and Potential
Conflicts with Marine Bird
Conservation</h4>

<p>Although extraction of hard mineral resources,
expansion of the timber industry,
and resultant increases in human pressures
along North Pacific and Arctic coasts will
ultimately affect northern marine bird populations,
current and proposed activities of the
petroleum industry pose the most immediate
threat to marine birds. Chronic degradation of
estuarine and marine coastal waters by logging
wastes, pulp mill and sewage effluents,
and bilge oils is an insidious process, the impacts
of which will be difficult, at best, to
quantify. Results of a major oil spill or even
low-level contamination of marine ecosystems
with oil will be more apparent, however. For
this reason, and the fact that the industry is
expanding rapidly into the North, most of this
discussion will be directed at the impacts of
oil development on northern marine birds.</p>

<p>Potential sources of adverse environmental
degradation affecting these birds resulting
from oil and gas exploration, development,
and production include: (1) oil discharges into
marine waters, both chronic and catastrophic,
(2) gravel excavation and dumping in coastal
areas, (3) seismic activities, (4) discharge of
drilling mud and drill cuttings into marine
waters, including toxic heavy metal constituents
of drilling mud, (5) disturbance resulting
from petrochemical activities, and (6) increased
human populations resulting in interference
with critical life processes and increased
hunting of game species. Each source
of environmental change will vary by latitu<span class="pagenum"><a name="Page_189" id="Page_189">[Pg 189]</a></span>dinal
and seasonal factors in their effects
upon the birds. We consider herein only
coastal and ocean floor developments and
their anticipated generalized impacts on
populations.</p>

<p>Although this is a discussion of "northern"
marine birds, it is important to remember
that we are considering a diverse avifauna
existing in an environmental gradient from
temperate to polar regions. In general, the
more southerly portions of this marine environment
are characterized by a greater diversity
of species, more complex food chains, and
a resultant greater stability (Dunbar 1968).
Arctic marine ecosystems, on the other hand,
are characterized by numerical dominance by
a few species, relatively simple food chains,
and an inherent instability or fragility (Dunbar
1968). According to Dunbar, arctic systems
are regulated primarily by temporal oscillations
in the physical environment, whereas
biological interactions (e.g., competition,
predation) are considered more significant in
the maintenance of temperate and tropical
ecosystems.</p>

<p>Because of their relative instability, arctic
ecosystems are more susceptible to alteration
by extreme environmental perturbation,
either natural or man-imposed (Burns and
Morrow 1973). Slow growth and maturation
rates of the avian constituents of these ecosystems
and resultant long recovery periods
(Ashmole 1971) further aggravate this
situation.</p>

<p>Regardless of their seasonal availability,
these arctic waters constitute some of the
most productive areas for seabirds in the
western hemisphere (Bartonek et al. 1974).
Upwelling, nutrient-rich waters, combined
with intense and prolonged incident radiation,
result in lush phytoplankton "blooms" that
form the foundation of relatively simple but
numerically strong plant and animal communities
(Ashmole 1971). A relatively small
number of avian species have evolved to take
advantage of this seasonally available food
supply, and the ability to migrate to lower
latitudes in winter is a characteristic of most
arctic-nesting species. Because summers are
short in arctic regions, early arrival and a synchronous
breeding schedule are necessary to
enable the young to leave the breeding
grounds before severe weather conditions prevail
(Ashmole 1971). Arrival of these birds
generally coincides closely with the earliest
availability of nesting habitat and food (Williamson
et al. 1966). Migration, molting, and
reproduction place tremendous stresses on
these birds, and as a result, arctic-nesting
species tend to reproduce less often and at
older ages than do those of more temperate regions
(Ashmole 1971).</p>

<p>In spite of these adaptations, arctic bird
species tread a thin line between extinction
and survival, and natural disasters take a
heavy toll. Bailey and Davenport (1972) reported
a massive mortality in a pelagic population
of common murres in Bristol Bay,
Alaska, during April 1970. They felt that this
disaster, resulting in the death of probably
100,000 or more birds, most likely resulted
from starvation precipitated by severe
weather. Barry (1968) reported a similar loss
to starvation of about 100,000 eiders along
the Beaufort Sea coast during the extremely
cold spring of 1964. Observers along Alaska's
Beaufort Sea reported finding eiders and oldsquaws
dead and dying from the effects of
cold weather in 1970 (Bartonek et al. 1971). It
is readily apparent that the tenuous existence
into which these birds have evolved leaves
them particularly vulnerable to the man-induced
stress of developments during the arctic
summer.</p>


<h5><em>Direct Effects of Oil Pollution</em></h5>

<p>The most obvious, and perhaps the most
disastrous consequence of petrochemical development
on northern marine bird populations
is that of a major oil spill or a well blowout
into marine waters. Although temperate
and tropical waters are apparently able to assimilate
oil spills and chronic pollution from
petroleum and its products (Nelson-Smith
1972), this has not been demonstrated to be
true for arctic waters. In fact, studies in the
Beaufort Sea have shown that the bacteria
that degrade oil do not use hydrocarbons at
the ambient temperatures of the Arctic
(Glaeser and Vance 1971). Therefore, a large
oil spill in the Arctic could persist for many
years. As demonstrated by Campbell and
Martin (1973), the diffusion and transport
mechanisms generated by the pack-ice dynamics
of the Beaufort Sea and the slow rate<span class="pagenum"><a name="Page_190" id="Page_190">[Pg 190]</a></span>
of oil biodegradation under arctic conditions
would combine to diffuse an oil spill over the
sea and eventually deposit oil on the ice surface.
This, in turn, would lower the natural albedo
over a large area and melt the ice in the
area of the spill. This pack ice supports an
under-ice community which is an important
food source for phalaropes, jaegers, gulls,
terns, and other seabirds (Watson and Divoky
1972).</p>

<p>As indicated by Nelson-Smith (1972) many
investigators have stated that a spot of oil
"no bigger than a dollar" on the breast of a
bird is enough to bring about death by exposure,
at least in the colder seas. It is easy to
see the relative vulnerability of already
stressed birds in arctic areas to a spill, and because
of the concentration of these birds in
available open-water areas, possibilities for
catastrophic mortalities are evident.</p>

<p>Such disasters already have occurred in
north Pacific waters. Dickason (1970) reported
an incident in which diesel oil reaching
the Alaska coast, probably from the sinking
of two Japanese freighters some distance offshore,
affected an estimated 90,000 murres.
J. G. King, Jr. (cited in Bartonek et al. 1971)
estimated that at least 100,000 birds, mostly
alcids and waterfowl, died in the vicinity of
Kodiak Island during winter 1970 as a result
of oil pollution (probably ballast dumped by
tankers entering Cook Inlet). It must not be
forgotten that chronic pollution in similar
areas where oil development and transport activities
are taking place probably kills more
birds every year than die after a single catastrophic
spill. Total annual losses due to oil in
the North Sea and North Atlantic, excluding
disasters, amount to 150,000 to 450,000 seabirds
(Nelson-Smith 1972).</p>

<p>That oil pollution, both chronic and catastrophic,
can dramatically affect populations
of marine birds has already been demonstrated
elsewhere. Uspenskii (1964) reported
that more than 30,000 wintering oldsquaws
perished from oil pollution near Botland Island
in the Baltic and that in later years this
species had almost disappeared from Swedish
Lapland. Jackass penguins <em>(Spheniscus
demersus)</em>, found only in South Africa, have
suffered losses from pollution caused by oil
traffic around the Cape of Good Hope
(Stander and Venter 1968). Their total population
was estimated at 100,000 in 1960, and in
two separate but not isolated incidents 1 to
2% of this number were known to have been
killed by oil. Unknown but considerable numbers
were uncounted or were lost at sea.
Colony nesters, including puffins <em>(Fratercula
arctica)</em>, razorbills <em>(Alca torda)</em>, and murres in
the southerly portions of the North Sea are
declining rapidly (Nelson-Smith 1972).
Puffins, which numbered 100,000 on Annet in
the Scilly Isles in 1907, were reduced to 100
birds by 1967; by then, colonies farther east
on the Great Britain coast were already extinct.
Pollution from the <em>Torrey Canyon</em> disaster
alone killed five-sixths of the puffins in the
main French colony on the Sept Isles in
Brittany and reduced the razorbills to a mere
50 birds, one-ninth of previous numbers
(Bourne 1970).</p>

<p>There is every reason to believe that similar
reductions in numbers could occur along the
tanker route from Valdez to Puget Sound,
with localized extirpation of colonies. Even
more disastrous, however, would be an inopportune
well blowout or other major spill in
arctic waters. Massed concentrations of birds,
already stressed by severe weather and food
shortages, would be extremely vulnerable to
this type of situation.</p>

<p>As pointed out by Nelson-Smith (1972),
peculiarities of bird behavior determine, to
some extent, the vulnerability of a species to
oil spills. Auks, murrelets, and puffins (all Alcidae),
loons (<em>Gavia</em> spp.), grebes (<em>Podiceps</em>
spp.), and diving ducks may be most susceptible
to oiling. Auks and loons, because
they float low in the water, may more readily
become completely covered by oil. Diving
species that become flightless during their
molt, such as alcids and waterfowl, or which
do not fly because of social bonds between
adults and flightless young (common murre)
and spend most of their lives on the water,
would be particularly vulnerable (J. M. Scott,
comments by Pacific Seabird Group on U.S.
Department of the Interior Draft Environmental
Statement 74-90). All divers can easily
surface into oil, and their reaction is to dive
again, which in a large spill could result in surfacing
into more oil. Phalaropes (<em>Phalaropus</em>
spp.), which flock to feed in eddies which concentrate
drift, may similarly be vulnerable to
adverse effects of oil that would also concen<span class="pagenum"><a name="Page_191" id="Page_191">[Pg 191]</a></span>trate
in these areas. On the other hand, gulls
swimming along the surface are likely to take
wing before becoming seriously contaminated.</p>

<p>Nelson-Smith (1972) reported that gannets
<em>(Morus bassana)</em>, which collected oiled sea-weed
for building nest mounds, contaminated
themselves and their eggs. Behavioral problems
associated with oil spills can be more
subtle, however, and Darling's (1938) conclusions
that the display of adjacent males contributes
to stimulation of the female during
courtship in seabirds breeding in massed colonies,
is a good example. If Darling was correct,
this behavioral characteristic could
further impede the recovery of a population of
auks, for example, from mortalities resulting
from catastrophic losses to spills.</p>

<hr class="tb" />

<p>On the basis of this information it is possible
to predict that alcids, which make up the
bulk of the birds inhabiting the coastal areas
during winter (Sanger 1972), would be very
susceptible to oil spills from future tanker
traffic in these waters. The potential exists,
therefore, for a tremendous impact (from a
single inopportune oil spill) upon these species
and upon the entire ecosystem. Sea ducks too,
because of their diving behavior, propensity
for flocking, and flightless molt period, would
be very vulnerable to oil spills. Wintering
flocks of oldsquaws and several species of
scoters along the coasts of Alaska, British
Columbia, and Washington can be expected to
dwindle as North Slope oil begins to be transported
to Puget Sound ports.</p>

<p>It is recognized now that seabirds transfer
and recycle nutrients and energy between trophic
levels and between regions of an ocean (Sowl
and Bartonek 1974). Although the significance
of this role in the marine ecosystem can
only be surmised at present, conservative estimates
by Sanger (1972) indicated that birds
consume from 0.6 to 1.2 million tons of food
and return from 0.12 to 0.24 million tons of
feces into the subarctic Pacific region annually.
G. A. Sanger's (personal communication)
revised estimates of these bird populations
indicated that his 1972 estimates should
be doubled. Regardless, it appears that the
disastrous effects of such a spill would extend
beyond the bird populations involved.</p>


<h5><em>Indirect Effects of Oil Pollution
and Petrochemical Developments</em></h5>

<p>By no means would direct losses attributable
to contamination by oil be the only threat
to marine bird populations as a result of
petrochemical expansions into these waters.
Some water birds that become contaminated
with nonlethal doses of petroleum during the
breeding season are not likely to breed (J. M.
Scott, comments by Pacific Seabird Group on
U.S. Department of the Interior Draft Environmental
Statement 74-90). Viability of embryos
is greatly reduced when the eggshell becomes
smeared with oil from the contaminated
plumage of the female (Hartung 1965).
Degradation of habitat, particularly to nesting
areas and food supplies, will certainly
occur, and its most pronounced effects will be
felt in the Arctic. Gravel removal for construction
of offshore drilling pads, causeways,
and onshore production facilities would displace
nesting birds and, combined with subsequent
discharge of drill cuttings, perhaps
have an adverse impact on bottom food organisms.
Nesting habitat loss through destruction
or the inability of birds to accept
disturbance could be substantial, particularly
along the Beaufort Sea coasts of Alaska and
Canada, where offshore barrier islands and
tundra-covered islands provide protection
from mammalian predators for nesting by
Pacific eiders, Sabine's gulls <em>(Xemia sabini)</em>,
Arctic terns <em>(Sterna paradisaea)</em>, black guillemots
<em>(Cepphus grylle)</em>, and other species (Arctic
Institute of North America 1974). Flaxman
Island near the mouth of the Canning
River is a tundra island supporting a nesting
population of whistling swans <em>(Olor columbianus)</em>,
and the only nesting colony of the
Alaska snow goose <em>(Chen caerulescens)</em> is on
Howe Island in the Sagavanirktok River
Delta (Arctic Institute of North America
1974).</p>

<hr class="tb" />

<p>Although there would probably be little actual
nesting habitat loss for cliff-nesting
species, human disturbance to colonies during
the nesting period, particularly from helicopter
and fixed-wing aircraft flybys, could
have considerable impact (Sowl and Bartonek
1974). The "living waterfall" effect of thou<span class="pagenum"><a name="Page_192" id="Page_192">[Pg 192]</a></span>sands
of seabirds pouring off a rookery is
truly spectacular, but each such occurrence
during incubation and brooding periods
causes a rain of eggs or young to fall from the
cliffs (Sowl and Bartonek 1974). Temporarily
abandoned chicks and eggs are susceptible to
predation by gulls or jaegers.</p>

<p>Even for species nesting on level ground,
aircraft overflights close to breeding colonies
may cause major losses to young and eggs.
Sladen and LeResche (1970) reported that
flights by an LH-34 helicopter (at 305 m altitude)
over an Adelie penguin <em>(Pygoscelis adeliae)</em>
colony caused some egg loss. Landing
this aircraft 183 m from the colony caused 50
to 80% of the birds to flee territories, resulting
in egg and chick loss. Disturbance caused
by visitors walking through or near nesting
areas of the South African gannet <em>(Sula
capensis)</em> on Bird Island, Lamberts Bay,
South Africa, caused desertion of nesting
sites (Jarvis and Cram 1971). Studies of disturbance
on breeding black brant, Pacific
eiders, glaucous gulls <em>(Larus hyperboreus)</em>,
and arctic terns at Nunaluk Spit and Phillips
Bay, Yukon, in July 1972 indicated that human
presence was the most critical form of
disturbance affecting incubating behavior of
these species (LGL Limited 1972<em>a</em>). Disturbance
by aircraft&mdash;especially helicopters&mdash;affected
the normal incubating behavior of all
species except Pacific eiders. Nesting success
of black brant and arctic terns was reduced by
this disturbance.</p>

<p>Disturbance can adversely affect molting
birds. The process of molting places heavy
energy demands on birds, and particularly on
waterfowl whose molt results in a flightless
period; few areas provide adequate protection
from predators necessary during this period.
Prime molting areas are scarce along the arctic
coast, yet are vital to the welfare of thousands
of sea ducks and seabirds. Studies conducted
by LGL Limited (1972<em>b</em>) indicated that
aircraft traffic over sea duck molting areas
altered normal behavior, and therefore had a
detrimental effect. Recommendations resulting
from these studies were that air traffic be
suspended over these areas during the molting
season.</p>

<p>For some arctic-nesting waterfowl, premigration
staging activity, during which fat reserves
to sustain southward migration are
stored, is a very important component of the
annual cycle (Delacour 1964). Snow geese,
breeding mainly in arctic Canada, concentrate
in large numbers on staging grounds along
the Beaufort Sea coast of eastern Alaska and
the Yukon. Because gas compressor stations
would be required along the proposed arctic
gas pipeline route, experimental studies were
conducted in September 1972 to determine
the effect of disturbance from sounds generated
by compressors (LGL Limited 1972<em>c</em>).
These studies indicated that compressor noise
was disruptive to staging geese.</p>

<p>Indirect effects on marine bird resources resulting
from development activities may ultimately
prove to be more detrimental than the
aforementioned direct factors. It is conceivable
that the impact of these industries,
mainly on the benthic and demersal fauna of
the coastal areas, could greatly lower the
carrying capacity of this habitat for marine
birds (Bartonek et al. 1974). Because of the
simplified and short arctic food chains and the
lack of alternative food sources in these areas,
arctic ecosystems would be particularly vulnerable
to this type of problem (Burns and
Morrow 1973).</p>

<p>Ecological or toxic influences on several
food species could result in substantial declines
in bird populations. In the Arctic,
where temperatures are low, and bacterial and
other decompositional activities are consequently
slow, spilled oil would persist for
many years, with concomitant deleterious effects
on the marine organisms of the area
(Burns and Morrow 1973). Reduced recruitment
of young would no longer balance inevitable
or density-independent population mortality
(Ashmole 1971). Although indications
are that arctic species are the most vulnerable
to this type of impact, the lack of knowledge
of the feeding niches of most seabirds discourages
further evaluation of this potential problem.
It is obvious, however, that ecology of
arctic birds is least understood, and these
species are the most vulnerable to the detrimental
effects of man-caused environmental
degradation.</p>


<h4>Conclusions</h4>

<p>Predictability of the impact of resource development
on marine birds in northern waters<span class="pagenum"><a name="Page_193" id="Page_193">[Pg 193]</a></span>
is limited by our relative ignorance of these
birds and their ecology. Just as there exists a
latitudinal gradient in the ecological stability
of the ecosystems involved, available knowledge
of these ecosystems is in inverse relationship
to the latitude at which they occur.
Arctic bird associations and their fragile environments
are least understood but are doubtless
the most vulnerable to the detrimental effects
of man-caused environmental degradation.
Existing technology and support system
capabilities of the oil industry are poorly defined
for Arctic areas, further compounding
this problem (Arctic Institute of North
America 1974).</p>

<p>Although activities associated with the extraction
of hard minerals and the timber industry
will ultimately affect northern seabirds,
petrochemical developments pose the most
immediate threat to this resource. Exploration
and development of many coastal and offshore
sedimentary basins with a potential for
oil or gas production are proceeding rapidly.
Within a few years, oil storage and loading
facilities at Valdez, Alaska, and supertankers
plying northern waters will probably result in
widespread chronic and localized catastrophic
contamination of northern marine environments.
Experience in other areas has demonstrated
that oil spills are a considerable potential
threat to these bird populations, directly
through widespread mortality and indirectly
through effects on the environment. This
threat is of such magnitude that entire populations
or species could be lost to a single spill
if it occurred at the wrong place at the wrong
time of year. Because many of these species
require 3 to 4 years for maturation and may
rear only one or two young per year, recovery
time for their populations is great (Ashmole
1971). For these and other reasons, the Council
on Environmental Quality (1974) concluded
that the Gulf of Alaska appeared more
vulnerable to major environmental damage
from outer continental shelf oil and gas development
than sites off the Atlantic coast.</p>

<p>As Bartonek et al. (1971) pointed out, it
would be a national tragedy if the great nongame
bird populations along Alaska's coast
were decimated during the "Environmental
Decade" without even being properly described.
Regardless of information amassed in
the future and precautionary measures taken
during exploitation of arctic petroleum reserves,
the potential for disastrous and perhaps
irrecoverable losses to northern marine
bird species and populations is great. Losses
of major magnitude could appreciably alter
the productivity of northern marine ecosystems,
to the detriment of other renewable
resources.</p>

<p>Knowledge of northern marine birds, their
environments, and their ecology must be
greatly expanded if the consequences of petrochemical
exploitation are to be predicted and
safeguards established against potential
problems. To the extent possible, oil exploration
and development activities should be
limited to temperate, more stable, marine ecosystems,
at least until more northerly areas
are better understood. Similarly, these activities
must be conducted in such places and at
such times that impact on the environment
will be minimized. State and federal governments
and the petroleum industry are ultimately
answerable for this responsibility.</p>

<p>The Nation must be aware of the potential
costs of energy independence set forth as a
goal of proposed oil and gas leasing of
Alaska's outer continental shelf. We must ask
ourselves if we are willing to risk extermination
of species to reach this goal, or if we can
afford the luxury of reducing the biological
productivity of these waters.</p>


<h4>References</h4>

<div class="hangindent">

<p>Arctic Institute of North America. 1974. The Alaskan
arctic coast&mdash;a background study of available
knowledge. 551 pp.</p>

<p>Ashmole, N. P. 1971. Seabird ecology and the
marine environment. Pages 223-286 <em>in</em> D. S.
Farner and J. R. King, eds. Avian biology. Vol. I.
Academic Press, New York.</p>

<p>Bailey, E. P., and G. H. Davenport. 1972. Die-off of
common murres on the Alaska Peninsula and
Unimak Island. Condor 74(2):215-219.</p>

<p>Barry, T. W. 1968. Observations on natural mortality
and native use of eider ducks along the
Beaufort Sea coast. Can. Field-Nat. 82(2):140-144.</p>

<p>Bartels, F. B. 1973. Bird survey techniques on
Alaska's north coast. M.S. Thesis. Iowa State
Univ., Ames. 47 pp.</p>

<p>Bartonek, J. C., J. G. King, and H. K. Nelson. 1971.
Problems confronting migratory birds in Alaska.
Trans. N. Am. Wildl. Nat. Resour. Conf. 36:345-361.</p>

<p><span class="pagenum"><a name="Page_194" id="Page_194">[Pg 194]</a></span></p>

<p>Bartonek, J. C., R. Elsner, and F. H. Fay. 1974.
Mammals and birds. Pages 23-28 <em>in</em> E. J. Kelley
and D. W. Hood, eds. Probes: a prospectus on
processes and resources of the Bering Sea shelf,
1975-1985. Univ. Alaska, Inst. Mar. Sci. Publ.
Inf. Bull. 74-1.</p>

<p>Bourne, W. R. P. 1970. Special review after the
"Torrey Canyon" disaster. Ibis 112:120-125.</p>

<p>Burns, J. J., and J. E. Morrow. 1973. The Alaskan
Arctic marine mammals and fisheries. Arctic oil
and gas problems and possibilities, 5th International
Congress. 24 pp.</p>

<p>Campbell, W. J., and S. Martin. 1973. Oil and ice in
the Arctic Ocean: possible large-scale interactions.
Science 181:56-58.</p>

<p>Council on Environmental Quality. 1974. OCS oil
and gas&mdash;an environmental assessment. Vol. I.
Washington, D.C. 214 pp.</p>

<p>Darling, F. F. 1938. Bird flocks and the breeding
cycle; a contribution to the study of avian sociability.
Cambridge Univ. Press, N.Y.</p>

<p>Delacour, J. 1964. The waterfowl of the world. Vol.
IV. Country Life Ltd., London. 364 pp.</p>

<p>Dickason, O. E. 1970. Center for short-lived phenomena
event report. Smithsonian Institution,
Cambridge, Massachusetts, 36(1970):926.</p>

<p>Dunbar, M. J. 1968. Ecological development in
polar regions. A study in evolution. Prentice-Hall,
Inc., Englewood Cliffs, N.J. 119 pp.</p>

<p>Glaeser, J. L., and G. P. Vance. 1971. A study of the
behavior of oil spills in the Arctic. U.S. Coast
Guard, Office of Research and Development,
Washington, D.C. 60 pp.</p>

<p>Hansen, H. A. 1962. Canada geese of coastal
Alaska. Trans. N. Am. Wildl. Nat. Resour. Conf.
27:301-320.</p>

<p>Hansen, H. A., and U. C. Nelson. 1957. Brant of the
Bering Sea&mdash;migration and mortality. Trans. N.
Am. Wildl. Nat. Resour. Conf. 22:237-256.</p>

<p>Harris, R. D. 1971. Further evidence of tree nesting
in the marbled murrelet. Can. Field-Nat. 85(1):67-68.</p>

<p>Hartung, R. 1965. Some effects of oiling on reproduction
of ducks. J. Wildl. Manage. 29(4):872-874.</p>

<p>Isleib, M. E., and B. Kessel. 1973. Birds of the north
gulf coast&mdash;Prince William Sound region, Alaska.
Biol. Pap. Univ. Alaska 14. 149 pp.</p>

<p>Jarvis, M. J. F., and D. L. Cram. 1971. Bird Island,
Lamberts Bay, South Africa: an attempt at conservation.
Biol. Conserv. 3(4):269-272.</p>

<p>Kinney, P. J., D. K. Button, D. M. Schell, B. R.
Robertson, and J. Groves. 1970. Quantitative assessment
of oil pollution problems in Alaska's
Cook Inlet. Univ. Alaska Inst. Mar. Sci. Rep.
R-69-16. 43 pp.</p>

<p>LGL Limited. 1972<em>a</em>. Disturbance studies of breeding
black brant, common eiders, glaucous gulls
and Arctic terns at Nunaluk Spit and Phillips
Bay, Yukon Territory. Report prepared for
Northern Engineering Service, Ltd. 50 pp.</p>

<p>LGL Limited. 1972<em>b</em>. Aircraft disturbance to molting
sea ducks, Herschel Island, Yukon Territory.
Report prepared for Northern Engineering Service,
Ltd. 31 pp.</p>

<p>LGL Limited. 1972<em>c</em>. Gas compressor noise simulator
disturbance to snow geese, Kamakuk
Beach, Yukon Territory. Report prepared for
Northern Engineering Service, Ltd.</p>

<p>McKnight, D. E., and B. L. Hiliker. 1970. The impact
of oil development on waterfowl populations
in Alaska. Proc. Western Assoc. State Game Fish
Comm. 50:286-297.</p>

<p>Nelson-Smith, A. 1972. Oil pollution and marine
ecology. Paul Elek, Ltd., London. 260 pp.</p>

<p>Sanger, G. A. 1972. Preliminary standing stock and
biomass estimates of seabirds on the subarctic
Pacific region. Pages 589-611 <em>in</em> A. Y. Takenouti
et al., eds. Biological oceanography of the North
Pacific. Idemitsu Shoten, Tokyo.</p>

<p>Savile, D. B. O. 1972. Evidence of tree nesting by
the marbled murrelet in the Queen Charlotte Islands.
Can. Field-Nat. 86(4):389-390.</p>

<p>Sladen, W. J. L., and R. E. LeResche. 1970. New
and developing techniques in antarctic ornithology.
Pages 585-616 <em>in</em> M. W. Holdgate, ed. Antarctic
Ecology. Vol. I. Academic Press.</p>

<p>Sowl, L. W., and J. C. Bartonek. 1974. Seabirds&mdash;Alaska's
most neglected resource. Trans. N. Am.
Wildl. Nat. Resour. Conf. 39:117-126.</p>

<p>Stander, G. H., and J. A. V. Venter. 1968. Oil pollution
in South Africa. Pages 251-259 <em>in</em> Proceedings
of the International Conference on Oil Pollution
of the Sea.</p>

<p>Swartz, L. G. 1966. Sea-cliff birds. Pages 611-678 <em>in</em>
N. J. Wilimovsky and J. N. Wolfe, eds. Environment
of the Cape Thompson region, Alaska.
U.S.A.E.C., Div. Tech. Inf., Oak Ridge, Tennessee.</p>

<p>Thompson, D. Q., and R. A. Person. 1963. The eider
pass at Point Barrow, Alaska. J. Wildl. Manage.
27(3):348-356.</p>

<p>Uspenskii, S. M. 1964. Present day problems of nature
conservation in the Arctic. Problems of the
North 7:171-178.</p>

<p>Watson, G. E., and G. J. Divoky. 1972. Pelagic bird
and mammal observations in the eastern Chukchi
Sea, early fall 1970. Pages 111-172 <em>in</em> C. I. Merton
et al., eds. An ecological survey in the eastern
Chukchi Sea. U.S. Coast Guard Oceanogr. Rep.
50.</p>

<p>Williamson, F. S. L., M. C. Thompson, and J. Q.
Hines. 1966. Pages 437-480 <em>in</em> N. J. Wilimovsky
and J. N. Wolfe, eds. Environment of the Cape
Thompson region, Alaska. U.S.A.E.C., Div. Tech.
Inf., Oak Ridge, Tennessee.</p></div>

<p><span class="pagenum"><a name="Page_195" id="Page_195">[Pg 195]</a></span></p>
<div class="chapter"></div>
<hr class="chap" />




<h3>Mortality to Marine Birds Through Commercial Fishing</h3>


<p class="center">by</p>

<p class="center">Warren B. King</p>

<p class="center">International Council for Bird Preservation<br />
Smithsonian Institution, Washington, D.C.</p>

<p class="center">R. G. B. Brown</p>

<p class="center">Canadian Wildlife Service<br />
Dartmouth, Nova Scotia, Canada</p>

<p class="center">and</p>

<p class="center">Gerald A. Sanger<a name="FNanchor_52_52" id="FNanchor_52_52"></a><a href="#Footnote_52_52" class="fnanchor">[52]</a></p>

<p class="center">U.S. National Marine Fisheries Service<br />
Seattle, Washington
</p>


<h4>Abstract</h4>

<div class="blockquot">

<p>Commercial fishing has been responsible for incidental mortality of seabirds
for centuries, but with the advent of offshore salmon gill-net fishing in the North
Pacific in 1952 and in the North Atlantic in 1965, the magnitude of this kill has
increased, and there is strong indication that populations of some seabirds are
being adversely affected. Murres (<em>Uria</em> spp.) are most frequently killed, although
several other species are caught in lesser numbers. The seabird resources of
several nations are involved in this mortality. Longline fishing and inshore gill-net
fishing for salmon and cod also are responsible for mortality of seabirds,
although usually not in significant numbers.</p></div>

<p>That the activities of commercial fishermen
have caused mortality of marine birds surprises
no one nowadays. Traditions of exploitation
of marine birds by fishermen date from
previous centuries, and fishing has contributed
to the extinction of some species. For
example, great auks <em>(Pinguinus impennis)</em> and
other birds were used as food by fishermen
fishing for Atlantic cod <em>(Gadus morhua)</em> on
the Grand Banks of Newfoundland since the
beginning of that fishery in the early 16th century
(Collins 1884; Lucas 1890). The last great
auk died in 1844, but smaller species, such as
storm-petrels (Hydrobatidae), greater shearwaters
<em>(Puffinus gravis)</em>, and black-legged
kittiwakes <em>(Rissa tridactyla)</em>, were used for
food until rather recently (Templeman 1945).
This practice has now lapsed, however.</p>


<h4>Inshore Fisheries</h4>

<p>Until the advent of the offshore salmon gill-net
fisheries in the North Pacific in 1952 and
the North Atlantic in 1965, most seabird mortality
in these areas was the result of local
fishing close to shore. Several records of such
bird mortality have been published. For
example, 8,000-10,000 seabirds&mdash;presumably
mostly alcids&mdash;were reported caught annually
off Hammerfest in northern Norway (Holgersen
1961). E. Brun (personal communication)
reported that the longline fishery off the coast
of Norway is having serious consequences on
Norwegian populations of murres.</p>
<p><span class="pagenum"><a name="Page_196" id="Page_196">[Pg 196]</a></span></p>
<p>Numbers of alcids are caught in nets set for
Atlantic salmon <em>(Salmo salar)</em> around the
coasts of Ireland and Scotland (Biddy 1971).
A similar situation exists along the west
Greenland coast, although it is overshadowed
there by the direct exploitation of huge numbers
of alcids by hunting. Nonetheless, in
1967 for example, 15,000 alcids were recovered
from fish nets in southwestern Greenland,
where they were sold as food (Evans and
Waterston 1976). The annual salmon catch of
the west Greenland inshore fishery has fluctuated
between 60 and 1,500 metric tons and
has averaged about 1,000 tons. There are no
data comparing the relative catch of birds and
fish in this fishery.</p>

<p>Atlantic cod follow the spawning capelin
<em>(Mallotus villosus)</em> inshore along the east
coast of Newfoundland in late June and early
July. They are traditionally fished with traps
and handlines along this coast, but there has
been a recent trend toward using drift nets set
on the bottom. Since alcids feed extensively
on capelin at this time, many are caught in the
cod nets set in areas close to the large colonies
off Witless Bay (D. N. Nettleship, personal
communication). Additionally, gill nets are
set at the surface for salmon in the same area.
Common murres <em>(Uria aalge)</em> are most affected,
but Atlantic puffins <em>(Fratercula arctica)</em>
are also taken.</p>

<p>There are as yet no estimates of the total alcid
mortality from this fishery, although the
annual catch of birds is believed to be smaller
during the present than during the last
decade because the fishing effort is reduced,
and fishermen in the area now avoid setting
nets near alcid concentrations because of the
annoyance of having to remove the birds from
their nets. The Witless Bay colonies contain
over 77,000 pairs of common murres, or 11%
of the total eastern North American population,
and over 235,000 pairs of Atlantic
puffins, or 71% of the North American population
outside of Greenland (Brown et al.
1975). The potential danger is obvious.</p>

<p>There are few data on mortality of seabirds
from inshore commercial fisheries in the
North Pacific. Some mortality of alcids has
been shown to take place in Cook Inlet,
Alaska, from beach-netting for Pacific salmon
(<em>Oncorhynchus</em> spp.) adjacent to seabird
rookeries and from drift-netting in the inlet
(D. A. Snarski, personal communication), but
this mortality has not been quantified.</p>

<p>Bilateral agreements between the United
States and Japan, the U.S.S.R. and the Republic
of Korea, concerning the use of inshore
waters adjacent to some of the Aleutian Islands,
Kodiak, Nunivak, St. Matthew, St.
George, Kayak, and Forrester Islands permit
trawling, longlining, and loading fish and fuel
in some of these areas and at certain periods.
Although these activities may affect the seabirds
of these areas, the extent of the effects
are not known (U.S. Department of the Interior,
Alaska Planning Group 1974). Murie
(1959) indicated, however, that the disappearance
of the ancient murrelet <em>(Synthliboramphus
antiquus)</em> from Sanak Island, Gulf of
Alaska, was probably due as much to fisheries
as to the blue fox industry. It has been suggested
that the Japanese murrelet <em>(Synthliboramphus
wumizusumi)</em> may have declined
as the result of fishing activities near breeding
sites off the coast of Japan (Bourne 1971).</p>


<h4>Atlantic Offshore
Gill-net Fishery</h4>

<p>In 1965, Denmark began an offshore gill-net
fishery for Atlantic salmon in the Davis
Strait off the coast of west Greenland. The
offshore fishery catch increased from 36
metric tons in 1965 to more than 1,200 metric
tons in 1969, and then gradually decreased.</p>

<p>The fact that large numbers of seabirds&mdash;almost
entirely thick-billed murres <em>(Uria lomvia)</em>&mdash;were
being drowned in the salmon gill
nets was brought to the attention of the International
Council for Bird Preservation at its
15th World Conference in 1970. The Council's
recommendation was submitted to the Danish
government and stated: "... having noted
that during the 1969 fishing season about
250,000 individuals of Brunnich's guillemot
or thick-billed murre <em>(Uria lomvia)</em>, a pelagic
diving bird, were caught in these drift nets
and drowned, which number represents no
less than 25 percent of the Greenland population
and exceeds its annual reproductive
capacity; urges the Danish Government, and
the national governments of all other countries
involved in this fishing, to take all possible
measures to eliminate this very serious
problem."</p>

<p><span class="pagenum"><a name="Page_197" id="Page_197">[Pg 197]</a></span></p>

<p>The figures in the recommendation were not
supported by research; they appeared instead
to have been derived from the observed mortality
on an offshore fishery vessel in 1965,
which was then related to the salmon catch on
that vessel and applied to the total catch of
the inshore fishery in 1964 (Anonymous
1969). Studies in 1969 and 1970 by the
Fisheries Research Board of Canada finally
gave a firm basis for the earlier, though
poorly substantiated concern. On the basis of
the assumption that the ratio of salmon to
murres caught in experimental fishing applied
to the commercial fishery, an estimate of an
annual mortality of 0.5 million murres
(&plusmn;50%) was made on the basis of a salmon
catch of 1,200 metric tons (Tull et al. 1972).</p>

<p>The birds being killed were from colonies in
west Greenland, the eastern Canadian Arctic,
and possibly east Greenland and Spitzbergen.
Coupled with other known causes of mortality
(particularly hunting on the Greenland and
Newfoundland coasts, an unknown but definitely
substantial kill from oil pollution, a calculated
mortality of pre-fledging young, and
an unknown natural post-fledging mortality)
there is no doubt that the estimated annual
production of 1.5 million chicks from west
Greenland and the Canadian Arctic was less
than the estimated total annual mortality
(Tull et al. 1972). Thus, it comes as no surprise
that recent surveys of murre populations of
west Greenland and the Canadian Arctic have
revealed massive declines in numbers (Evans
and Waterston 1976; D. N. Nettleship, personal
communication). It is therefore encouraging
news that, as a result of an agreement
between the United States and Denmark, the
offshore salmon gill-net fishery was terminated
at the end of the 1975 season. The inshore
fishery remained in operation, however,
but was restricted to a total annual salmon
catch of 1,100 metric tons.</p>


<h4>Pacific Offshore Salmon
Gill-net Fishery</h4>

<p>In the north Pacific Ocean, the Japanese
gill-net fisheries for salmon (<em>Oncorhynchus</em>
spp.), which have operated since 1952, might
be expected to have an even more destructive
effect on seabirds, since the annual salmon
catch by the three Japanese salmon drift-net
fisheries was about one hundred times that in
west Greenland in recent years. The first, the
mothership fishery, comprising about 369
catcher-boats<a name="FNanchor_53_53" id="FNanchor_53_53"></a><a href="#Footnote_53_53" class="fnanchor">[53]</a> serviced by 11 mother-ships,
operates west of 175&deg;W and generally north
of 46&deg;N during the summer. The second, the
land-based fishery of about 325 ocean-going
vessels, operates west of 175&deg;W and south of
46&deg;N; and the third, the coastal fishery, made
up of about 1,380 short-haul vessels, operates
off Hokkaido. The relative salmon catches of
these three fisheries is on the order of
1:1.34:0.65.</p>

<p>Data collected on U.S. National Marine
Fisheries Service research vessels in 1974 (obtained
through the cooperation of Francis M.
Fukuhara and Richard Bakkala, Northwest
Fisheries Center, Seattle, Washington) give,
for the first time, an estimate of the magnitude
of the incidental seabird kill of the Japanese
salmon gill-net fishery. The kill data are
available only from the mothership area and
from an area east of it to 165&deg;W. The Japanese
salmon fishery is restricted to waters
west of 175&deg;W by agreement with the United
States. Bird kills from the other two areas
may be estimated by the relative salmon
catch figures for the areas, assuming that seabird
densities, species composition, and catch
effort are similar.</p>

<p>An estimate of the total kill of seabirds in
the mothership area may be made by calculating
the bird mortality per length of gill-net set
by research vessels, multiplied by the total
length of gill nets set by the 369 catcher-boats
of the Japanese mothership fishery. About
4,666 km of nets are set and retrieved daily
during the approximately 65-day fishing season.
The estimated annual mortality in the
mothership area is about 75,000 to 250,000
birds. The lower number is based on data from
10 cruises (450 km of nets set) west of 175&deg;W,
within the area of the mothership fishery. The
higher number is based on data from 20
cruises, including those in the first figure,
west of 165&deg;W, and covering the period 18
April to 3 September 1974 (956 km of nets
<span class="pagenum"><a name="Page_198" id="Page_198">[Pg 198]</a></span>set), whereas the mothership fishery usually
operates between mid-May and late July. Assuming
similar seabird densities and catch per
unit of effort in the areas of the land-based
and coastal fisheries, the estimated annual
mortality is between 214,500 and 715,000
birds. Since 1952, as many as 4.7 million birds
may have been killed by the Japanese salmon
gill-net fishery. It must be stressed that seabird
densities and catch per unit of effort are
not known to be similar for the areas in question;
consequently the projection of bird kill
figures from one area to all three is
speculative.</p>

<p>In the mothership area and adjacent seas to
the east, in addition to murres (48% of birds
killed), significant numbers of shearwaters,
<em>Puffinus</em> spp. (27%); puffins (9%); and fulmars,
<em>Fulmarus glacialis</em> (5%) are killed, as
are lesser numbers of small alcids, albatrosses
(<em>Diomedea</em> spp.), and storm-petrels. The
murres and puffins taken in the mothership
area are of U.S. and U.S.S.R. origin, and the
shearwaters come from New Zealand, Australia,
and Chile. In the coastal fishery area,
Japanese and U.S.S.R. alcids are taken.
Available knowledge of the populations of the
species making up the bulk of the kill, which
has been taking place for 20 years, is insufficient
to suggest whether their annual reproduction
can tolerate such losses. Prohibition
of fishing within 160 km of North Pacific seabird
breeding islands would help to decrease
losses of alcids of U.S. origin, but would not
help the shearwaters from the southern
hemisphere.</p>

<p>Comparison of statistics of the salmon
fisheries and associated bird kills from the
North Atlantic and the North Pacific shows
that the North Atlantic salmon fishery is concentrated
in a relatively small area which is
also along a major migration pathway of
murres. Virtually all seabird mortality is confined
to one species. Enough information is at
hand to indicate that this cause of mortality,
in conjunction with others known to be significant,
is causing a drastic decline in the
thick-billed murre population.</p>

<p>In the North Pacific, on the other hand, the
fishery is more widely dispersed and the ratio
of seabirds to salmon caught is much lower.
Furthermore, several species are subject to
mortality. No information is available to indicate
whether alcid populations (which make
up two-thirds of the kill) are stable or decreasing.
The shearwaters, primarily sooty <em>(Puffinus
griseus)</em> and slender-billed <em>(P. tenuirostris)</em>,
appear to be able to sustain not only
these losses but also a sizable harvest of birds
of the year (the so-called muttonbirds) on
their New Zealand and Australian breeding
grounds. Thus, although the latest estimates
of the total standing stock of seabirds in the
North Pacific in summer may be as high as
100 million (Sanger and King, this volume),
and thus only about 1 of every 200 birds in the
North Pacific region may be caught, the fact
that a few species, particularly murres, are selectively
caught raises questions about the
impact of this fishery on populations of these
species.</p>

<p>The U.S.-Japan Migratory Bird Convention
of 1973 specifically protects all of the species
thought to be subject to gill-net mortality in
the Pacific. Thus, the Japanese salmon fleet
apparently operates in constant violation of
this convention.</p>


<h4>Mortality of Albatrosses</h4>

<p>A recent analysis of recoveries of Laysan
albatrosses <em>(Diomedea immutabilis)</em> and
black-footed albatrosses <em>(D. nigripes)</em> banded
on the northwest Hawaiian chain from 1937
to 1969 showed that of a sample of 532 recovered
birds, 57.4% of the Laysan species
and 49.5% of the black-footed species were
caught on fishhooks or in nets, and the means
of recovery of many additional birds was
thought to have been the same (Robbins and
Rice 1974). It is likely that the large majority
are taken on Japanese and U.S.S.R. longline
tuna fishing gear. Although this cause of mortality
is insignificant in terms of the total
population of either species (only 0.2% of
banded Laysan and 0.8% of banded black-footed
albatrosses have been recovered by
any means away from their breeding
grounds), these species are protected by the
U.S.-Japan Migratory Bird Convention.
Furthermore, the possibility exists that individuals
of the endangered short-tailed albatross
<em>(Diomedea albatrus)</em> might be killed in
this manner.</p>

<p><span class="pagenum"><a name="Page_199" id="Page_199">[Pg 199]</a></span></p>


<h4>Long-term Effects of
Developing Capelin Fishery in
Northwest Atlantic</h4>

<p>Capelin are important food fish for many
seabirds in the northwest Atlantic, and the
development and expansion of this fishery off
eastern Canada must be carefully monitored.
In theory, the capelin fishery ought not to
seriously affect the birds because it is designed
to exploit a surplus of capelin artificially
created by the overfishing of Atlantic
cod, the capelin's most important predator. It
is hoped that there is no prospect of the overfishing
that may have contributed to the recent
drastic decline of the Peruvian anchovy
<em>(Engraulis ringens)</em> and the seabird species dependent
on it (Paulik 1971). However, the relative
influence of overfishing and "El Ni&ntilde;o"
oceanographic conditions on the decline remains
unclear. North Atlantic seabirds are, in
any case, more versatile in their feeding
habits (Belopol'skii 1961). But, the threat
may be a subtle one. The important point to
the seabirds may well be not merely the survival
of a reasonably large capelin stock, but
the presence of capelin schools in high densities
in certain areas or at certain seasons.
Lower densities might, for example, reduce
the foraging efficiency of breeding birds, and
hence their nesting success. The very large
common murre colony on Funk Island, Newfoundland
(500,000 pairs: Tuck 1960), might
be particularly vulnerable. It lies close to an
area where capelin are especially abundant
and one which is already being exploited by
the developing fishery.</p>


<h4>References</h4>

<div class="hangindent">

<p>Anonymous. 1969. Seabird slaughter. Sports Fish.
Inst. Bull. 203:5.</p>

<p>Belopol'skii, L. O. 1961. Ecology of sea colony birds
of the Barents Sea. (Transl. from Russian.) Israel
Program for Scientific Translations, Jerusalem.
346 pp.</p>

<p>Biddy, C. J. 1971. Auks drowned by fishnets. Seabird
Rep. No. 2.</p>

<p>Bourne, W. R. P. 1971. General threats to seabirds.
ICBP [Int. Counc. Bird Preservation] Bull.
11:200-219.</p>

<p>Brown, R. G. B., D. N. Nettleship, P. Germain, C. E.
Tull, and T. Davis. 1975. Atlas of eastern Canadian
seabirds. Canadian Wildlife Service, Ottawa.
220 pp.</p>

<p>Collins, J. W. 1884. Notes on the habits and methods
of capture of various species of seabirds that
occur on the fishing banks off the east coast of
North America and which are used as bait for
catching codfish by New England fishermen. U.S.
Comm. Fish Fish. Rep. 1882:311-335.</p>

<p>Evans, P., and G. Waterston. 1976. The decline of
the thick-billed murre in Greenland. Polar Rec.
18:283-286.</p>

<p>Holgersen, H. 1961. On the movements of Norwegian
<em>Uria aalge</em>. (In Norwegian, English summary.)
Sterna 4:229-240.</p>

<p>Lucas, F. A. 1890. Expedition to the Funk Island,
with observations on the history and anatomy of
the Great Auk. Rep. U.S. Natl. Mus., 1887-1888:493-529.</p>

<p>Murie, O. J. 1959. Fauna of the Aleutian Islands
and Alaska Peninsula. U.S. Fish Wildl. Serv., N.
Am. Fauna 61. 406 pp.</p>

<p>Paulik, A. J. 1971. Anchovies, birds, and fishermen
in the Peru Current. Pages 156-185 <em>in</em> W. W. Murdoch,
ed. Environmental Resources and Society.
Sinauer Associates, Inc., Stamford, Conn.</p>

<p>Robbins, C. S., and D. W. Rice. 1974. Recoveries of
banded Laysan albatrosses <em>(Diomedea immutabilis)</em>
and black-footed albatrosses <em>(D. nigripes)</em>.
Pages 232-271 <em>in</em> W. B. King, ed. Pelagic studies
of seabirds in the Central and Eastern Pacific
Ocean. Smithson. Contrib. Zool. 158.</p>

<p>Templeman, W. 1945. Observations on some Newfoundland
seabirds. Can. Field-Nat. 59:136-147.</p>

<p>Tuck, L. M. 1960. The murres. Canadian Wildlife
Service, Ottawa. 260 pp.</p>

<p>Tull, C. E., P. Germain, and A. W. May. 1972. Mortality
of thick-billed murres in the west Greenland
salmon fishery. Nature (Lond.) 237 (5349):42-44.</p>

<p>U.S. Department of the Interior, Alaska Planning
Group. 1974. Final environmental impact statement,
proposed Alaska Coastal National Wildlife
Refuges. 678 pp.</p></div>

<p><span class="pagenum"><a name="Page_200" id="Page_200">[Pg 200]</a></span></p>


<div class="footnotes"><h4>FOOTNOTES:</h4>

<div class="footnote">

<p><a name="Footnote_52_52" id="Footnote_52_52"></a><a href="#FNanchor_52_52"><span class="label">[52]</span></a> Present address: U.S. Fish and Wildlife Service,
Office of Biological Services&mdash;Coastal Ecosystems.
1011 E. Tudor Road, Anchorage, Alaska
99503.</p></div>

<div class="footnote">

<p><a name="Footnote_53_53" id="Footnote_53_53"></a><a href="#FNanchor_53_53"><span class="label">[53]</span></a> This figure is based on data through 1971. Since
then, the number of catcher-boats has decreased
to 332 in 1974 (F. M. Fukuhara, personal communication).</p>

<p><span class="pagenum"><a name="Page_201" id="Page_201">[Pg 201]</a></span></p></div></div>
<div class="chapter"></div>
<hr class="chap" />




<h3><a name="Interactions_Among_Marine_Birds_and_Commercial" id="Interactions_Among_Marine_Birds_and_Commercial">Interactions Among Marine Birds and Commercial
Fish in the Eastern Bering Sea</a></h3>


<p class="center">by</p>

<p class="center">Richard R. Straty and Richard E. Haight</p>

<p class="center">National Marine Fisheries Service<br />
Auke Bay Fisheries Laboratory<br />
Auke Bay, Alaska 99821
</p>


<h4>Abstract</h4>

<div class="blockquot">

<p>The high primary and secondary productivity of the eastern Bering Sea makes
it one of the greatest producers of commercial fish and largest congregating
areas of marine birds in the world. The fish and birds are so interrelated that fluctuations
in the abundance of one may well be responsible for changes in the abundance
of the other. The seasonal and annual variation in the impact of birds on
fish is a function of the life history, food habits, growth rate, and final size of the
fish species of concern and of the distribution, abundance, and feeding habits of
bird populations&mdash;plus the effects of the environment on these factors. Stages in
the life history of some of the important commercial fish and shellfish of the
Bering Sea directly or indirectly influenced by marine birds are identified.</p></div>

<p>The eastern Bering Sea is one of the world's
richest fish-producing areas and is also one of
the world's major congregating areas for
marine birds. The large extent of the continental
shelf and the climatic and oceanographic
characteristics of the eastern Bering
Sea combine to make this region extremely
productive biologically. The distribution and
abundance of plankton, benthos, and fish determine
the distribution, time, and character
of the migration of marine birds in the eastern
Bering Sea (Shuntov 1961). Several studies
have illustrated the close relation between
marine birds and the biological properties of
surface waters (Tuck 1960; Bourne 1963; Solomensen
1965). Spatial and temporal variations
in the abundance of the fish families Clupeidae
(herring), Gadidae (codfish), Osmeridae
(capelin), and Ammodytidae (sand lance) are
thought to be major determinants of the
breeding seasons, breeding places, and movements
of boreal seabirds (Ashmole 1971). The
timing of breeding among larids and alcids is
related to the seasonal changes in the surface
waters inhabited by Ammodytidae and Clupeidae
in the North Sea (Pearson 1968).</p>

<p>The eastern Bering Sea contains members
of these and other fish families that are extensively
exploited by man; the fish are also important
as forage for other species of commercial
fish, marine mammals, and marine birds.
During some part of their life cycles, all fish
species feed on plankton, nekton, benthos, or
other fishes.</p>

<p>The incidental use or dependence of marine
birds on commercial fish and the items on
which the fish feed account for the major interaction
between man and these two groups
of animals.</p>

<p>In this paper, we consider how marine birds
and fish interact. Although some of what we
present is only speculative, we identify certain
areas that have received little or no scientific
study, areas in which further research is
needed for a better understanding of the role
of commercial fish in the ecology and dynamics
of marine birds in the eastern Bering
Sea.</p>

<p><span class="pagenum"><a name="Page_202" id="Page_202">[Pg 202]</a></span></p>


<h4>Commercial Fish Resources
of the Eastern Bering Sea</h4>

<p>Most of the fishing in the eastern Bering
Sea is done by Japan and the Soviet Union.
Japan resumed fishing in the Bering Sea in
1953 (7 years after World War II), the Soviet
Union started fishing in the region in 1959,
and since the early 1960's both nations have
accelerated their exploitation of Bering Sea
fish stocks (Chitwood 1969).</p>

<p>Species of major concern to Japan and the
Soviet Union include fish&mdash;walleye pollock
<em>(Theragra chalcogramma)</em>, yellowfin sole
<em>(Limanda aspera)</em>, Pacific cod <em>(Gadus macrocephalus)</em>,
Pacific ocean perch <em>(Sebastes
alutus)</em>, Pacific herring <em>(Clupea harengus pallasi)</em>,
and sablefish <em>(Anoplopoma fimbria)</em>&mdash;and
snow crabs (<em>Chionoecetes</em> spp.). The distribution
of the principal species being harvested
in Bristol Bay and the eastern Bering
Sea are shown in Figs. 1, 2, and 3. The weight
of each of the major species in the total
catches made by foreign and domestic fishermen
in 1973 is shown in Table 1. In 1972, the
catch of commercial finfish in the eastern
Bering Sea alone amounted to 5% of the total
world catch of marine fishes (H. Larkins, personal
communication).</p>

<p>Most species of commercial fish in the
Bering Sea are in a state of decline or in a depressed
condition from overexploitation
(Table 1). This is indicated by a reduction in
the catch per unit of effort and in the mean
size of fish in the commercial catch (H.
Larkins, personal communication). The notable
exception is the king crab (<em>Paralithodes</em>
sp.), which has increased in abundance in recent
years as a result of reduced foreign fishing.</p>

<div class="center">
<table border="0" cellpadding="4" cellspacing="0" summary="Table 1.
Foreign and domestic catch of fish and shellfish in the eastern Bering
Sea, including Bristol Bay, 1973.">
  <caption>Table 1. <em>Foreign and domestic catch of fish and
  shellfish in the eastern Bering Sea, including Bristol Bay,
  1973.</em></caption>
  <tr>
    <th>Species</th>
    <th>Catch (metric tons)</th>
  </tr>
  <tr>
    <td>Fish</td>
  </tr>
  <tr>
    <td class="tdp">Pollock</td>
    <td class="tdr">1,500,000</td>
  </tr>
  <tr>
    <td class="tdp">Flatfish</td>
    <td class="tdr">125,000</td>
  </tr>
  <tr>
    <td class="tdp">Pacific cod</td>
    <td class="tdr">45,000</td>
  </tr>
  <tr>
    <td class="tdp">Herring</td>
    <td class="tdr">35,033</td>
  </tr>
  <tr>
    <td class="tdp">Salmon</td>
    <td class="tdr">11,785</td>
  </tr>
  <tr>
    <td class="tdp">Sablefish</td>
    <td class="tdr">7,000</td>
  </tr>
  <tr>
    <td class="tdp">Pacific halibut</td>
    <td class="tdr">222</td>
  </tr>
  <tr>
    <td class="tdp">Other</td>
    <td class="tdr">40,000</td>
  </tr>
  <tr>
    <td>Shellfish</td>
  </tr>
  <tr>
    <td class="tdp">King crabs</td>
    <td class="tdr">26,798</td>
  </tr>
  <tr>
    <td class="tdp">Snow crabs</td>
    <td class="tdr">17,694</td>
  </tr>
  <tr>
    <td class="tdp">Shrimp</td>
    <td class="tdr">Minor</td>
  </tr>
</table></div>

<div class="figcenter" style="width: 700px;">
<img src="images/p_202a.jpg" width="700" height="454" alt="" />
<div class="caption"><p><strong>Fig. 1.</strong> Areas of major concentrations of ground fish
(Pacific pollock, halibut, yellowfin sole, rock sole,
flathead sole, Pacific ocean perch, and Pacific
cod) in Bristol Bay and the Bering Sea.</p></div>
</div>

<div class="figcenter" style="width: 700px;">
<img src="images/p_202b.jpg" width="700" height="471" alt="" />
<div class="caption"><p><strong>Fig. 2.</strong> Areas of major winter and spring concentrations
of Pacific herring in Bristol Bay and the
Bering Sea.</p></div>
</div>

<div class="figcenter" style="width: 700px;">
<img src="images/p_202c.jpg" width="700" height="476" alt="" />
<div class="caption"><p><strong>Fig. 3.</strong> Areas of major concentrations of king and
snow crab in Bristol Bay and the Bering Sea.</p></div>
</div>

<p><span class="pagenum"><a name="Page_203" id="Page_203">[Pg 203]</a></span></p>


<h4>Routes of Interaction
Between Marine Birds and
Commercial Fish</h4>

<p>The obvious ways in which marine birds and
fish of commercial importance interact in the
eastern Bering Sea are illustrated by the simplified
food web diagram in Fig. 4. The major
animal groups and species included in two of
the categories in this figure&mdash;secondary producers
(invertebrate forage) and intermediate
carnivores (commercial and forage marine fish
and shellfish)&mdash;are as follows:</p>


<ul><li class="ifrst"><em>Secondary producers</em></li>

<li class="indx">Zooplankton and micronekton</li>
<li class="isub1">Copepods</li>
<li class="isub2"><em>Calanus</em> spp.</li>
<li class="isub2"><em>Eucalanus</em> spp.</li>

<li class="isub1 indx">Euphausiids</li>
<li class="isub2"><em>Thysanoessa</em> spp.</li>

<li class="isub1 indx">Amphipods</li>
<li class="isub2"><em>Parathemisto</em> spp.</li>
<li class="isub2"><em>Gammarus</em> spp.</li>

<li class="isub1 indx">Pteropods</li>
<li class="isub2"><em>Spiratella</em> spp.</li>
<li class="isub2"><em>Clione</em> spp.</li>

<li class="isub1 indx">Chaetognaths</li>
<li class="isub2"><em>Sagitta</em> spp.</li>

<li class="indx">Benthos</li>
<li class="isub1">Polychaetes</li>
<li class="isub2"><em>Nereis</em> spp.</li>
<li class="isub2"><em>Euroe</em> spp.</li>

<li class="isub1 indx">Molluscs</li>
<li class="isub2"><em>Mytilus edulis</em></li>
<li class="isub2"><em>Tonicella</em> spp.</li>
<li class="isub2"><em>Fusitriton oregonensis</em></li>

<li class="isub1 indx">Echinodermata</li>
<li class="isub2"><em>Strongylocentrotus</em> spp.</li>

<li class="isub1 indx">Crustacea</li>
<li class="isub2">Gammaridae</li>
<li class="isub2">Mysidae</li>
<li class="isub2"><em>Idothea</em> spp.</li>
<li class="isub2"><em>Pagurus</em> spp.</li>
<li class="isub2"><em>Hapalogaster</em> spp.</li>
<li class="isub2"><em>Sclerocrangon</em> spp.</li>

<li class="indx"><em>Intermediate carnivores</em></li>

<li class="indx">Eggs (littoral, adhesive)</li>
<li class="isub1">Clupeidae</li>

<li class="indx">Pelagic larvae</li>
<li class="isub1">Gadidae</li>
<li class="isub1">Pleuronectidae</li>
<li class="isub1">Osmeridae</li>
<li class="isub1">Ammodytidae</li>
<li class="isub1">Salmonidae</li>
<li class="isub1">Gadidae</li>
<li class="isub1">Pandalidae</li>

<li class="indx">Juvenile and small adults</li>
<li class="isub1">Clupeidae</li>
<li class="isub1">Osmeridae</li>
<li class="isub1">Ammodytidae</li>
<li class="isub1">Salmonidae</li>
<li class="isub1">Gadidae</li>
<li class="isub1">Pandalidae</li>

<li class="indx">Large adults</li>
<li class="isub1">Clupeidae</li>
<li class="isub1">Gadidae</li>
<li class="isub1">Pleuronectidae</li>
<li class="isub1">Salmonidae</li>
<li class="isub1">Scorpaenidae</li>
<li class="isub1">Lithodidae</li>
<li class="isub1">Majidae</li>
<li class="isub1">Pandalidae</li>

<li class="indx">Marine birds</li>
<li class="isub1">Alcidae</li>
<li class="isub1">Procellariidae</li>
<li class="isub1">Laridae</li>
<li class="isub1">Phalacrocoracidae</li>
</ul>

<div class="figcenter" style="width: 700px;">
<img src="images/p_203.jpg" width="700" height="407" alt="" />
<div class="caption"><p>Fig. 4. Food web in the eastern Bering Sea, showing
routes of interaction between marine birds and
the various life history stages of commercial fish
and shellfish.</p></div>
</div>

<p><span class="pagenum"><a name="Page_204" id="Page_204">[Pg 204]</a></span></p>

<p>In our discussion, we mainly consider predation
by birds on commercial fish and competition
between birds and commercial fish
for food. The extent of these interactions determines
the potential for birds and fish to influence
each other's abundance. The extent of
the interactions also determines the impact of
man's commercial harvest of fish on the abundance
of birds or of the bird's harvest on the
abundance of fish.</p>

<p>The extent of the interaction between
marine birds and commercial fish depends on
the abundance, distribution, feeding habits,
and life history of the fish species of concern.
We have limited our discussion to examples of
the major commercial pelagic and demersal
fish and shellfish of the eastern Bering Sea.
We also use as examples those species of
marine birds whose abundance in the eastern
Bering Sea and feeding habits give them the
greatest potential for influence on, or being influenced
by, fish abundance.</p>


<h4>Abundance and Feeding Habits
of Marine Birds in the
Eastern Bering Sea</h4>

<p>Information on the general abundance and
distribution of the most important marine
birds in the eastern Bering Sea in the summer
and winter is scattered among many published
and unpublished reports: Shuntov
(1961, 1966), Sanger (1972), Bartonek and
Gibson (1972), and Ogi and Tsujita (1973); and
surveys by D. T. Montgomery and W. E. Oien
("Bristol Bay waterbird survey, 1972," unpublished
report of the U.S. Bureau of Sport
Fisheries and Wildlife, Alaska area) and by
J. G. King and D. E. McKnight (1969, "A
waterbird survey in Bristol Bay and proposals
for future studies," unpublished report
of the U.S. Bureau of Sport Fisheries and
Wildlife and the Alaska Department of Fish
and Game, Juneau, Alaska).</p>

<p>In summer, the most abundant birds appear
to be the procellariids, mainly the slender-billed
shearwater <em>(Puffinus tenuirostris)</em> and
Pacific fulmar <em>(Fulmarus glacialis)</em>; the alcids,
mainly the common murre <em>(Uria aalge)</em>,
thick-billed murre <em>(U. lomvia)</em>, tufted puffin
<em>(Lunda cirrhata)</em>, horned puffin <em>(Fratercula
corniculata)</em>, and the ancient murrelet <em>(Synthliboramphus
antiquus)</em>; and the larids, mainly
the glaucous-winged gull <em>(Larus glaucescens)</em>
and the black-legged kittiwake <em>(Rissa
tridactyla)</em>.</p>

<p>In winter, the alcids and larids appear to be
the most abundant groups, the procellariids
having been reduced by the departure of the
slender-billed shearwaters for breeding
grounds in the southern hemisphere. The selection
of the types of food to be consumed by
these marine birds is a function of their morphological
and physiological adaptations and
of the resultant feeding behavior. Ashmole
(1971) classified the feeding behavior of
various genera of marine birds and the relative
importance of the kinds of food eaten by
each group; this information for some of the
Bering Sea bird species occurring in the
genera listed by Ashmole (1971) is summarized
in Fig. 5.</p>

<p>Fish and invertebrates are evidently of
moderate to major importance in the diet of
these marine birds (Fig. 5). The extent to
which a given fish species is fed upon by or is
in competition with marine birds for food is
determined by the life history of the fish.
Most pelagic and some demersal fish and
shellfish are more subject to predation by pursuit
diving birds than by birds restricted to
the near-surface waters. Invertebrates appear
to be equal to or more important than fish in
the diets of birds feeding in near-surface
waters (Fig. 5).</p>


<h4>Predation by Marine Birds</h4>

<p>The literature contains numerous accounts
of marine birds feeding on marine fish and
shellfish of commercial importance. Some
studies quantify the impact of some bird
species on certain species of commercial fish
(Outram 1958; Shaefer 1970; Wiens and Scott
1976) and shellfish (Glude 1967). Other
studies have shown that in some regions the
value of guano produced by birds may exceed
the value of the commercial fish they consume
(Jarvis 1970). Some fish of worldwide commercial
importance that are important in the
diets of marine birds are listed in Table 2.</p>

<p><span class="pagenum"><a name="Page_205" id="Page_205">[Pg 205]</a></span></p>

<div class="center">
<table border="0" cellpadding="4" cellspacing="0" summary="Table 2. Fish of worldwide commercial importance in the diets of some marine birds.">
    <caption>Table 2. <em>Fish of worldwide commercial importance in the diets of some marine birds.</em></caption>
  <tr>
    <th>Fish</th>
    <th>Shearwaters</th>
    <th> Murres</th>
    <th> Puffins</th>
    <th> Fulmars</th>
    <th> Gulls</th>
  </tr>
  <tr>
    <td>Anchovy</td>
    <td class="tdc"> X</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
  </tr>
  <tr>
    <td>Sardines</td>
    <td class="tdc"> X</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
  </tr>
  <tr>
    <td>Herring</td>
    <td class="tdc"> X</td>
    <td class="tdc"> X</td>
    <td class="tdc"> X</td>
    <td class="tdc"> X</td>
    <td class="tdc"> X</td>
  </tr>
  <tr>
    <td>Sprat</td>
    <td class="tdc"> X</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
  </tr>
  <tr>
    <td>Pilchard</td>
    <td class="tdc"> X</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
  </tr>
  <tr>
    <td>Capelin</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc"> X</td>
    <td class="tdc"> X</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc"> X</td>
  </tr>
  <tr>
    <td>Salmon</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc"> X</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
  </tr>
  <tr>
    <td>Mackerel</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc"> X</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
  </tr>
  <tr>
    <td>Pollock</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc"> X</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc"> X</td>
    <td class="tdc">&mdash;</td>
  </tr>
  <tr>
    <td>Haddock</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc"> X</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
  </tr>
  <tr>
    <td>Cod</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc"> X</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
  </tr>
</table></div>

<p>The significance of bird predation on pelagic
or demersal fish and shellfish (Fig. 5) depends
on the feeding behavior of the birds and on the
life history of the fish (e.g., distribution, abundance,
growth, and adult size). Pursuit diving
birds, such as murres and puffins, can consume
fish at greater depths than can birds
that feed near the surface, such as shearwaters,
kittiwakes, fulmars, and gulls.</p>

<div class="figcenter" style="width: 700px;">
<img src="images/p_205.jpg" width="700" height="446" alt="" />
<div class="caption"><p>Fig. 5. Feeding behavior and relative importance of
food of some groups of marine birds that occur in
the eastern Bering Sea.</p></div>
</div>


<h4>Aspects of the Life
Histories of Fish Related to
Predation by Marine Birds</h4>

<p>Fish that are pelagic during part of their
lives, such as salmon and herring, and forage
fish like smelt, capelin, and sand lance, are
vulnerable to greater predation by a wider
variety of marine birds than are bottom-dwelling
demersal fish, such as pollock, cod, sole,
ocean perch, and halibut, as well as king and
snow crabs. Some species that live on the bottom
as adults have pelagic stages during
which they are vulnerable to predation by
marine birds. Juveniles of some demersal
species (pollock, cod, halibut, some species of
sole, and king crabs) are sometimes found in
shallow water where they might be subject to
predation by birds.</p>


<h5><em>Demersal Fish and Shellfish</em></h5>

<p>The early life histories of the commercially
important demersal fish of the eastern Bering
Sea are quite different (Table 3). For example,
the eggs and larvae of Pacific halibut <em>(Hippoglossus
stenolepis)</em> generally occur at depths
greater than 100 m (Hart 1973), whereas
those of pollock and yellowfin sole are found
at or near the surface (Musienko 1963, 1970).
The eggs of Pacific cod are demersal, but the
larvae are oceanic (pelagic) and occur from 25-150
m (Mukhacheva and Zviagina 1960).</p>

<p>In their juvenile stages, many demersal fish
frequent the near-surface waters (Table 3),
where they become vulnerable to predation by
piscivorous marine birds. Juvenile pollock, for
example, form into small schools that usually
move about close to the bottom but sometimes
move into areas as shallow as 3 m.
Juvenile Pacific cod prefer the warmer water
close to shore and may be found within 10 m
of the surface (Moiseev 1953). The young of
many species of flatfish, such as yellowfin
sole, rock sole <em>(Lepidopsetta bilineata)</em>, and
flathead sole <em>(Hippoglosoides elassodon)</em>, remain
for a time in shallow warm water after
assuming a demersal existence. Yellowfin sole
2-2.5 cm in total length may be found in abundance
in areas as shallow as 5 m (Fadeev 1965;
Moiseev 1953).</p>

<p><span class="pagenum"><a name="Page_206" id="Page_206">[Pg 206]</a></span></p>

<div class="center">
<table border="1" cellpadding="4" cellspacing="0" summary="Table 3. Informal listing of life history information on
  selected species of commercial and forage fish and shellfish to
  show vulnerability to predation by marine birds.">
  <caption>Table 3. <em>Informal listing of life history information on
  selected species of commercial and forage fish and shellfish to
  show vulnerability to predation by marine birds.</em> (? indicates
  no information available.)</caption>
<thead>
  <tr>
    <th colspan="2">Fecundity</th>
    <th colspan="2">Spawning season</th>
  </tr>
  <tr>
    <th>Length of female (cm)<a name="FNanchor_54_54" id="FNanchor_54_54"></a><a href="#Footnote_54_54" class="fnanchor">[54]</a></th>
    <th>Mean no. of eggs</th>
    <th>Total period</th>
    <th>Peak period</th>
    <th>Life stage</th>
    <th>Total length<br />(cm)<a name="FNanchor_56_56" id="FNanchor_56_56"></a><a href="#Footnote_56_56" class="fnanchor">[56]</a></th>
    <th>Depth from surface<br />(m)</th>
    <th>Seasonal period of pelagic life</th>
    <th>Duration of life stages<br />(days)</th>
    <th>Source of data</th>
  </tr>
</thead>
<tbody>
  <tr>
    <td colspan="0"><strong>Walleye pollock</strong> (<em>Theragra chalcogramma</em> Pallas)</td>
  </tr>
  <tr>
    <td rowspan="3" class="tdc">31-35</td>
    <td rowspan="3" class="tdr">95,700</td>
    <td rowspan="3">Feb.- June</td>
    <td rowspan="3">April-May</td>
    <td rowspan="2">Egg</td>
    <td rowspan="2" class="tdc">0.1-0.2</td>
    <td rowspan="2" class="tdc">0-10</td>
    <td rowspan="2">Feb.-June</td>
    <td class="tdc">12 at 6-7&deg;C</td>
    <td rowspan="7">Yusa 1954; Tanino et al. 1959; Kobayashi 1963; Musienko 1963, 1970; Serobaba 1968; Hart 1973<a name="FNanchor_55_55" id="FNanchor_55_55"></a><a href="#Footnote_55_55" class="fnanchor">[55]</a></td>
  </tr>
  <tr>
    <td class="tdc">20.5 at 3.4&deg;C<a name="FNanchor_57_57" id="FNanchor_57_57"></a><a href="#Footnote_57_57" class="fnanchor">[57]</a></td>
  </tr>
  <tr>
    <td>Larval</td>
    <td class="tdc">0.4-0.9</td>
    <td class="tdc">10-25</td>
    <td>March-?</td>
    <td class="tdc">&gt; 25 at 6-7&deg;C</td>
  </tr>
  <tr>
    <td class="tdc">46-50</td>
    <td class="tdr">324,400</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td>Larval</td>
    <td class="tdc">0.9-?</td>
    <td class="tdc">25-?</td>
    <td>?-Sept.</td>
    <td class="tdc">?</td>
  </tr>
  <tr>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td>Juvenile</td>
    <td class="tdc">2.2-4.1</td>
    <td class="tdc">0-?<a name="FNanchor_58_58" id="FNanchor_58_58"></a><a href="#Footnote_58_58" class="fnanchor">[58]</a></td>
    <td>Summer</td>
    <td class="tdc">&mdash;</td>
  </tr>
  <tr>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td>Juvenile</td>
    <td class="tdc">6.0-30.0</td>
    <td class="tdc">4-37</td>
    <td>Summer</td>
    <td class="tdc">&mdash;</td>
  </tr>
  <tr>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td>Adult</td>
    <td class="tdc">30.0-70.0</td>
    <td class="tdc">0-386</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
  </tr>
  <tr>
    <td colspan="4"><strong>Pacific cod</strong> (<em>Gadus macrocephalus</em> Tilesius)</td>
    <td rowspan="3">Egg</td>
    <td rowspan="3" class="tdc">0.1-0.11</td>
    <td rowspan="3" class="tdc">100-250</td>
    <td rowspan="3">Demersal</td>
    <td class="tdc">8-9 at 11&deg;C</td>
    <td rowspan="6">Moiseev 1953; Mukhacheva and Zviagina
    1960; Musienko 1970; Hart 1973<a href="#Footnote_55_55"
    class="fnanchor">[55]</a></td>
  </tr>
  <tr>
    <td rowspan="3" class="tdc">60</td>
    <td rowspan="3" class="tdr">1,200,000</td>
    <td rowspan="3">Jan.-March</td>
    <td rowspan="3" class="tdc">?</td>
    <td class="tdc">17 at 5&deg;C</td>
  </tr>
  <tr>
    <td class="tdc">28 at 2&deg;C</td>
  </tr>
  <tr>
    <td>Larval</td>
    <td class="tdc">0.5-3.2</td>
    <td class="tdc">25-150</td>
    <td>Feb.-Aug.</td>
    <td class="tdc">?</td>
  </tr>
  <tr>
    <td class="tdc">78</td>
    <td class="tdr">3,300,000</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td>Juvenile</td>
    <td class="tdc">?</td>
    <td class="tdc">10-?</td>
    <td>Summer</td>
    <td class="tdc">&mdash;</td>
  </tr>
  <tr>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td>Adult</td>
    <td class="tdc">40.0-99.0</td>
    <td class="tdc">0-900</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
  </tr>
  <tr>
    <td colspan="4"><strong>Pacific herring</strong> (<em>Clupea harengus pallasi</em> Valenciennes)</td>
    <td>Egg</td>
    <td class="tdc">0.1-0.2</td>
    <td class="tdc">0-12</td>
    <td>Demersal</td>
    <td class="tdc">10-20<a href="#Footnote_57_57" class="fnanchor">[57]</a></td>
    <td rowspan="6">Stevenson 1962; Musienko 1970; Rumyantsev and Darda
    1970; Reid 1972; Hart 1973<a href="#Footnote_55_55"
    class="fnanchor">[55]</a></td>
  </tr>
  <tr>
    <td class="tdc">20.5-22.0</td>
    <td class="tdr">26,600</td>
    <td>May-June</td>
    <td>Varies</td>
    <td>Larval</td>
    <td class="tdc">0.9</td>
    <td class="tdc">0.5-8</td>
    <td>May-June</td>
    <td rowspan="3" class="tdc">42-56</td>
  </tr>
  <tr>
    <td class="tdc">28.0-31.0</td>
    <td class="tdr">77,800</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td>Larval</td>
    <td class="tdc">1.3</td>
    <td class="tdc">0.5-8</td>
    <td>June-July </td>
  </tr>
  <tr>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td>Larval</td>
    <td class="tdc">2.5</td>
    <td class="tdc">1-6</td>
    <td>July-Aug. </td>
  </tr>
  <tr>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td>Juvenile</td>
    <td class="tdc">2.5-20.5</td>
    <td class="tdc">0-?</td>
    <td>March-Nov.</td>
    <td class="tdc">&mdash;</td>
  </tr>
  <tr>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td>Adult</td>
    <td class="tdc">20.5-31.0</td>
    <td class="tdc">0-140</td>
    <td>March-Nov.</td>
    <td class="tdc">&mdash;</td>
  </tr>
  <tr>
    <td colspan="4"><strong>Capelin</strong> (<em>Mallotus villosus</em> (Muller))</td>
    <td>Egg</td>
    <td class="tdc">0.1</td>
    <td class="tdc">&lt;20</td>
    <td>Demersal</td>
    <td class="tdc">14-?</td>
    <td rowspan="5">Clemens and Wilby 1961; Musienko 1970; Hart 1973</td>
  </tr>
  <tr>
    <td class="tdc">?</td>
    <td class="tdr">3,000</td>
    <td>June-July</td>
    <td class="tdc">?</td>
    <td>Larval</td>
    <td class="tdc">0.5-?</td>
    <td class="tdc">?</td>
    <td>June-?</td>
    <td class="tdc">?</td>
  </tr>
  <tr>
    <td class="tdc">?</td>
    <td class="tdr">6,000</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td>Juvenile</td>
    <td class="tdc">?</td>
    <td class="tdc">?</td>
    <td>March-Nov.(est.)</td>
    <td class="tdc">&mdash;</td>
  </tr>
  <tr>
    <td class="tdc">10.3</td>
    <td class="tdr">6,670</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td colspan="0"></td>
  </tr>
  <tr>
    <td class="tdc">?</td>
    <td class="tdr">60,000</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td>Adult</td>
    <td class="tdc">?</td>
    <td class="tdc">0-?</td>
    <td>March-Nov.</td>
    <td class="tdc">&mdash;<span class="pagenum"><a name="Page_207" id="Page_207">[Pg 207]</a></span></td>
  </tr>
  <tr>
    <td colspan="0"><strong>Pacific sand lance</strong> (<em>Ammodytes hexapterus</em> Pallas)</td>
  </tr>
  <tr>
    <td class="tdc">&mdash;</td>
    <td class="tdc">?</td>
    <td>June- Aug.</td>
    <td class="tdc"><a name="FNanchor_59_59" id="FNanchor_59_59"></a><a href="#Footnote_59_59" class="fnanchor">[59]</a></td>
    <td>Egg</td>
    <td class="tdc">?</td>
    <td class="tdc">?</td>
    <td>Demersal</td>
    <td class="tdc">?</td>
    <td rowspan="4">Musienko 1963, 1970; Kashkina 1970; Hart 1973</td>
  </tr>
  <tr>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td>Larval</td>
    <td class="tdc">0.7-3.4</td>
    <td class="tdc">0-?</td>
    <td>June-Sept.</td>
    <td class="tdc">?</td>
  </tr>
  <tr>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td>Juvenile</td>
    <td class="tdc">3.6-9.6</td>
    <td class="tdc">0-?</td>
    <td class="tdc">?</td>
    <td class="tdc">&mdash;</td>
  </tr>
  <tr>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td>Adult</td>
    <td class="tdc">26</td>
    <td class="tdc">0-?</td>
    <td class="tdc">?</td>
    <td class="tdc">&mdash;</td>
  </tr>
  <tr>
    <td colspan="0"><strong>Pacific ocean perch</strong> (<em>Sebastes alutus</em> (Gilbert))</td>
  </tr>
  <tr>
    <td class="tdc">26</td>
    <td class="tdr">10,000</td>
    <td>March-May</td>
    <td class="tdc">?</td>
    <td>Egg<a name="FNanchor_61_61" id="FNanchor_61_61"></a><a href="#Footnote_61_61" class="fnanchor">[61]</a></td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td rowspan="6">Paraketsov 1963; Lisovenko 1965; Lyubimova 1965;
    Kashkina 1970<a name="FNanchor_60_60" id="FNanchor_60_60"></a><a
    href="#Footnote_60_60" class="fnanchor">[60]</a></td>
  </tr>
  <tr>
    <td class="tdc">44</td>
    <td class="tdr">180,000</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td>Larval<a name="FNanchor_62_62" id="FNanchor_62_62"></a><a href="#Footnote_62_62" class="fnanchor">[62]</a></td>
    <td class="tdc">0.6-?</td>
    <td class="tdc"><a href="#Footnote_62_62" class="fnanchor">[62]</a></td>
    <td>March-Aug.</td>
    <td class="tdc">?</td>
  </tr>
  <tr>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td>Juvenile</td>
    <td class="tdc">6.2</td>
    <td class="tdc">37-128</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
  </tr>
  <tr>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td>Juvenile</td>
    <td class="tdc">10.4</td>
    <td class="tdc">37-154</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
  </tr>
  <tr>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td>Juvenile</td>
    <td class="tdc">14.7-21.3</td>
    <td class="tdc">37-230</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
  </tr>
  <tr>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td>Adult</td>
    <td class="tdc">21.3-51.0</td>
    <td class="tdc">37-420</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
  </tr>
  <tr>
    <td colspan="0"><strong>Pacific halibut</strong> (<em>Hippoglossus stenolepis</em> Schmidt)</td>
  </tr>
  <tr>
    <td class="tdc">75</td>
    <td class="tdr">101,723</td>
    <td>Oct.-March</td>
    <td class="tdc">?</td>
    <td>Egg</td>
    <td class="tdc">0.3-0.4</td>
    <td class="tdc">40-935</td>
    <td>Oct.-March</td>
    <td class="tdc">48 at ?</td>
    <td rowspan="5">Novikov 1964; Hart 1973</td>
  </tr>
  <tr>
    <td class="tdc">135</td>
    <td class="tdr">2,800,837</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td>Larval</td>
    <td class="tdc">0.8-1.5</td>
    <td class="tdc">&gt;200</td>
    <td>Nov.-May</td>
    <td rowspan="2" class="tdc">70-98</td>
  </tr>
  <tr>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td>Larval</td>
    <td class="tdc">1.5-2.9</td>
    <td class="tdc">&lt;100</td>
    <td>May-Sept.</td>
  </tr>
  <tr>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td>Juvenile</td>
    <td class="tdc">3.4-4.2</td>
    <td class="tdc">7-43</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
  </tr>
  <tr>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td>Juvenile</td>
    <td class="tdc">19-25</td>
    <td class="tdc">7-45</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
  </tr>
  <tr>
    <td colspan="0"><strong>Yellowfin sole</strong> (<em>Limanda aspera</em> (Pallas))</td>
  </tr>
  <tr>
    <td class="tdc">26.1-28.0</td>
    <td class="tdr">1,295,000</td>
    <td>June-Aug.</td>
    <td>July</td>
    <td>Egg</td>
    <td class="tdc">0.07-0.09</td>
    <td class="tdc">&gt;0</td>
    <td>June-Aug.</td>
    <td class="tdc">9.4 at 13.1&deg;C<a href="#Footnote_57_57" class="fnanchor">[57]</a></td>
    <td rowspan="3">Moiseev 1953; Pertseva-Ostraumova 1954; Musienko
    1963; Fadeev 1965; Kashkina 1965<em>a</em>, 1965<em>b</em><a
    href="#Footnote_55_55" class="fnanchor">[55]</a></td>
  </tr>
  <tr>
    <td class="tdc">40.1-42.0</td>
    <td class="tdr">3,319,500</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td>Larval</td>
    <td class="tdc">0.2-1.2</td>
    <td class="tdc">&gt;0</td>
    <td>July-Oct.</td>
    <td class="tdc">?</td>
  </tr>
  <tr>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td>Juvenile </td>
    <td class="tdc">2.1-2.5</td>
    <td class="tdc">5-15</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
  </tr>
  <tr>
    <td colspan="0"><strong>King crabs</strong> (<em>Paralithodes camtschatica</em> (Tilesius))</td>
  </tr>
  <tr>
    <td class="tdc">9.4</td>
    <td class="tdr">55,408</td>
    <td>April-June</td>
    <td class="tdc">?</td>
    <td>Egg</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">100-200<a name="FNanchor_63_63" id="FNanchor_63_63"></a><a href="#Footnote_63_63" class="fnanchor">[63]</a></td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">?</td>
    <td rowspan="5">Kurata 1960, 1964; Korolev 1964; Rodin 1970</td>
  </tr>
  <tr>
    <td class="tdc">17.1</td>
    <td class="tdr">444,651</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td>Zoeal</td>
    <td rowspan="2" class="tdc">0.55-0.65</td>
    <td rowspan="2" class="tdc">?</td>
    <td rowspan="2">April-July</td>
    <td class="tdc">33 at 7-10&deg;C</td>
  </tr>
  <tr>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td>Zoeal</td>
    <td class="tdc">23 at 12.3-12.5&deg;C</td>
  </tr>
  <tr>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td>Glaucothoeal </td>
    <td class="tdc">0.38x0.18</td>
    <td class="tdc">?</td>
    <td>May-?</td>
    <td class="tdc">?</td>
  </tr>
  <tr>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td>Juvenile</td>
    <td class="tdc">?</td>
    <td class="tdc">1-?</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">?</td>
  </tr>
  <tr>
    <td colspan="0"><strong>Snow crabs</strong> (<em>Chionoecetes bairdi</em> Rathbun)</td>
  </tr>
  <tr>
    <td class="tdc">?</td>
    <td class="tdc">?</td>
    <td class="tdc">?<a name="FNanchor_65_65" id="FNanchor_65_65"></a><a href="#Footnote_65_65" class="fnanchor">[65]</a></td>
    <td class="tdc">?</td>
    <td>Egg</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">100<a href="#Footnote_63_63" class="fnanchor">[63]</a></td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">?</td>
    <td rowspan="8">Haynes 1973<a href="#Footnote_55_55"
    class="fnanchor">[55]</a> Jewett and Haight<a
    name="FNanchor_64_64" id="FNanchor_64_64"></a><a
    href="#Footnote_64_64" class="fnanchor">[64]</a></td>
  </tr>
  <tr>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td>Prezoeal</td>
    <td class="tdc">0.22-0.28</td>
    <td class="tdc">?</td>
    <td>May-?</td>
    <td class="tdc">1-2 at 2.5&deg;C</td>
  </tr>
  <tr>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td>1st zoeal</td>
    <td class="tdc">0.50-0.56</td>
    <td class="tdc">?</td>
    <td>Summer</td>
    <td class="tdc">?</td>
  </tr>
  <tr>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td>2d zoeal</td>
    <td class="tdc">?</td>
    <td class="tdc">0-10</td>
    <td>Summer</td>
    <td class="tdc">?</td>
  </tr>
  <tr>
    <td rowspan="2" class="tdc">&mdash;</td>
    <td rowspan="2" class="tdc">&mdash;</td>
    <td rowspan="2" class="tdc">&mdash;</td>
    <td rowspan="2" class="tdc">&mdash;</td>
    <td rowspan="2">Megalopal</td>
    <td class="tdc">0.30-0.35x</td>
    <td rowspan="2" class="tdc">?</td>
    <td rowspan="2">Summer</td>
    <td rowspan="2" class="tdc">&mdash;</td>
  </tr>
  <tr>
    <td class="tdc">0.18-0.21</td>
  </tr>
  <tr>
    <td rowspan="2" class="tdc">&mdash;</td>
    <td rowspan="2" class="tdc">&mdash;</td>
    <td rowspan="2" class="tdc">&mdash;</td>
    <td rowspan="2" class="tdc">&mdash;</td>
    <td rowspan="2">Juvenile</td>
    <td class="tdc">0.44-0.48x</td>
    <td rowspan="2" class="tdc">?</td>
    <td rowspan="2" class="tdc">&mdash;</td>
    <td rowspan="2" class="tdc">&mdash;</td>
  </tr>
  <tr>
    <td class="tdc">0.32-0.35<span class="pagenum"><a name="Page_208" id="Page_208">[Pg 208]</a></span></td>
  </tr>
  <tr>
    <td colspan="4"><strong>Snow crabs</strong> (<em>Chionoecetesopilio</em> (O. Fabricius))</td>
    <td>Egg</td>
    <td class="tdc">?</td>
    <td class="tdc">93<a href="#Footnote_60_60" class="fnanchor">[60]</a></td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">?</td>
    <td rowspan="0">Ito 1968; Kon 1970; Haynes 1973; Motoh 1973; Jewett and Haight<a href="#Footnote_64_64" class="fnanchor">[64]</a></td>
  </tr>
  <tr>
    <td class="tdc">?</td>
    <td class="tdc">?</td>
    <td class="tdc">?<a href="#Footnote_65_65" class="fnanchor">[65]</a></td>
    <td class="tdc">?</td>
    <td>Prezoeal</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">?</td>
    <td>May-?</td>
    <td rowspan="5" class="tdc">63-66 at 11-13&deg;C</td>
  </tr>
  <tr>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td>1st zoeal </td>
    <td class="tdc">0.48-0.54</td>
    <td class="tdc">?</td>
    <td>Summer</td>
  </tr>
  <tr>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td>2d zoeal</td>
    <td class="tdc">0.62-0.71</td>
    <td class="tdc">?</td>
    <td>Summer </td>
  </tr>
  <tr>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td>Megalopal </td>
    <td class="tdc">0.29-0.33</td>
    <td class="tdc">?</td>
    <td>Summer </td>
  </tr>
  <tr>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td></td>
    <td class="tdc">0.19</td>
    <td colspan="0"></td>
  </tr>
  <tr>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td>Juvenile</td>
    <td class="tdc">4.4-4.8x</td>
    <td class="tdc">?</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
  </tr>
  <tr>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td></td>
    <td class="tdc">3.2-3.5</td>
    <td colspan="0"></td>
  </tr>
</tbody>
</table></div>

<div class="pagenum"><a name="Page_209" id="Page_209">[Pg 209]</a></div>

<p>The commercially important king and snow
crabs of the eastern Bering Sea also have larval
stages that are pelagic (Table 3). Zoeae
and megalopa of snow crabs are found near
the surface where they are vulnerable to
plankton-feeding marine birds. The eggs of
king crabs are attached to the abdomen of the
female, but after hatching, the larvae become
pelagic and occur near the surface. They are
planktonic through five larval stages before
settling to the bottom to take up demersal
residence (Kurata 1960, 1964). These larvae
attain a length of 5.5-6.5 mm and spend 33
days or more in the plankton (Kurata 1960).
Even after the young king crabs have settled
to the bottom, they may still frequent water
shallow enough to make them vulnerable to
predation by some marine birds. Juvenile king
crabs 1 and 2 years of age appear to prefer
shallower water than do older crabs. In southeastern
Alaska, during the spring, small juvenile
crabs have been observed in pods at
depths as little as 1 m below the low tide level.</p>

<p>The available life stages of king and snow
crabs and commercially important demersal
fish (Table 3) represent an enormous food
supply for other fishes and marine birds. Predation
by marine birds on pelagic eggs and on
the larval and juvenile stages of demersal fish
is not well documented, probably because the
rapid digestion rate of birds makes species
identification of these stages difficult. Investigators
must often depend on the presence of
the hard parts of fish (such as scales and otoliths)
in the stomachs of birds to identify the
species eaten. Because these hard parts have
not yet formed in the larvae and most juveniles,
predation by marine birds on older fish
is more apparent on examination of stomach
contents. Full understanding of predation by
marine birds on demersal fish and shellfish requires
additional data on when and where the
egg, larval, and juvenile stages are present.</p>


<h5><em>Pelagic Fish</em></h5>

<p>Many fish, such as herring, capelin, smelt,
and salmon, are pelagic for part of their lives,
particularly during the spring and summer
feeding periods. The extent of predation by
marine birds on these species depends primarily
on the location of their spawning
grounds, their growth rates, and the size of
the adults. The spawning location determines
the extent of predation on eggs, whereas
growth rate and adult size determine during
how much of its lifetime a given fish species is
vulnerable to the wide variety of marine birds.</p>

<p>Herring spawn in intertidal and subtidal
zones and spend most of their post-larval lives
in bays or estuaries near the coast. They deposit
their adhesive eggs primarily on vegetation,
and the eggs are particularly vulnerable
to predation by a wide variety of marine and
terrestrial birds. Outram (1958) estimated
that gulls alone accounted for 39% of the egg
loss on the spawning grounds at Vancouver
Island, British Columbia. When herring larvae
hatch, they are between 0.7 and 0.8 cm
long; when they metamorphose about 6-8
weeks later, they are between 2.6 and 3.5 cm
long. Thereafter, juvenile herring grow
rapidly and reach a length of about 7-10 cm
before winter. Although herring as old as 13
years and up to 38 cm long have been reported
in Alaska, they seldom exceed 30 cm and 11
years of age (Rounsefell 1929). During spring
and summer, herring are commonly within
10 m of the surface, but in winter, they are in
water 100-140 m deep. Although herring are
particularly vulnerable to predation in spring
and summer, they are available to marine
birds during most of their life.</p>

<p>The life history of capelin is somewhat different
than that of herring&mdash;they live in the
open sea near the surface and throughout the
water column most of their lives. Sometime in
June or early July, they migrate in large
schools toward shore to spawn (Musienko
1970). In British Columbia, capelin bury their
eggs in coarse sand and gravel in the intertidal
and subtidal zones. The larvae are 0.5-0.7
cm long at hatching and are carried by
currents to the open sea where they develop in
the plankton. Capelin attain an age of 5 years
and a maximum length of about 22 cm; their
small size makes them vulnerable to predation
by marine birds most of their lives, and
they are an important pelagic food fish for
other commercial fish in the Bering Sea.</p>

<p>The sand lance reaches a maximum size of
20-26 cm and is vulnerable to bird predation
during most of its life. Little information is
available on the maximum age attained by<span class="pagenum"><a name="Page_210" id="Page_210">[Pg 210]</a></span>
this species in the Bering Sea, but because of
its size, it is an important forage fish for many
commercial fish species.</p>

<p>The five species of Pacific salmon of the
eastern Bering Sea spawn in fresh water, unlike
herring, capelin, and sand lance. Their
eggs are not vulnerable to extensive predation
by marine birds; gulls take mainly salmon
eggs which have been dislodged from the
gravel and are drifting or being rolled along
the stream bottom by the current (Moyle
1966). After a few months to several years in
fresh water, the juvenile salmon (5-14 cm
long) enter the Bering Sea during late spring
or early summer and migrate through these
waters to feeding grounds, primarily in the
north Pacific Ocean. At maturity, the survivors
return to their home streams and
rivers to spawn. It is during the seaward migratory
phase of their life cycle that salmon
are most vulnerable to predation by marine
birds.</p>

<p>The sockeye salmon <em>(Oncorhynchus nerka)</em>
is the most abundant and valuable species
harvested by American fishermen in the
waters adjacent to the Bering Sea and, as a result,
the one that has been most extensively
studied during early marine life. Juvenile
sockeye salmon are between 8 and 14 cm long
when they enter the Bering Sea between late
May and early July. They are most abundant
in the upper 1 m of water at night and the
upper 2 m during the day (Straty 1974)&mdash;well
within the regime that can be exploited by
many species of marine birds.</p>

<p>The numbers of juvenile sockeye salmon migrating
seaward from the Bristol Bay region
of the Bering Sea in a single year has ranged
between 46.3 and 370.4 million (H. Jaenicke,
personal communication). This is equivalent
to between 409 and 3,267 metric tons (on the
basis of the mean weight of the juveniles when
they enter the Bering Sea). These large numbers
of juvenile sockeye salmon, plus juvenile
chinook salmon <em>(O. tshawytscha)</em>, coho salmon
<em>(O. kisutch)</em>, chum salmon <em>(O. keta)</em>, and
pink salmon <em>(O. gorbuscha)</em> from all other
rivers entering the Bering Sea, represent a
considerable input of energy from fresh water
in the form of prime forage fish for other
fishes, marine birds, and mammals. Young
salmon enter the Bering Sea each year over a
period of only 6 to 8 weeks and may follow
rather discrete coastal migration routes
through the Bering Sea (Fig. 6), with the result
that predators have access to an abundant
but transient food supply.</p>

<div class="figcenter" style="width: 700px;">
<img src="images/p_210.jpg" width="700" height="493" alt="" />
<div class="caption"><p>Fig. 6. Distribution of juvenile sockeye salmon in
Bristol Bay and the eastern Bering Sea (adapted
from Straty 1974).</p></div>
</div>

<p>The only published account of predation by
marine birds on juvenile salmon in the Bering
Sea is that of Ogi and Tsujita (1973). They
found juvenile sockeye salmon in the
stomachs of murres captured in gill nets in
the eastern Bering Sea. The predation did not
appear extensive, but most of the birds were
captured outside or on the fringes of the main
seaward migration route of the salmon. The
foods of marine birds should be studied in conjunction
with studies of the migrations of
juvenile salmon.</p>


<h4>Influence of Growth Rate
and Adult Size of Fish on
the Extent of Predation</h4>

<p>Incubation time for fish eggs, the length of
the pelagic larval period (Table 3), and the
growth rate of juvenile fish are species-specific
and temperature-dependent. The extent
to which a fish species is subjected to predation
by marine birds is directly related to
the rate at which development and growth
occur. For example, the less time it takes the
pelagic eggs of demersal fish and shellfish to
hatch and complete pelagic larval life, the less
is the time they will be preyed on by marine
birds. For fish species that are pelagic during
their entire life, the rate of growth will deter<span class="pagenum"><a name="Page_211" id="Page_211">[Pg 211]</a></span>mine
how long they remain small enough for
birds to eat. Some of the smaller pelagic fish,
such as herring, capelin, and smelt, are vulnerable
to bird predation most of their lives;
larger pelagic species like salmon may be
preyed on for only a very short time. The
maximum size fish that can be eaten by
marine birds is, therefore, important in
evaluating predation on a given species of
fish.</p>

<p>The literature on the food habits of marine
birds contains little on the sizes of fish consumed.
Tuck (1960) stated that murres probably
will take fish up to 18 cm long. Ogi and
Tsujita (1973) estimated the lengths of Pacific
pollock in the stomachs of murres taken in the
eastern Bering Sea at 24 cm.</p>

<p>Herring in the eastern Bering Sea reach an
age of 11 years and grow to about 33 cm.
Herring could, therefore, be taken during
most of their lives by murres but during only
the first few years by smaller birds such as
fulmars and shearwaters. Capelin and some
species of smelt would be vulnerable to birds
during all their lives. Although the size of
adult Pacific salmon varies with the species,
they are all so large that they are not preyed
upon by marine birds. Once in the ocean, juvenile
salmon grow at such a rapid rate that
they are probably not very vulnerable to
marine birds after their first 4 to 6 months at
sea. Limited studies on the growth of juvenile
sockeye salmon in the eastern Bering Sea
(Straty 1974) indicate they may double their
size in their first 8 weeks at sea. A sockeye salmon
that entered the Bering Sea at 12 cm in
mid-June would be 24 cm long in August&mdash;the
maximum size that a murre could eat; the fish
could be eaten by smaller marine birds for
much less time. Pink and chum salmon enter
the sea at a smaller size than sockeye salmon
and would be vulnerable to predation both by
a greater variety of marine birds and for a
longer period of time.</p>


<h4>Competition Between
Commercial Fish and
Marine Birds</h4>

<p>We do not know the importance of competition
between marine birds and commercial
fish in the eastern Bering Sea. Only a few investigators
have even alluded to competition
between marine birds and fish for food. Ogi
and Tsujita (1973) mentioned that competition
seemed to exist between murres and juvenile
sockeye salmon for euphausiids in the
eastern Bering Sea. We have listed some of
the types of forage fish and invertebrates
eaten by commercial fish (Table 4) and marine
birds (Table 5) in the eastern Bering Sea; comparison
of these two tables clearly indicates
that competition could occur.</p>

<p>The principal factors determining the extent
of competition between marine birds and
fish are the numbers of birds and fish, the
length of time that various life history stages
of the fish are in association with the birds,
and the abundance of the preferred foods at
these times. The impact of competition depends
on the adaptability of the birds and fish
to alternative types of food.</p>

<p>The types and sizes of food eaten by fish
vary with the life history stage&mdash;especially
with size at each stage. For instance, very
young herring eat the eggs and nauplii of
copepods or small copepodite stages and barnacles.
As herring grow, their diet includes
small fish and larger zooplankton, such as
mature copepods, amphipods, euphausiids,
and pteropods. Pacific cod shorter than 9 cm
feed on small crustaceans (Moiseev 1953),
whereas larger cod eat young crabs, shrimp,
and fish. Small juvenile sockeye salmon feed
mainly on larval stages of euphausiids (Straty
1974), but larger juveniles also eat the more
adult forms, which eventually make up a significant
part of their diet (Nishiyama 1974).</p>

<p>The change in the diet of fishes with growth
results in competition with a changing variety
of marine birds. For example, deep-diving
birds may replace surface feeders as the major
bird competitors of the Pacific cod and pollock
as these fish increase in size and seek
deeper waters. The diet of cod changes from
small crustaceans in shallow water to progressively
larger food that eventually includes
herring, sand lance, shrimp, and crabs. The
change to herring and sand lance, and quite
possibly small crabs, places the adult cod in
competition with both the surface feeders and
pursuit diving birds, but adult cod do not
compete with birds for zooplankton.</p>

<p><span class="pagenum"><a name="Page_212" id="Page_212">[Pg 212]</a></span></p>

<div class="center">
<table border="0" cellpadding="4" cellspacing="0" summary="Table 4.
Food items eaten by the adult stage of seven commercially important
species of fish in the eastern Bering Sea.">
    <caption>Table 4. <em>Food items eaten by the adult stage of seven
    commercially important species of fish in the eastern Bering
    Sea.</em></caption>
  <tr>
    <th>Food item</th>
    <th>Herring</th>
    <th>Salmon</th>
    <th>Walleye pollock</th>
    <th>Pacific cod</th>
    <th>Pacific ocean perch</th>
    <th>Yellowfin sole</th>
    <th>Pacific halibut</th>
  </tr>
  <tr>
    <td colspan="0">Invertebrates</td>
  </tr>
  <tr>
    <td class="tdp">Pteropods</td>
    <td class="tdc">X</td>
    <td class="tdc">X</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">X</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
  </tr>
  <tr>
    <td class="tdp">Squid</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">X</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">X</td>
    <td class="tdc">X</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">X</td>
  </tr>
  <tr>
    <td class="tdp">Polychaetes</td>
    <td class="tdc">X</td>
    <td class="tdc">X</td>
    <td class="tdc">X</td>
    <td class="tdc">X</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">X</td>
    <td class="tdc">X</td>
  </tr>
  <tr>
    <td class="tdp">Copepods</td>
    <td class="tdc">X</td>
    <td class="tdc">X</td>
    <td class="tdc">X</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
  </tr>
  <tr>
    <td class="tdp">Amphipods</td>
    <td class="tdc">X</td>
    <td class="tdc">X</td>
    <td class="tdc">X</td>
    <td class="tdc">X</td>
    <td class="tdc">X</td>
    <td class="tdc">X</td>
    <td class="tdc">&mdash;</td>
  </tr>
  <tr>
    <td class="tdp">Euphausiids</td>
    <td class="tdc">X</td>
    <td class="tdc">X</td>
    <td class="tdc">X</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">X</td>
    <td class="tdc">X</td>
    <td class="tdc">&mdash;</td>
  </tr>
  <tr>
    <td class="tdp">Decapods</td>
    <td class="tdc">X</td>
    <td class="tdc">X</td>
    <td class="tdc">X</td>
    <td class="tdc">X</td>
    <td class="tdc">X</td>
    <td class="tdc">X</td>
    <td class="tdc">X</td>
  </tr>
  <tr>
    <td colspan="0">Fish</td>
  </tr>
  <tr>
    <td class="tdp">Capelin</td>
    <td class="tdc">X</td>
    <td class="tdc">X</td>
    <td class="tdc">X</td>
    <td class="tdc">X</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">X</td>
    <td class="tdc">&mdash;</td>
  </tr>
  <tr>
    <td class="tdp">Sand lance</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">X</td>
    <td class="tdc">X</td>
    <td class="tdc">X</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">X</td>
  </tr>
</table></div>

<div class="center">
<table border="0" cellpadding="4" cellspacing="0" summary="Table 5.
Forage fish and invertebrate foods eaten by seven species of marine
birds in the eastern Bering Sea.">
    <caption>Table 5. <em>Forage fish and invertebrate foods eaten by
    seven species of marine birds in the eastern Bering Sea.</em></caption>
  <tr>
    <th>Food item</th>
    <th>Shearwaters</th>
    <th>Murres</th>
    <th>Puffins</th>
    <th>Murrelets</th>
    <th>Fulmars</th>
    <th>Kittiwakes</th>
    <th>Gulls</th>
  </tr>
  <tr>
    <td colspan="0">Forage fish</td>
  </tr>
  <tr>
    <td class="tdp">Sand lance</td>
    <td class="tdc">X</td>
    <td class="tdc">X</td>
    <td class="tdc">X</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">X</td>
    <td class="tdc">X</td>
  </tr>
  <tr>
    <td class="tdp">Capelin</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">X</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
  </tr>
  <tr>
    <td colspan="0">Invertebrates</td>
  </tr>
  <tr>
    <td class="tdp">Copepods</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">X</td>
    <td class="tdc">&mdash;</td>
  </tr>
  <tr>
    <td class="tdp">Euphausiids</td>
    <td class="tdc">X</td>
    <td class="tdc">X</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">X</td>
    <td class="tdc">&mdash;</td>
  </tr>
  <tr>
    <td class="tdp">Amphipods</td>
    <td class="tdc">X</td>
    <td class="tdc">X</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">X</td>
    <td class="tdc">&mdash;</td>
  </tr>
  <tr>
    <td class="tdp">Decapods</td>
    <td class="tdc">X</td>
    <td class="tdc">X</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">X</td>
    <td class="tdc">&mdash;</td>
  </tr>
  <tr>
    <td class="tdp">Pteropods</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">X</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
  </tr>
  <tr>
    <td class="tdp">Chaetognaths</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
  </tr>
  <tr>
    <td class="tdp">Polychaetes</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">X</td>
    <td class="tdc">X</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">X</td>
    <td class="tdc">&mdash;</td>
  </tr>
  <tr>
    <td class="tdp">Squid</td>
    <td class="tdc">X</td>
    <td class="tdc">X</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">X</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
  </tr>
</table></div>

<p>As pollock increase in size, they continue to
feed mainly on zooplankton, but they change
from copepods near the surface to euphausiids
at mid-depths and near the bottom. Euphausiids
are large and abundant zooplankters
which, for the most part, are available only to
deep-diving birds. Adult pollock also consume
herring, sand lance, capelin, and other small
fish.</p>

<p>Both marine birds and fish are capable of
exploiting a wide variety of food, and often
their stomach contents reflect the relative
abundance of food items in the area. Ogi and
Tsujita (1973) illustrated the differences in
the food taken by murres captured at different
locations in the eastern Bering Sea.
Carlson (1977) and Ogi and Tsujita (1973) reported
on differences in the diet of juvenile
sockeye salmon captured at various locations
in Bristol Bay and the eastern Bering Sea.
The diets of many species of birds and fish,
however, seem to be largely determined by
their physiological and morphological adaptations
and resultant feeding behavior. For instance,
adult sockeye and pink salmon have
well-developed gill rakers and feed largely on
zooplankton, whereas chinook and coho salmon
have poorly developed gill rakers and
feed almost entirely on fish. In the eastern
Bering Sea, murres appear to prefer the
Pacific sand lance, whereas the slender-billed
shearwater consumes mainly euphausiids (Ogi
and Tsujita 1973). Thus, murres may be
greater competitors with piscivorous fish
than are shearwaters. Shearwaters are probably
more important as competitors with zoo<span class="pagenum"><a name="Page_213" id="Page_213">[Pg 213]</a></span>plankton-eating
fish that inhabit shallow
water in juvenile stages and with pelagic fish
species (such as pollock, herring, salmon, and
capelin) that are heavily dependent on
euphausiids.</p>

<p>Some species of marine birds may interact
with fish as predators and competitors. As an
example, pursuit diving birds, such as murres
and puffins, may be important predators on
juvenile salmon in the eastern Bering Sea, but
these same birds may compete for food with
adult salmon. Surface-feeding birds, such as
fulmars, shearwaters, kittiwakes, and gulls,
may be important as both predators and competitors
with herring and capelin and some
demersal fish.</p>


<h4>Dependency of Marine Birds
on Commercial Fish</h4>

<p>The interactions of commercial fish and
marine birds of the Bering Sea can be determined
only if we know their distribution,
abundance, and food habits, especially while
they are associated with one another. Information
is particularly lacking for all life history
stages of commercial fish species and the
seasonal movements of birds. We have some
knowledge of the distribution and abundance
of the various life history stages and the food
habits of commercial fish in the Bering Sea.
Little is known of the abundance, seasonal
movements, and food habits of marine birds
in this region, however, probably because
marine birds have had little direct commercial
value in the northern hemisphere. Food
studies on marine birds are particularly difficult
because their rapid digestion soon destroys
the identity of the food.</p>

<p>We can make a reasonable guess as to some
bird-fish associations for two regions of the
Bering Sea where we have information on the
distribution of marine birds and the various
life history stages of commercial fish. For
example, piscivorous birds, such as murres,
puffins, black-legged kittiwakes, and slender-billed
shearwaters, are extremely abundant in
the summer along the seaward migration
route of juvenile sockeye salmon (Fig. 7); the
juvenile salmon, kittiwakes, and shearwaters
all feed on plankton. Shuntov (1961) showed
that kittiwakes are most abundant along the
edge of the continental shelf in the Bering Sea
in the summertime. This distribution coincides
with the distribution of the eggs and larvae
of pollock, certain flatfish, rockfish, sablefish,
and several other species. These birds
both exploit the fish directly (predation) and
compete with them for plankton. Not enough
information is available on the food habits of
birds at the time fish eggs and larvae are
present to evaluate this interaction.</p>


<h4>Environmental Influence on
Predation and Competition
Between Marine Birds
and Commercial Fish</h4>

<p>Because fish are cold-blooded animals, temperature,
through its influence on the rate of
metabolism, is a major variable in determining
the amount of energy needed for maintenance
and for performing such essential activities
as swimming and feeding&mdash;fish are
less active, feed less, and grow more slowly in
cold waters. For example, growth in young
sockeye salmon is very slow at temperatures
lower than 4&deg;C (Donaldson and Foster 1941),
and temperature profoundly affects their
swimming speed (Brett et al. 1958). The rates
of development of the eggs of some flatfish
are closely correlated with water temperature
(Ketchen 1956)&mdash;flatfish developed more
rapidly at higher temperatures (Fig. 8). At
lower temperatures, the rate of growth is also
slower and, therefore, the duration of pelagic
larval life is longer for demersal fish and shellfish.</p>

<p>Variations in sea temperature should, therefore,
influence the extent to which fish are vulnerable
to predation and competition. For
example, eggs would take a longer time to
hatch in colder than in warmer sea water. In
both pelagic fish such as herring, whose eggs
are laid in the intertidal zone, and in demersal
fish with pelagic eggs such as the sole, the
period of vulnerability of eggs to bird predation
would be extended. At lower temperatures
the length of the pelagic life of demersal
fish and shellfish and their vulnerability to
predation would also be greater than at higher
temperatures. For example, the number of<span class="pagenum"><a name="Page_214" id="Page_214">[Pg 214]</a></span>
days between molts of the zoeal stages of
snow crabs is temperature-dependent&mdash;the
warmer the water, the less the time between
molts (Kon 1970).</p>

<div class="figcenter" style="width: 657px;">
<img src="images/p_214.jpg" width="657" height="700" alt="" />
<div class="caption"><p>Fig. 7. Distribution and numbers of birds observed in Bristol Bay along seaward migration route of sockeye
salmon (from Bartonek and Gibson 1972).</p></div>
</div>

<p>Temperature, through its effects on swimming
speed, feeding activity, and growth of
juvenile fish, might influence the magnitude
of predation by birds on pelagic fish in the following
ways: (1) lower sea temperatures
would increase the vulnerability of juvenile
fish to bird predation because swimming
speed would decrease, and the time the fish
are of a size that could be eaten by would-be
predators would increase; (2) lower sea temperatures
would reduce the feeding by fish
and decrease the competition by fish for food
exploited by birds; and (3) higher sea temperatures
would have the opposite effect&mdash;the
feeding by fish would increase consumption of
the foods that birds feed on.</p>

<p>In the eastern Bering Sea, water temperatures
may vary greatly between years for the
same month (Fig. 9). Such variation should result
in variation in the temperature-de<span class="pagenum"><a name="Page_215" id="Page_215">[Pg 215]</a></span>pendent
activities of fish and, in turn, in magnitude
of marine bird predation and competition.</p>

<div class="figcenter" style="width: 556px;">
<img src="images/p_215.jpg" width="556" height="700" alt="" />
<div class="caption"><p>Fig. 8. The relation of temperature to the rate of development
to hatching of lemon sole, as compared
with two European flatfishes (Ketchen 1956).</p></div>
</div>


<h4>Possible Influences of Man
on the Interaction of Marine
Birds with Commercial Fish</h4>

<p>We have noted that the abundance and age
and size composition of major stocks of fish in
the Bering Sea have been drastically reduced
by commercial fishing. This has resulted in
the reduction in numbers of fish at all life history
stages, including those on which marine
birds and other fishes depend for food. What
effect this reduction has had on the abundance
and distribution of marine birds in the
Bering Sea is unknown. It depends in part on
the ability of birds to eat other fish or increase
their use of zooplankton or nekton.</p>

<p>We can hypothesize on probable changes in
bird and fish abundance that resulted from
the heavy commercial harvest of fish but any
such changes cannot be documented or quantified.
A reduction in stocks of a fish species
could result in a reduced supply of food for a
species of bird and cause a shift in the diet of
this bird to other species of fish or to more
zooplankton. For a bird species with specific
food preferences, this could mean a reduction
in its abundance to a level supportable by the
available food supply. For bird species with
less specific food requirements, a reduction in
a species of fish could mean a reduction in
competition for food with that fish&mdash;which
could increase survival of the birds.</p>

<p>Man's intentional harvest of marine birds,
such as the shearwater in parts of the southern
hemisphere, and his inadvertent harvest
of other bird species which are entangled or
caught in fishing gear reduce predation and
competition by marine birds. This, in turn,
may aid the survival of the fish stocks in the
Bering Sea.</p>

<p>The status of most stocks of commercial
fish and shellfish in the Bering Sea is such
that reductions in harvest are warranted,
have been proposed, or are in effect. If the
200-mile (61-km) limit of jurisdiction over the
marine resources by adjacent coastal States is
implemented, either as a result of the Law of
the Sea Conferences or unilaterally by the
United States, we can expect commercial fishing
in the eastern Bering Sea to be more
tightly regulated. Such action should result in
a reduction in harvest of those fish species
now in a depleted condition, which, in turn,
could influence the abundance of marine
birds. Now is an opportune time to implement
the studies required to increase our knowledge
of the abundance, distribution, and seasonal
movements of marine birds and their
relationship to commercial fish resources of
the eastern Bering Sea.</p>


<h4>Conclusions</h4>

<p>&#8226; The eastern Bering Sea is a region of high
biological productivity; it is one of the world's
great producers of commercial fish and major
congregating areas for marine birds.</p>

<p>&#8226; The vulnerability of fish to predation by
marine birds depends on life history features,
such as place of spawning, duration of larval
stages, growth rate, sea temperature, and
adult size of fish, and on the distribution,
feeding behavior, and food habits of marine
birds.</p>

<p><span class="pagenum"><a name="Page_216" id="Page_216">[Pg 216]</a></span></p>

<div class="figcenter" style="width: 536px;">
<img src="images/p_216.jpg" width="536" height="700" alt="" />
<div class="caption"><p><strong>Fig. 9.</strong> Sea temperatures in Bristol Bay and southeastern Bering Sea in mid-June and early July of 1967
and 1971 (from Straty 1974).</p></div>
</div>

<p><span class="pagenum"><a name="Page_217" id="Page_217">[Pg 217]</a></span></p>

<p>&#8226; The most apparent predation by marine
birds on fish is on fish large or mature enough
that some hard body parts persist and can be
found in the stomach samples of birds.</p>

<p>&#8226; Little is known of the extent of bird predation
on the pelagic eggs and larvae of
demersal fish and shellfish in the Bering Sea
because of lack of investigation and the rapid
digestion of eggs and larvae by birds.</p>

<p>&#8226; Predation by marine birds on juvenile
salmon is not well documented because of the
lack of investigation in areas where both birds
and fish are present.</p>

<p>&#8226; Marine birds and commercial fish eat similar
zooplankton and fish in the eastern Bering
Sea. The food exploited by both generally reflects
the relative abundance of the types of
food in the area, but food preference is displayed
by some species of fish and birds.</p>

<p>&#8226; More is known about the food habits of the
commercial fish than of the marine birds of
the Bering Sea.</p>

<p>&#8226; Sea water temperature may be a major environmental
factor in the Bering Sea since it
influences both the extent to which fish are
vulnerable to predation and the amount of
competition with marine birds. Sea temperatures
may vary greatly from year to year in
the Bering Sea, and this may result in variations
in the magnitude of predation and competition
between birds and fish.</p>

<p>&#8226; The distribution of marine birds and the
various stages in the life history of commercial
fish are not well known for the eastern
Bering Sea. Where these have been studied,
they are intimately related. Such knowledge is
required to gain some insight into even the
potential for predation and competition in the
dynamics of the marine bird and commercial
fish populations of this region. In two instances,
it is known that the occurrence of
marine birds and the early life history stages
of fish coincide so as to result in both potential
predation on the fish by the birds and
competition for food between the fish and the
birds.</p>

<p>&#8226; The possibility exists that the commercial
fish resources of the eastern Bering Sea will
eventually come under the jurisdiction of the
United States. This could mean reduced harvests
of fish to restore depleted stocks. Such
action could result in changes in the abundance
of the marine birds of this region by
creating an increased food supply for some
and decreased supply for others.</p>


<h4>Acknowledgments</h4>

<p>We thank J. C. Bartonek and H. R. Carlson,
H. Jaenicke, H. Larkins, and B. L. Wing for
supplying various materials presented in this
paper.</p>


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from the eastern Bering Sea and Bristol Bay,
June-August 1970 and 1971. Jpn. J. Ecol.
23(5):201-209.</p>

<p>Outram, D. N. 1958. The magnitude of herring
spawn losses due to bird predation on the west
coast of Vancouver Island. Fish. Res. Board Can.,
Prog. Rep. Pac. Coast Stn. 111:9-13.</p>

<p>Paraketsov, I. A. 1963. On the biology of <em>Sebastodes
alutus</em> of the Bering Sea. Part 1, pages 319-327
<em>in</em> Soviet fisheries investigations in the
northeast Pacific Ocean. (Transl. from Russian.)
Israel Program for Scientific Translations,
Jerusalem.</p>

<p>Pearson, T. H. 1968. The feeding biology of sea-bird
species breeding on the Farne Islands, Northumberland.
J. Anim. Ecol. 37:521-552.</p>

<p><span class="pagenum"><a name="Page_219" id="Page_219">[Pg 219]</a></span></p>

<p>Pertseva-Ostraumova, T. A. 1954. Material on the
development of the far-eastern flatfish. 1. Development
of the yellowfin sole. Tr. Inst.
Okeanol. Akad. Nauk., SSSR-11.</p>

<p>Reid, G. J. 1972. Alaska's fishery resources, the
Pacific herring. U.S. Natl. Mar. Fish. Serv., Fish.
Facts 2, 20 pp.</p>

<p>Rodin, V. E. 1970. An estimation of the state of the
king crab (<em>Paralithodes camtschatica</em> Tilesius)
stock in the southeastern Bering Sea. Part 5,
pages 149-156 <em>in</em> Soviet fisheries investigations
in the northeast Pacific Ocean. (Transl. from Russian.)
Israel Program for Scientific Translations,
Jerusalem.</p>

<p>Rounsefell, G. A. 1929. Contribution to the biology
of the Pacific herring, <em>Clupea pallasii</em>, and the
condition of the fishery in Alaska. Bull. U.S. Bur.
Fish. 45:227-320.</p>

<p>Rumyantsev, A. I., and M. A. Darda. 1970. Summer
herring in the eastern Bering Sea. Part 5, pages
409-441 <em>in</em> Soviet fisheries investigations in the
northeastern Pacific Ocean. (Transl. from Russian.)
Israel Program for Scientific Translations,
Jerusalem.</p>

<p>Sanger, G. A. 1972. Preliminary standing stock and
biomass estimates of seabirds in the subarctic
Pacific region. Pages 499-611 <em>in</em> A. Y. Takenouti,
ed. Biological oceanography of the northern
north Pacific Ocean.</p>

<p>Serobaba, I. I. 1968. On the spawning of Alaska pollock
<em>Theragra chalcogramma</em> (Pallas) in the
northeastern part of the Bering Sea. Vopr.
Ikhtiol. 6(53):992-1003. (Transl. from Russian.)
Israel Program for Scientific Translations,
Jerusalem.</p>

<p>Shaefer, M. B. 1970. Men, birds, and anchovies in
the Peru current&mdash;dynamic interactions. Trans.
Am. Fish. Soc. 99(3):461-467.</p>

<p>Shuntov, V. P. 1961. Migration and distribution of
marine birds in the southeastern Bering Sea during
spring-summer season. Zool. Zh. 40(7):1058-1069.
(In Russian, summary in English.)</p>

<p>Shuntov, V. P. 1966. Concerning wintering of birds
in the far eastern seas and in the northern part of
the Pacific Ocean. Zool. Zh. 45(11):698-711.
(Transl. from Russian.) Can. Wildl. Serv.</p>

<p>Solomensen, F. 1965. The geographical variation of
the fulmar <em>(Fulmarus glacealis)</em> and the zones of
marine environment in the North Atlantic. Auk
82:327-355.</p>

<p>Stevenson, J. C. 1962. Distribution and survival of
herring larvae (<em>Clupea pallasi</em> Valenciennes) in
British Columbia waters. J. Fish. Res. Board
Can. 19(5):735-810.</p>

<p>Straty, R. R. 1974. Ecology and behavior of juvenile
sockeye salmon <em>(Oncorhynchus nerka)</em> in
Bristol Bay and the eastern Bering Sea. Inst.
Mar. Sci., Univ. Alaska, Occas. Publ. 2:285-320.</p>

<p>Tanino, Y., H. Tsujisaki, K. Nakamichi, and K.
Kyushin. 1959. On the maturity of Alaska pollock,
<em>Theragra chalcogramma</em> (Pallas). Bull. Hokkaido
Reg. Fish. Res. Lab. 20:145-164.</p>

<p>Taylor, F. H. C. 1967. The relationship of midwater
trawl catches to sound scattering layers off the
coast of northern British Columbia. J. Fish. Res.
Board Can. 25(3):457-472.</p>

<p>Tuck, L. M. 1960. The murres: their distribution,
populations, and biology, a study of the genus
<em>Uria</em>. Can. Wildl. Ser. 1. 260 pp.</p>

<p>Wiens, J. A., and J. M. Scott. 1976. Model estimation
of energy flow in Oregon coastal seabird
populations. Condor 77(4):439-452.</p>

<p>Yusa, T. 1954. On the normal development of the
fish, <em>Theragra chalcogramma</em> (Pallas), Alaska pollock.
Bull. Hokkaido Reg. Fish. Res. Lab. 10.
15 pp.</p></div>

<p><span class="pagenum"><a name="Page_220" id="Page_220">[Pg 220]</a></span></p>


<div class="footnotes"><h4>FOOTNOTES:</h4>

<div class="footnote">

<p><a name="Footnote_54_54" id="Footnote_54_54"></a><a href="#FNanchor_54_54"><span class="label">[54]</span></a> For crabs, this measurement is carapace width.</p></div>

<div class="footnote">

<p><a name="Footnote_55_55" id="Footnote_55_55"></a><a href="#FNanchor_55_55"><span class="label">[55]</span></a> Authors' data.</p></div>

<div class="footnote">

<p><a name="Footnote_56_56" id="Footnote_56_56"></a><a href="#FNanchor_56_56"><span class="label">[56]</span></a> For crabs, the measurements are total length for zoeal stages and carapace length and width for postzoeal stages.</p></div>

<div class="footnote">

<p><a name="Footnote_57_57" id="Footnote_57_57"></a><a href="#FNanchor_57_57"><span class="label">[57]</span></a> The incubation period for an egg is temperature dependent. Embryo development is faster at higher temperatures.</p></div>

<div class="footnote">

<p><a name="Footnote_58_58" id="Footnote_58_58"></a><a href="#FNanchor_58_58"><span class="label">[58]</span></a> Juvenile pollock have diurnal migrations.</p></div>

<div class="footnote">

<p><a name="Footnote_59_59" id="Footnote_59_59"></a><a href="#FNanchor_59_59"><span class="label">[59]</span></a> The peak period varies with latitude: to 55&deg;N&mdash;June; to 55-60&deg;N&mdash;July; to north of 60&deg;N&mdash;August.</p></div>

<div class="footnote">

<p><a name="Footnote_60_60" id="Footnote_60_60"></a><a href="#FNanchor_60_60"><span class="label">[60]</span></a> H. R. Carlson and R. E. Haight (in preparation), Juvenile life of Pacific ocean perch, <em>Sebastes alutus</em>, in coastal fiords of southeastern Alaska:
their environment, growth, food habits, and schooling behavior.</p></div>

<div class="footnote">

<p><a name="Footnote_61_61" id="Footnote_61_61"></a><a href="#FNanchor_61_61"><span class="label">[61]</span></a> The genus <em>Sebastes</em> is a live bearer.</p></div>

<div class="footnote">

<p><a name="Footnote_62_62" id="Footnote_62_62"></a><a href="#FNanchor_62_62"><span class="label">[62]</span></a> Rockfish larvae resemble each other quite closely, and complete descriptions for the 10 species in the Bering Sea do not exist. The following
depth distribution for rockfish larvae may or may not include <em>S. alutus</em>: 45-365 m (Taylor 1967) off British Columbia; 0-88 m (Ahlstrom 1959,
1961) off California and Baja California.</p></div>

<div class="footnote">

<p><a name="Footnote_63_63" id="Footnote_63_63"></a><a href="#FNanchor_63_63"><span class="label">[63]</span></a> In crabs, the eggs are attached to the female.</p></div>

<div class="footnote">

<p><a name="Footnote_64_64" id="Footnote_64_64"></a><a href="#FNanchor_64_64"><span class="label">[64]</span></a> S. C. Jewett and R. E. Haight (in preparation), A description of megalopa of the snow crab, <em>Chionoecetes bairdi</em> Rathbun (Majidae, subfamily
Oregoniinae).</p></div>

<div class="footnote">

<p><a name="Footnote_65_65" id="Footnote_65_65"></a><a href="#FNanchor_65_65"><span class="label">[65]</span></a> Spawning occurs in May in the eastern Bering Sea, but the total period is not known.</p>

<p><span class="pagenum"><a name="Page_221" id="Page_221">[Pg 221]</a></span></p></div></div>
<div class="chapter"></div>
<hr class="chap" />




<h3>Interrelations Between Seabirds and Introduced Animals</h3>


<p class="center">by</p>

<p class="center">Robert D. Jones, Jr.</p>

<p class="center">U.S. Fish and Wildlife Service<br />
1011 East Tudor Road<br />
Anchorage, Alaska 99507</p>

<p class="center">and</p>

<p class="center">G. Vernon Byrd<a name="FNanchor_66_66" id="FNanchor_66_66"></a><a href="#Footnote_66_66" class="fnanchor">[66]</a></p>

<p class="center">U.S. Fish and Wildlife Service<br />
Aleutian Islands National Wildlife Refuge<br />
Adak, Alaska
</p>


<h4>Abstract</h4>

<div class="blockquot">

<p>Animals introduced to insular seabird habitats are of both intentional and accidental
origin. The results of the introductions&mdash;particularly of herbivores&mdash;cannot
be predicted, but may range from severely destructive to beneficial. Herbivores
are of both domestic and wild stocks of ungulates, hares, and rabbits. Rats
are the most commonly introduced omnivore on a worldwide basis. In Alaska the
commonest carnivore introduction has been the red fox <em>(Vulpes fulva)</em> and arctic
fox <em>(Alopex lagopus)</em>, and the first of these were made in the early 19th century
by the Russian-American Company. These foxes nearly extirpated the Aleutian
Canada goose <em>(Branta canadensis leucopareia)</em> from its nesting grounds. Black
flies (Simuliidae), which are vectors of avian blood parasites, have been introduced
to three of the Aleutian Islands.</p></div>

<p>The purpose of this paper is to discuss some
influences of introduced animals, primarily
mammals, on seabirds and their nesting habitat,
with emphasis on the coasts of Washington,
British Columbia, and Alaska. Our discussion
focuses on island introductions partly
because a large proportion of seabirds choose
island nesting sites, and because islands present
ecosystems vulnerable to such introductions.</p>

<p>Flightless animals have no means of immigration,
hence little probability of colonizing
islands. In these circumstances marine birds
evolve populations in relatively simple ecosystems
(Carlquist 1965; MacArthur and Wilson
1967), though the degree of simplicity depends
on several variables, including the island's
size and its distance from a source of
immigrants. These systems have achieved
ecological homeostasis through reciprocal
adaptation over an extended period. Experience
has shown that introductions to such
systems result in severe perturbations (Odum
1971:221).</p>

<p>The introductions can be categorized as
being either intentional or accidental events.
Effects of such introductions have varied
widely, depending on the type of animal introduced,
the types of birds present and the habitat
they occupy, the size and shape of the island,
the type of nesting area used by the
birds, and the status of their populations before
the introduction. An example drawn from
our Aleutian experience with gallinaceous
birds illustrates the interaction of these variables.
The dark phase of the arctic fox <em>(Alopex
lagopus)</em> was introduced to Adak and Amchitka
islands, both of which had native populations
of the rock ptarmigan, <em>Lagopus mutus</em>
<span class="pagenum"><a name="Page_222" id="Page_222">[Pg 222]</a></span>(Gabrielson and Lincoln 1959). Foxes were released
on Adak in 1924, and on Amchitka in
1921. Adak has an area of 751 km<sup>2</sup> and Amchitka
350 km<sup>2</sup>. Adak is irregular in shape
with extensive precipitous shorelines, relatively
few beaches, and a large, central mountainous
hinterland which foxes rarely penetrated.
Amchitka, on the other hand, presents
a zone of marine planation on its eastern two
thirds, low mountains on the rest, shelving
beaches around most of the island, and a long,
linear, narrow shape that foxes explored completely.
By 1949 ptarmigan were difficult to
find on Amchitka, and then only in the
highest, steepest section of the mountains.
They were extirpated from the low, eastern
two thirds of the island. The foxes flourished
on Amchitka, but did much less well on Adak,
where the ptarmigan population fluctuated in
a normal cyclic manner, apparently uninfluenced
by the foxes. Then the foxes were eradicated
on Amchitka in the 1950's, and by 1962
the ptarmigan had spread over the whole of
the island and become one of the most conspicuous
avian features of the landscape.</p>


<h4>Animal Introductions</h4>


<h5><em>Non-predatory Animals</em></h5>

<p>Man has taken ungulates with him to many
islands. Although numerous records of livestock
introductions are available, few provide
information relating to the effects of these
animals on the habitat and their fauna unless
the impact has been severe.</p>

<p>A most noteworthy example of destruction
by ungulates occurred on Guadalupe Island
off the coast of Baja California. Domestic
goats <em>(Capra hircus)</em> were introduced in the
unrecorded past with the result that little of
the once abundant vegetation remains. In its
place introduced species capable of withstanding
heavy grazing are abundant over most of
the island. Several endemic avian species are
now considered extinct, including the Guadalupe
storm-petrel, <em>Oceanodroma macrodactyla</em>
(Howell and Cade 1954; Jehl 1972).</p>

<p>Sheep <em>(Ovis aries)</em> have been introduced to
seabird nesting islands with varying results.
In Bass Strait, Australia, Norman (1970)
studied the effects of introduced sheep on
vegetation and birds. He cited various papers
attributing destruction of colonies of shearwaters
(<em>Puffinus</em> sp.) to the activities of sheep,
primarily their treading on the burrows. He
found, however, that on Big Green Island and
Phillip Island, sheep were not responsible for
declines in shearwater breeding success, nor
did they prevent expansion of colonies.</p>

<p>Other authors have reported damage to seabird
nesting areas by sheep. One such
example in the eastern North Pacific region
concerns Protection Island, Washington. According
to Richardson (1961), 100 to 300
sheep grazed freely on the island since 1958.
He reported damage by grazing and frequent
trampling of nesting areas of rhinoceros auklets
<em>(Cerorhinca monocerata)</em>. Landslides were
initiated by these activities, rendering the
slopes unusable by auklets. Of the burrows in
his study area, 46% were buried by slides. He
did not determine mortality.</p>

<p>Other avian consequences may flow from
sheep introductions. Husbandry of these
ungulates has been practiced with varying
success for many years in the Aleutian
Islands, most notably on Umnak and Unalaska
islands, both of which have large native
populations of bald eagles, <em>Haliaeetus
leucocephalus</em> (Gabrielson and Lincoln 1959).
Before the introduction of sheep, these raptors
were oriented to the sea, hunting fish and
seabirds. Sheep presented a new resource and
presently the industry found itself confronted
by a formidable predator, and demanded that
eagles be destroyed (letter to William Egan,
Governor of Alaska, from James S. Bynum,
Secretary-treasurer, Umnak Company, Inc.).</p>

<p>Other ungulates introduced on Alaska islands
include cattle <em>(Bos taurus)</em> on Chernofski
and Chernabura islands; caribou <em>(Rangifer
tarandus)</em> on Adak; reindeer on St. Matthew,
Nunivak, Atka, Umnak, St. Paul, St.
Lawrence, Hagemeister, and Kodiak as well
as many interior locations; deer <em>(Odocoileus
hemionus)</em> on Kodiak and Afognak; elk
<em>(Cervus canadensis)</em> on Afognak; and musk
oxen <em>(Ovibos moschatus)</em> on Nunivak. All
these animals have maintained populations on
islands for a time, and some appear likely to
do so into the distant future. Specific effects
on seabirds is generally not known, but
trampling of grassy slopes such as that re<span class="pagenum"><a name="Page_223" id="Page_223">[Pg 223]</a></span>ported
for sheep develops in some cases.
Bailey et al. (1933) speculated that nests of
the snow goose <em>(Anser caerulescens)</em> were destroyed
by reindeer or their herdsmen in the
Point Barrow area.</p>

<p>The destruction of vegetation by introduced
rabbits and hares has been documented for
many areas in the world. This destruction has
often extended to seabirds. Perhaps the most
dramatic example occurred on Laysan Island
in the Hawaiian archipelago, where rabbits of
unknown species were introduced in 1903. According
to Warner (1963) it took less than 20
years for the rabbits to remove every green
plant but three patches of <em>Sesuvium portulacastrum</em>.
The Laysan duck <em>(Anas laysanensis)</em>
was brought perilously close to extinction.
The rabbits were eliminated in the
1920's, and the population of ducks increased
to over 600 by 1963, a figure thought to approximate
the pre-disturbance population.</p>

<p>European hares <em>(Lepus europaeus)</em> were introduced
on Smith, San Juan, and Long islands,
in Washington. On Smith Island, these
burrowing animals apparently grazed nearly
all the succulent vegetation close to the
ground. By 1924, their burrows riddled the
bluffs, causing them to cave into the ocean
(Couch 1929). Couch found no seabirds nesting
on the island, but found numerous tufted
puffins <em>(Lunda cirrhata)</em> present on the bluffs,
but not nesting. A removal campaign was directed
against the hares in 1924 and in a few
years they were gone. Smith Island now supports
nesting pelagic birds (D. Manuwal, personal
communication).</p>

<p>Accounts of hare and rabbit introductions
to islands are legion, but not all such introductions
have drastically affected seabirds.
Manana Island, Hawaii, is such a case.
Tomich et al. (1968) believed that introduced
rabbits <em>(Oryctolagus cuniculas)</em> were not even
indirectly detrimental to the nesting noddies
<em>(Anous tolidus)</em> and sooty terns <em>(Sterna fuscata)</em>.
In some situations, introduced lagomorphs
have been credited with benefiting
seabirds. Lockley (1942) suggested that
rabbits helped to open new breeding colonies
of manx shearwaters <em>(Puffinus puffinus)</em> at
Skomer and in west Wales in general. In
Alaska rabbits were introduced to Middleton
Island in 1952 (Rausch 1958) and to Ananiuliak
Island at an earlier unrecorded date. Both
have developed sustaining populations in the
presence of large seabird populations without
measurable effect on the birds. On Ananiuliak
glaucous-winged gulls <em>(Larus glaucescens)</em>
have been observed feeding on rabbits (W. S.
Laughlin, personal communication).</p>

<p>Invertebrates have been introduced on
three islands in the Aleutians. The black fly
(<em>Simulium</em> sp.) reached Adak by 1958,
Shemya by 1964, and Amchitka in connection
with activities of the Atomic Energy Commission
in 1968. Apparently the insects were
transported on jet aircraft. The pest appears
well established on Adak, but its status on the
other two islands is uncertain. Like the mosquito,
the female black fly sucks blood from
warm-blooded animals, and in the process becomes
the vector of a <em>Leucocytozoan</em> blood
parasite of birds. In years of black fly abundance
at Seney (Michigan) National Wildlife
Refuge the blood parasite has been responsible
for reproductive failure in Canada geese
(<em>Branta canadensis</em>; Sherwood 1968). If black
fly problems reach such a scale in the Aleutians,
the parasites might prove limiting to
pelagic birds as well as to waterfowl. Winds,
for which the Aleutian region is famous, constitute
a limiting factor for obligate blood-feeding
Simuliids and may control the
severity of this problem.</p>


<h5><em>Predatory Animals</em></h5>

<p>The list of introduced animals that prey on
seabirds is extensive. Often several animals
have been introduced to the same island. For
example, in 1951 Amchitka Island in the
Aleutians supported populations of feral dogs
<em>(Canis familiaris)</em> and cats <em>(Felis catus)</em>, rats
<em>(Rattus norvegicus)</em>, and arctic fox. Their
presence resulted from three of the usual
sources of predator introductions: escape of
pets, escape from visiting ships (and aircraft),
and commercial introductions. Add introductions
to control pests, such as that of the mongoose
<em>(Herpestes auropunctatus)</em> to the
Hawaiian Islands, and only one source remains&mdash;the
desire of man to improve on nature.
In the Aleutians this impulse has taken
the more innocuous form of fish and plant introductions,
such as rainbow trout <em>(Salmo
gairdneri)</em> on Adak and Shemya, and trees<span class="pagenum"><a name="Page_224" id="Page_224">[Pg 224]</a></span>
(mostly Sitka spruce, <em>Picea sitkensis</em>) on
every military base in the "Chain."</p>

<p>Rats appear to be the most commonly introduced
predators on a worldwide scale. Ships
furnish the traditional source of their introduction,
but one of us (R.D.J.) has observed
them disembarking from a military aircraft at
Cold Bay on the Alaska Peninsula. These animals
probably entered the plane at Adak,
which received rats from military ships early
in World War II.</p>

<p>Rats may be able to exploit a larger percentage
of the seabird species on a given island
than other introduced predators because they
can enter crevices and burrows in search of
the birds and their eggs and young. They also
destroy ground-nesters, and cliff-nesters may
not be altogether safe from them. Clayton M.
White (personal communication) found that
<em>Rattus norvegicus</em> had ravaged every one of
16 eyries of the peregrine falcon <em>(Falco peregrinus)</em>
that he checked in 1971 at Amchitka
Island, Alaska. Only one egg had tooth
marks, however. Kenyon (1961) ascribed the
disappearance of the song sparrow <em>(Melospiza
melodia maxima)</em> and the winter wren <em>(Troglodytes
troglodytes kiskensis)</em> from Amchitka
to predation by rats.</p>

<p>Many authors have mentioned potential rat
damage, but few have quantitatively documented
it. Imber (1974), however, provided
data concerning the magnitude of rat predation
on diving petrels and storm-petrels on
some New Zealand islands. He found that rats
were taking between 10 and 35% of the chicks
of gray-faced petrels <em>(Pterodroma macroptera
gouldi)</em> on Whale Island in the parts of the
colonies where burrows were dense. On those
parts of the island with a very low density of
petrel burrows, rats were believed to have
killed virtually every chick. Imber revealed
that where Polynesian rats <em>(Rattus exulans)</em>
occur, diving petrels and storm-petrels are
rare or absent, though they are widespread on
neighboring islands. Imber concluded from
his studies of the ecology of petrels and Polynesian
and Norway rats that a petrel colony is
endangered if invaded by a species of rat
whose maximum weight approaches or exceeds
the mean adult weight of the petrel.
Harris (1970), who worked with dark-rumped
petrels <em>(Pterodroma phacopygia)</em> on Santa
Cruz in the Galapagos Islands, indicated that
black rats <em>(Rattus rattus)</em> were responsible for
the extremely low nesting success of the
petrels there.</p>

<p>In British Columbia, Campbell (1968) recorded
predation by the Alexandrian rat
<em>(R. rattus)</em> on ancient murrelets <em>(Synthliboramphus
antiquus)</em> at Langara Island. The
extent of damage to the murrelet population
is not known.</p>

<p>The animals most widely introduced in
Alaska seabird habitat are the red fox <em>(Vulpes
fulva)</em> and the arctic fox. The red fox is native
to the Alaska Peninsula and to the easternmost
group of islands in the Aleutians, known
as the Lissii or Fox Islands (Berkh 1823;
Murie 1959). At the other end of the archipelago,
in the group known as the Near Islands,
Attu Island has a native population of the
arctic fox (Tikhmenev 1861; Bancroft 1886).
Between Umnak Island, the westernmost island
of the Fox group, and Attu there are no
native terrestrial mammals, and substantial
avian populations evolved an ecology in the
absence of mammalian predation (Murie
1959).</p>

<p>At the time of Russian contact with the
Aleutians, both fox species were dominantly
dark phase, and the early introductions
(about 1836) by the Russian-American Company
were of both species (Tikhmenev 1861).
Initially both species were successful in developing
insular populations, but in the long
run the arctic fox proved the more successful.
At Great Sitkin, Adak, and Kanaga, introduced
red foxes maintained populations that
were eliminated in the 1920's to be replaced
by arctic foxes (unpublished records of the
Aleutian Islands National Wildlife Refuge).
Differential harvest of the preferred dark
phase had in the meantime altered the genetic
makeup of the population, and the light phase
had become dominant. In the arctic fox populations,
the dark phase remained generally
dominant at about 95%, but in some small islands
with limited genetic stock (e.g., the
Semichis) the proportion approached one to
one (unpublished records of the Aleutian Islands
National Wildlife Refuge).</p>

<p>By 1936, the Aleutian archipelago constituted
a large-scale fox farm, which in its 23
years of existence as a refuge had produced
25,641 fox pelts with a value of $1,162,826.
During the same period, and perhaps earlier,<span class="pagenum"><a name="Page_225" id="Page_225">[Pg 225]</a></span>
arctic foxes were introduced on almost every
island from the Aleutians to Prince William
Sound, and on some of the islands in southeastern
Alaska. The Aleutian Islands National
Wildlife Refuge maintained records
from which the above figures are quoted, but
though records of other islands' use for fur
farms exist in the archives of the Alaska
Game Commission, no record of the fur values
was kept.</p>

<p>Murie (1959) assessed the influence of the
foxes by examining 2,501 fox droppings collected
in 1936 and 1937 from 22 of the Aleutian
Islands. He reported 57.8% of the food
items in these droppings was avian&mdash;48.9%
seabirds. The result of his investigations was
the adoption of new policies governing issuance
of permits for fox farming in the
Refuge. The essential feature of these policies
was the revocation of certain existing permits,
with a view to reserving the islands concerned
for wildlife use. The decision proved
academic, for fur prices declined until no market
for Aleutian arctic fox pelts could be
found. But the foxes remained.</p>

<p>The most obvious damage has been the
nearly complete extermination of the Aleutian
Canada goose <em>(Branta canadensis leucopareia)</em>.
It has vanished from its former nesting
range in the Aleutian and Kuril Islands,
except for Buldir Island in the western Aleutians
(Jones 1963). Clark (1910) described this
goose as extremely abundant on Agattu Island
in 1909; however, foxes from Attu were
introduced there in 1923, 1925, and 1929.
Murie (1959) found "probably less than six
pairs" in 4 days of traveling over the island in
1937.</p>

<p>In our main area of interest, cats appear to
have been widely introduced, but we found no
record of extensive predation on marine birds.
Jehl (1972) attributed the extinction of the
Guadalupe petrel to predation by cats, in combination
with the destruction of vegetation by
goats. Imber (1974) reported that "serious
predation by cats upon a colony of gray-faced
petrels on Little Barrier Island, New Zealand
was observed in 1950. Since that time, the
colony has become extinct."</p>

<p>Though feral dogs are reported present on
islands in our area of interest, they do not appear
to have significant influence on seabirds.
On Attu Island, the pet dogs of personnel of
the Coast Guard LORAN station are reported
to take common eiders <em>(Somateria
mollissima)</em>.</p>


<h4>Conclusions</h4>

<p>Ecological consequences of animal introductions
to islands are rarely well documented.
Usually no thought is devoted to such consequences
until redress becomes difficult or
quite impossible. Many of the introductions
stem from a period before ecological understanding,
and the introduced animal has acquired
the status of a native. The arctic fox in
the Aleutians fits all of these conditions. Until
we conducted a thorough search of the literature,
some of it difficult to secure and written
in several languages, the original status of
this animal was not known. Its elimination,
now under way on selected islands, is difficult
and expensive. Rapid recovery of some avian
species, including certain passerines, has been
observed. However, ecological homeostasis is
the product of evolution, and restoration in
the Aleutians must follow that course. It is
not likely to proceed rapidly to a point
thought desirable by man. The accidental introductions
of animals such as rats and black
flies in the Aleutians constitute particularly
irksome events because they cannot be reversed.
The new ecology of Amchitka, from
which the foxes have been removed, must
evolve in the presence of these species. Its
face will look very different than if they were
not there. We would like to suggest a means
by which such introductions may be prevented,
but it seems likely that more, not less,
can be expected.</p>

<p>Preventing the introduction of ungulates
seems more likely to be successful, especially
if the islands lie within a National Wildlife
Refuge. Even this, however, becomes less certain
with an expanding human population
and, with it, demands for more land on which
to produce food.</p>

<p>Legal restrictions have been suggested as a
means to control or prevent introductions,
but in the northern islands, little enforcement
is likely. There is a phrase bearing on this,
said to have governed human behavior in the
early years of Caucasoid occupation of the
Aleutian Islands, "Heaven is too high and the
Czar too far away."</p>

<p><span class="pagenum"><a name="Page_226" id="Page_226">[Pg 226]</a></span></p>


<h4>References</h4>

<div class="hangindent">

<p>Bailey, A. M., C. D. Brower, and L. B. Bishop. 1933.
Birds of the region of Point Barrow, Alaska.
Prog. Act. Chic. Acad. Sci. 4(2):14-40.</p>

<p>Bancroft, H. H. 1886. History of Alaska, 1730-1885.
San Francisco.</p>

<p>Berkh, V. 1823. The chronological history of the discovery
of the Aleutian Islands or the exploits of
the Russian merchants. N. Grech., St. Petersburg.</p>

<p>Campbell, R. W. 1968. Alexandrian rat predation on
ancient murrelet eggs. Murrelet 49:38.</p>

<p>Carlquist, S. 1965. Island life: a natural history of
the islands of the world. Natural History Press,
New York.</p>

<p>Clark, A. H. 1910. The birds collected and observed
during the cruises of the United States Fisheries
Steamer "Albatross" in the North Pacific Ocean
and in the Bering, Okhotsk, Japan, and Eastern
seas from April to December 1906. Proc. U.S.
Natl. Mus. 38:25-74.</p>

<p>Couch, L. K. 1929. Introduced European rabbits in
the San Juan Islands, Washington. J. Mammal.
10:334-336.</p>

<p>Gabrielson, I. N., and F. C. Lincoln. 1959. The birds
of Alaska. Stackpole Co. and Wildlife Management
Institute, Washington, D.C.</p>

<p>Harris, M. P. 1970. The biology of an endangered
species, the dark-rumped petrel (<em>Pterodroma
phacopygia)</em> in the Galapagos Islands. Condor
72:76-84.</p>

<p>Howell, T. R., and T. J. Cade. 1954. The birds of
Guadalupe Island in 1953. Condor 56:283-291.</p>

<p>Imber, M. J. 1974. The rare and endangered species
of the New Zealand region and the policies that
exist for their management: petrels and predators.
Paper presented to the International Council
for Bird Preservation (XVI World Conference),
Canberra.</p>

<p>Jehl, J. R. 1972. On the cold trail of an extinct
petrel. Pac. Discovery 25:24-28.</p>

<p>Jones, R. D. 1963. Buldir Island, site of a remnant
breeding population of Aleutian Canada geese.
Wildfowl Trust Annu. Rep. 14:80-84.</p>

<p>Kenyon, K. W. 1961. Birds of Amchitka Island,
Alaska. Auk 78(3):305-326.</p>

<p>Lockley, R. M. 1942. Shearwaters. Devin-Adair,
New York.</p>

<p>MacArthur, R. H., and E. O. Wilson. 1967. The
theory of island biogeography. Princeton Univ.
Press, Princeton, N.J.</p>

<p>Murie, O. J. 1959. Fauna of the Aleutian Islands
and Alaska Peninsula. U.S. Fish Wildl. Serv., N.
Am. Fauna 61:1-364.</p>

<p>Norman, F. I. 1970. The effects of sheep on the
breeding success and habitat of the short-tailed
shearwater <em>(Puffinus tenuirostris)</em> (Temminch).
Aust. J. Zool. 18:215-242.</p>

<p>Odum, E. P. 1971. Fundamentals of ecology. W. B.
Saunders Co., Philadelphia, Pa. 574 pp.</p>

<p>Rausch, R. 1958. The occurrence and distribution of
birds on Middleton Island, Alaska. Condor
60(4):227-242.</p>

<p>Richardson, F. 1961. Breeding biology of the rhinoceros
auklet on Protection Island. Condor
63(6):456-473.</p>

<p>Sherwood, G. A. 1968. Factors limiting production
and expansion of local populations of Canada
geese. Pages 73-85 <em>in</em> R. L. Hine and C. Schoenfeld,
eds. Canada goose management. Dembar
Educational Research Service, Madison, Wis.</p>

<p>Tikhmenev, P. 1861. Historical review of the origin
of the Russian-American Co. and its activity up
to and the present time. Edward Weimar, St.
Petersburg.</p>

<p>Tomich, P. Q., N. Wilson, and C. H. Lamoureux.
1968. Ecological factors on Manana Island,
Hawaii. Pac. Sci. 12:352-368.</p>

<p>Warner, R. E. 1963. Recent history and ecology of
the Laysan duck. Condor 65(1):2-23.</p></div>

<div class="footnotes"><h4>FOOTNOTES:</h4>

<div class="footnote">

<p><a name="Footnote_66_66" id="Footnote_66_66"></a><a href="#FNanchor_66_66"><span class="label">[66]</span></a> Present address: Hawaiian Islands National Wildlife
Refuge, Kilauea, Hawaii.</p>

<p><span class="pagenum"><a name="Page_227" id="Page_227">[Pg 227]</a></span></p></div></div>
<div class="chapter"></div>
<hr class="chap" />




<h3>Oil Vulnerability Index for Marine Oriented Birds</h3>


<p class="center">by</p>

<p class="center">James G. King</p>

<p class="center">U.S. Fish and Wildlife Service<br />
P. O. Box 1287<br />
Juneau, Alaska 99802</p>

<p class="center">and</p>

<p class="center">Gerald A. Sanger</p>

<p class="center">U.S. Fish and Wildlife Service<br />
Anchorage, Alaska
</p>


<h4>Abstract</h4>

<div class="blockquot">

<p>The 176 species of birds using marine habitats of the Northeast Pacific are
graded on the basis of 20 factors that affect their survival. A score of 0, 1, 3, or 5,
respectively, representing no, low, medium, or high significance is assigned for
each factor. The total score is the Oil Vulnerability Index (OVI). The OVI's range
from 1 to 100, an index of 100 indicating the greatest vulnerability. Using this
system, one can rank the avifauna of different areas according to their vulnerability
to environmental hazards as an aid in making management decisions.</p></div>

<p>Today's decision makers require an ever-increasing
array of information and planning
documents. The Federal Government's requirement
for environmental impact statements
under the National Environmental Protection
Act of 1969 is but one example of this
trend. These documents generally consider
the effects of proposed actions on waterfowl
and a few other species of birds, but the bulk
of the avifauna is usually only listed, or sometimes
ignored completely. A simple system
for evaluating and presenting avian data is
badly needed so that those interested in birds,
whether technically trained or not, can easily
grasp the implications of proposed actions. It
is incumbent on biologists to devise new ways
of presenting their knowledge so that it can be
easily and effectively used by decision
makers, who are often less informed. In short,
biologists must do for the environmental impact
statement assessors what Roger Tory
Peterson did for the bird watchers by giving
them a simple and comprehensible system.</p>

<p>The need for a system to evaluate relative
vulnerabilities of bird populations is particularly
great for birds that are being increasingly
affected by marine oil pollution. The system
needs to allow comparisons of potential
impacts to birds resulting from various oil development
projects in different locations and
served by various modes of transport. The Oil
Vulnerability Index (OVI) is our attempt to
fulfill this informational need on the avifauna
of the Northeast Pacific. Insofar as we know,
this approach to assessing a wildlife management
problem has been attempted only for
ranking endangered species in a numeric ranking
system that identified where restoration
efforts could best be directed (Sparrowe and
Wight 1975).</p>

<p>We are indebted to Gene Ruhr and Keith
Schreiner for ideas generated in their work
with endangered species. Frank Pitelka,
James Bartonek, Kent Wohl, and Mary Lou
King reviewed portions of the manuscript and
offered helpful suggestions. Jack Hodges
helped prepare the OVI tables.</p>

<p><span class="pagenum"><a name="Page_228" id="Page_228">[Pg 228]</a></span></p>


<h4>Methods</h4>

<p>A list of 176 species of birds using marine
habitats in or near the States of Washington
and Alaska and the Province of British
Columbia (Table 1, left column) was compiled
from checklists by the American Ornithologists'
Union (AOU 1957) and Gibson (1970).
Nomenclature is from AOU (1957). The scientific
names of three species of shorebirds recently
identified in the Aleutian Islands that
were not listed by the AOU (1957) came from
Peterson et al. (1967).</p>

<p>Each bird was scored on 20 factors that affect
its survival (Table 1). Point scores for
most birds were either 0, 1, 3, or 5, indicating
no, low, medium, or high importance, respectively,
in their biology or habits as related to
Northeast Pacific oil development. Rare or
accidental species were given only one point
for occurrence, and endangered species 99
points for population size plus 1 point for occurrence.
Thus the potential range of the
OVI's is from 1 to 100.</p>

<p>The factors in Table are largely self-explanatory.
The items under "range" apply to
the entire world population of the species.
"Productivity" is derived from a combination
of clutch size and age at first nesting. Specialization
is used in the biological sense to compare
a versatile species like mallards (<em>Anas
platyrhynchos)</em> with a less versatile species
such as the trumpeter swan <em>(Olor buccinator)</em>.
Mortality under "history of oiling" is based
on our knowledge that some species (e.g., alcids)
have been more involved than others
such as gulls. Exposure relates to the level of
exposure within the Pacific area in any
season.</p>

<p>Information on many of the factors for
many species is scanty at best, and subjective
appraisals were made by us when information
was lacking. Opinions as to appropriate
scores will vary among experts. References
used, in part, in preparing Table 1 were: AOU
1957; Fay and Cade 1959; Gabrielson and
Lincoln 1959; Isleib and Kessel 1973; Kortright
1942; Murie 1959; Palmer 1962; Robbins
et al. 1966; Sanger 1972; and Stout et al. 1967.</p>


<h4>Results</h4>

<p>The OVI for each of 176 bird species is
listed in Table 1. The average OVI for 22
avian families comprising 128 species that are
neither rare stragglers nor endangered ranged
from 19 to 88, with a mean of 51 (Table 3).</p>

<p>Tables 4 and 5 show a possible use for the
OVI by comparing impacts in two large,
widely separated areas. A species list from
Southeast Alaska (U.S. Forest Service and
Alaska Department of Fish and Game 1970) is
compared with a list from the Aleutian Islands
(U.S. Fish and Wildlife Service 1974).
Only commonly occurring species are included.
These tables graphically display
rather strong differences in the vulnerability
of the avifauna of each area. A person explaining
comparative impacts of projects might
use the tables in the following way:</p>

<p>&#8226; Column 1, with scores from 1 to 20 points,
indicates birds with a low level of project involvement,
where damage or future costs
would not be expected. As this will normally
be the longest list, as in Tables 4 and 5, one
would expect an immediate rise of interest on
the part of the planning agency, which is probably
eager to learn where problems will be
fewest.</p>

<p>&#8226; Column 2 (21 to 40 points) indicates birds
for which there is a low level of concern. Perhaps
all that is needed is a review to determine
if special characteristics of the project
might be detrimental to these species.</p>

<p>&#8226; Column 3 (41 to 60 points) might be called
"trial and error" species. If some birds are adversely
affected, it will not be catastrophic.
As the project develops it will be merely necessary
to monitor these to make sure their
status is not adversely affected. If it is, there
will be time to develop conservation
measures.</p>

<p>&#8226; Columns 4 and 5 (61 to 80 points and 81 to
100 points, respectively) include the species
where concern is high. It is for these species
that research money will be needed, where
project modifications may be required, where
a contingency plan in case of disaster is
needed, where a conservation technology will
be needed, and where periodic project shutdown
could be called for.</p>

<p><span class="pagenum"><a name="Page_229" id="Page_229">[Pg 229]</a></span></p>

<div class="center">
<table border="1" cellpadding="4" cellspacing="0" summary="Table 1. Oil Vulnerability Index (OVI) for waterbirds in the Northeast Pacific Region.">
    <caption>Table 1. <em>Oil Vulnerability Index (OVI) for waterbirds in the Northeast Pacific Region.</em></caption>
<col></col>
<col span="21" width="4%"></col>

<thead>
  <tr>
    <th rowspan="2">Family, common (AOU) name and scientific name</th>
    <th colspan="4" class="br">Range</th>
    <th colspan="2" class="br">Pop&shy;u&shy;la&shy;tion</th>
    <th colspan="6" class="br">Hab&shy;its</th>
    <th colspan="4" class="br">Mor&shy;tal&shy;i&shy;ty</th>
    <th colspan="4" class="br">An&shy;nu&shy;al ex&shy;po&shy;sure</th>
    <th>OVI</th>
  </tr>
  <tr>
    <th>B<br />r<br />e<br />e<br />d<br />i<br />n<br />g<br /> <br />r<br />a<br />n<br />g<br />e<br /> <br />s<br />i<br />z<br />e</th>
    <th>M<br />i<br />g<br />r<br />a<br />t<br />i<br />o<br />n<br /> <br />l<br />e<br />n<br />g<br />t<br />h</th>
    <th>W<br />i<br />n<br />t<br />e<br />r<br /> <br />r<br />a<br />n<br />g<br />e<br /> <br />s<br />i<br />z<br />e</th>
    <th>M<br />a<br />r<br />i<br />n<br />e<br /> <br />o<br />r<br />i<br />e<br />n<br />t<br />a<br />t<br />i<br />o<br />n</th>
    <th>P<br />o<br />p<br />u<br />l<br />a<br />t<br />i<br />o<br />n<br /> <br />s<br />i<br />z<br />e</th>
    <th>P<br />r<br />o<br />d<br />u<br />c<br />t<br />i<br />v<br />i<br />t<br />y</th>
    <th>R<br />o<br />o<br />s<br />t<br />i<br />n<br />g</th>
    <th>F<br />o<br />r<br />a<br />g<br />i<br />n<br />g</th>
    <th>E<br />s<br />c<br />a<br />p<br />e</th>
    <th>F<br />l<br />o<br />c<br />k<br />i<br />n<br />g<br /> <br />o<br />n<br /> <br />w<br />a<br />t<br />e<br />r</th>
    <th>N<br />e<br />s<br />t<br />i<br />n<br />g<br /> <br />d<br />e<br />n<br />s<br />i<br />t<br />y</th>
    <th>S<br />p<br />e<br />c<br />i<br />a<br />l<br />i<br />z<br />a<br />t<br />i<br />o<br />n</th>
    <th>H<br />u<br />n<br />t<br />e<br />d<br /> <br />b<br />y<br /> <br />m<br />a<br />n</th>
    <th>A<br />n<br />i<br />m<br />a<br />l<br /> <br />d<br />e<br />p<br />r<br />e<br />d<br />a<br />t<br />i<br />o<br />n<br />s</th>
    <th>N<br />o<br />n<br />-<br />o<br />i<br />l<br /> <br />p<br />o<br />l<br />l<br />u<br />t<br />i<br />o<br />n</th>
    <th>H<br />i<br />s<br />t<br />o<br />r<br />y<br /> <br />o<br />f<br /> <br />o<br />i<br />l<br />i<br />n<br />g</th>
    <th>S<br />p<br />r<br />i<br />n<br />g</th>
    <th>S<br />u<br />m<br />m<br />e<br />r</th>
    <th>F<br />a<br />l<br />l</th>
    <th>W<br />i<br />n<br />t<br />e<br />r</th>
    <th>T<br />o<br />t<br />a<br />l<br /> <br />P<br />o<br />i<br />n<br />t<br />s</th>
  </tr>
</thead>
<tbody>
  <tr>
    <td colspan="0">Gaviidae</td>
  </tr>
  <tr>
    <td class="tdp">Common loon <em>(Gavia immer)</em></td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr"> 1</td>
    <td class="tdr">5</td>
    <td class="tdr">5</td>
    <td class="tdr">5</td>
    <td class="tdr">5</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">0</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">47</td>
  </tr>
  <tr>
    <td class="tdp">Yellow-billed loon <em>(G. adamsii)</em></td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">5</td>
    <td class="tdr">3</td>
    <td class="tdr"> 5</td>
    <td class="tdr">5</td>
    <td class="tdr">5</td>
    <td class="tdr">5</td>
    <td class="tdr">5</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">0</td>
    <td class="tdr">3</td>
    <td class="tdr">5</td>
    <td class="tdr">1</td>
    <td class="tdr">5</td>
    <td class="tdr">5</td>
    <td class="tdr">65</td>
  </tr>
  <tr>
    <td class="tdp">Arctic loon <em>(G. arctica)</em></td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr"> 3</td>
    <td class="tdr">5</td>
    <td class="tdr">5</td>
    <td class="tdr">5</td>
    <td class="tdr">5</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">58</td>
  </tr>
  <tr>
    <td class="tdp">Red-throated loon <em>(G. stellata)</em></td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">5</td>
    <td class="tdr"> 1</td>
    <td class="tdr">5</td>
    <td class="tdr">5</td>
    <td class="tdr">5</td>
    <td class="tdr">5</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">0</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">49</td>
  </tr>
  <tr>
    <td colspan="0">Podicipedidae</td>
  </tr>
  <tr>
    <td class="tdp">Red-necked grebe (<em>Podiceps grisegena</em>)</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr"> 1</td>
    <td class="tdr">3</td>
    <td class="tdr">5</td>
    <td class="tdr">5</td>
    <td class="tdr">5</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">0</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">0</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">44</td>
  </tr>
  <tr>
    <td class="tdp">Horned grebe <em>(P. auritus)</em></td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr"> 1</td>
    <td class="tdr">3</td>
    <td class="tdr">5</td>
    <td class="tdr">5</td>
    <td class="tdr">5</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">0</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">0</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">48</td>
  </tr>
  <tr>
    <td class="tdp">Western grebe (<em>Aechmophorus occidentalis</em>)</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">5</td>
    <td class="tdr"> 1</td>
    <td class="tdr">3</td>
    <td class="tdr">5</td>
    <td class="tdr">5</td>
    <td class="tdr">5</td>
    <td class="tdr">5</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">0</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">5</td>
    <td class="tdr">1</td>
    <td class="tdr">0</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">56</td>
  </tr>
  <tr>
    <td colspan="0">Diomedeidae</td>
  </tr>
  <tr>
    <td class="tdp">Short-tailed albatross <em>(Diomedea albatrus)</em></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdr">99</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdr">1</td>
    <td class="tdr">100</td>
  </tr>
  <tr>
    <td class="tdp">Black-footed albatross <em>(D. nigripes)</em></td>
    <td class="tdr">5</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">5</td>
    <td class="tdr"> 3</td>
    <td class="tdr">5</td>
    <td class="tdr">5</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">5</td>
    <td class="tdr">5</td>
    <td class="tdr">0</td>
    <td class="tdr">0</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">50</td>
  </tr>
  <tr>
    <td class="tdp">Laysan albatross <em>(D. immutabilis)</em></td>
    <td class="tdr">5</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">5</td>
    <td class="tdr"> 3</td>
    <td class="tdr">5</td>
    <td class="tdr">5</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">5</td>
    <td class="tdr">5</td>
    <td class="tdr">0</td>
    <td class="tdr">0</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">52</td>
  </tr>
  <tr>
    <td colspan="0">Procellaridae</td>
  </tr>
  <tr>
    <td class="tdp">Fulmar <em>(Fulmarus glacialis)</em></td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">5</td>
    <td class="tdr"> 1</td>
    <td class="tdr">5</td>
    <td class="tdr">5</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">5</td>
    <td class="tdr">3</td>
    <td class="tdr">0</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">57</td>
  </tr>
  <tr>
    <td class="tdp">Pink-footed shearwater <em>(Puffinus creatopus)</em></td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">5</td>
    <td class="tdr"> 1</td>
    <td class="tdr">5</td>
    <td class="tdr">5</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">5</td>
    <td class="tdr">3</td>
    <td class="tdr">0</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">47</td>
  </tr>
  <tr>
    <td class="tdp">Pale-footed shearwater <em>(P. carneipes)</em></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdr">1</td>
    <td class="tdr"> 1</td>
  </tr>
  <tr>
    <td class="tdp">New Zealand shearwater <em>(P. bulleri)</em></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdr">1</td>
    <td class="tdr"> 1</td>
  </tr>
  <tr>
    <td class="tdp">Sooty shearwater <em>(P. griseus)</em></td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">5</td>
    <td class="tdr"> 1</td>
    <td class="tdr">5</td>
    <td class="tdr">5</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">5</td>
    <td class="tdr">5</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">5</td>
    <td class="tdr">1</td>
    <td class="tdr">0</td>
    <td class="tdr">51</td>
  </tr>
  <tr>
    <td class="tdp">Slender-billed shearwater <em>(P. tenuirostris)</em></td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">5</td>
    <td class="tdr"> 1</td>
    <td class="tdr">5</td>
    <td class="tdr">5</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">5</td>
    <td class="tdr">5</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">5</td>
    <td class="tdr">1</td>
    <td class="tdr">0</td>
    <td class="tdr">53</td>
  </tr>
  <tr>
    <td class="tdp">Scaled petrel (<em>Pterodroma inexpectata</em>)</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdr">1</td>
    <td class="tdr"> 1</td>
  </tr>
  <tr>
    <td class="tdp">Cooks petrel <em>(P. cookii)</em></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdr">1</td>
    <td class="tdr"> 1</td>
  </tr>
  <tr>
    <td colspan="0">Hydrobatidae</td>
  </tr>
  <tr>
    <td class="tdp">Fork-tailed storm-petrel <em>(Oceanodroma furcata)</em></td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">5</td>
    <td class="tdr"> 1</td>
    <td class="tdr">5</td>
    <td class="tdr">5</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">5</td>
    <td class="tdr">3</td>
    <td class="tdr">0</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">5</td>
    <td class="tdr">5</td>
    <td class="tdr">5</td>
    <td class="tdr">5</td>
    <td class="tdr">67</td>
  </tr>
  <tr>
    <td class="tdp">Leach's storm-petrel <em>(O. leucorhoa)</em></td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">5</td>
    <td class="tdr"> 1</td>
    <td class="tdr">5</td>
    <td class="tdr">5</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">5</td>
    <td class="tdr">3</td>
    <td class="tdr">0</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">5</td>
    <td class="tdr">5</td>
    <td class="tdr">5</td>
    <td class="tdr">5</td>
    <td class="tdr">63 <span class="pagenum"><a name="Page_230" id="Page_230">[Pg 230]</a></span></td>
  </tr>
  <tr>
    <td colspan="0">Pelecanidae</td>
  </tr>
  <tr>
    <td class="tdp">Brown pelican <em>(Pelecanus occidentalis)</em></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdr">1</td>
    <td class="tdr"> 1</td>
  </tr>
  <tr>
    <td colspan="0">Phalacrocoracidae</td>
  </tr>
  <tr>
    <td class="tdp">Double-crested cormorant <em>(Phalacrocorax auritus)</em></td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr"> 3</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">5</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">0</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">5</td>
    <td class="tdr">3</td>
    <td></td>
    <td class="tdr">3</td>
    <td class="tdr">5</td>
    <td class="tdr"> 52</td>
  </tr>
  <tr>
    <td class="tdp">Brandt's cormorant <em>(P. penicillatus)</em></td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">5</td>
    <td class="tdr"> 3</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">5</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">0</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">5</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr"> 57</td>
  </tr>
  <tr>
    <td class="tdp">Pelagic cormorant <em>(P. pelagicus)</em></td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">5</td>
    <td class="tdr"> 3</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">5</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">0</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">5</td>
    <td class="tdr">5</td>
    <td class="tdr">1</td>
    <td class="tdr">5</td>
    <td class="tdr">5</td>
    <td class="tdr"> 63</td>
  </tr>
  <tr>
    <td class="tdp">Red-faced cormorant <em>(P. urile)</em></td>
    <td class="tdr">5</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">5</td>
    <td class="tdr"> 3</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">5</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">0</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">5</td>
    <td class="tdr">5</td>
    <td class="tdr">5</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr"> 63</td>
  </tr>
  <tr>
    <td colspan="0">Ardeidae</td>
  </tr>
  <tr>
    <td class="tdp">Great blue heron <em>(Ardea herodias)</em></td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr"> 3</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">0</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr"> 29</td>
  </tr>
  <tr>
    <td colspan="0">Anatidae</td>
  </tr>
  <tr>
    <td class="tdp">Whooper swan <em>(Olor cygnus)</em></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdr">1</td>
    <td class="tdr"> 1</td>
  </tr>
  <tr>
    <td class="tdp">Whistling swan <em>(O. columbianus)</em></td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr"> 3</td>
    <td class="tdr">3</td>
    <td class="tdr">5</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">5</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">0</td>
    <td class="tdr">3</td>
    <td class="tdr">0</td>
    <td class="tdr"> 50</td>
  </tr>
  <tr>
    <td class="tdp">Trumpeter swan <em>(O. buccinator)</em></td>
    <td class="tdr">5</td>
    <td class="tdr">5</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr"> 5</td>
    <td class="tdr">3</td>
    <td class="tdr">5</td>
    <td class="tdr">5</td>
    <td class="tdr">1</td>
    <td class="tdr">5</td>
    <td class="tdr">1</td>
    <td class="tdr">5</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">0</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr"> 63</td>
  </tr>
  <tr>
    <td class="tdp">Canada goose <em>(Branta canadensis)</em></td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr"> 5</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">5</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr"> 34</td>
  </tr>
  <tr>
    <td class="tdp">Black Brant <em>(B. nigricans)</em></td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">5</td>
    <td class="tdr"> 3</td>
    <td class="tdr">3</td>
    <td class="tdr">5</td>
    <td class="tdr">5</td>
    <td class="tdr">3</td>
    <td class="tdr">5</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">5</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">5</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">5</td>
    <td class="tdr">3</td>
    <td class="tdr"> 70</td>
  </tr>
  <tr>
    <td class="tdp">Emperor goose <em>(Philacte canagica)</em></td>
    <td class="tdr">3</td>
    <td class="tdr">5</td>
    <td class="tdr">5</td>
    <td class="tdr">5</td>
    <td class="tdr"> 3</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">5</td>
    <td class="tdr">5</td>
    <td class="tdr">3</td>
    <td class="tdr">5</td>
    <td class="tdr">5</td>
    <td class="tdr"> 70</td>
  </tr>
  <tr>
    <td class="tdp">White-fronted goose <em>(Anser albifrons)</em></td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr"> 3</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">5</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr"> 36</td>
  </tr>
  <tr>
    <td class="tdp">Snow goose <em>(Chen hyperborea)</em></td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr"> 3</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">5</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr"> 32</td>
  </tr>
  <tr>
    <td class="tdp">Mallard <em>(Anas platyrhynchos)</em></td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr"> 1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">5</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr"> 36</td>
  </tr>
  <tr>
    <td class="tdp">Gadwall <em>(A. strepera)</em></td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr"> 1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">5</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr"> 38</td>
  </tr>
  <tr>
    <td class="tdp">Pintail <em>(A. acuta)</em></td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr"> 1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">5</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr"> 36</td>
  </tr>
  <tr>
    <td class="tdp">Common teal <em>(A. crecca)</em></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdr">1</td>
    <td></td>
    <td></td>
    <td class="tdr"> 1</td>
  </tr>
  <tr>
    <td class="tdp">Green-winged teal <em>(A. carolinensis)</em></td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr"> 1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">5</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr"> 34</td>
  </tr>
  <tr>
    <td class="tdp">Blue-winged teal <em>(A. discors)</em></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdr">1</td>
    <td></td>
    <td></td>
    <td class="tdr"> 1</td>
  </tr>
  <tr>
    <td class="tdp">Cinnamon teal <em>(A. cyanoptera)</em></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdr">1</td>
    <td></td>
    <td></td>
    <td class="tdr"> 1</td>
  </tr>
  <tr>
    <td class="tdp">European wigeon <em>(Mareca penelope)</em></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdr">1</td>
    <td></td>
    <td></td>
    <td class="tdr"> 1</td>
  </tr>
  <tr>
    <td class="tdp">American wigeon <em>(M. americana)</em></td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr"> 1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">5</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr"> 36</td>
  </tr>
  <tr>
    <td class="tdp">Shoveler <em>(Spatula clypeata)</em></td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr"> 1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">5</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr"> 34</td>
  </tr>
  <tr>
    <td class="tdp">Redhead <em>(Aythya americana)</em></td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr"> 5</td>
    <td class="tdr">3</td>
    <td class="tdr">5</td>
    <td class="tdr">5</td>
    <td class="tdr">5</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">5</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr"> 52</td>
  </tr>
  <tr>
    <td class="tdp">Ring-necked duck <em>(A. collaris)</em></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdr">1</td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdr"> 1</td>
  </tr>
  <tr>
    <td class="tdp">Canvasback <em>(A. valisineria)</em></td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr"> 5</td>
    <td class="tdr">3</td>
    <td class="tdr">5</td>
    <td class="tdr">5</td>
    <td class="tdr">5</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">5</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr"> 52</td>
  </tr>
  <tr>
    <td class="tdp">Greater scaup <em>(A. marila)</em></td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">5</td>
    <td class="tdr"> 1</td>
    <td class="tdr">3</td>
    <td class="tdr">5</td>
    <td class="tdr">5</td>
    <td class="tdr">5</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">5</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr"> 52</td>
  </tr>
  <tr>
    <td class="tdp">Lesser scaup <em>(A. affinis)</em></td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr"> 1</td>
    <td class="tdr">3</td>
    <td class="tdr">5</td>
    <td class="tdr">5</td>
    <td class="tdr">5</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">5</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr"> 50 <span class="pagenum"><a name="Page_231" id="Page_231">[Pg 231]</a></span></td>
  </tr>
  <tr>
    <td class="tdp">Common goldeneye <em>(Bucephala clangula)</em></td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr"> 1</td>
    <td class="tdr">3</td>
    <td class="tdr">5</td>
    <td class="tdr">5</td>
    <td class="tdr">5</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr"> 48</td>
  </tr>
  <tr>
    <td class="tdp">Barrow's goldeneye <em>(B. islandica)</em></td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr"> 1</td>
    <td class="tdr">3</td>
    <td class="tdr">5</td>
    <td class="tdr">5</td>
    <td class="tdr">5</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr"> 56</td>
  </tr>
  <tr>
    <td class="tdp">Bufflehead <em>(B. albeola)</em></td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr"> 1</td>
    <td class="tdr">3</td>
    <td class="tdr">5</td>
    <td class="tdr">5</td>
    <td class="tdr">5</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr"> 52</td>
  </tr>
  <tr>
    <td class="tdp">Oldsquaw <em>(Clangula hyemalis)</em></td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">5</td>
    <td class="tdr"> 1</td>
    <td class="tdr">3</td>
    <td class="tdr">5</td>
    <td class="tdr">5</td>
    <td class="tdr">5</td>
    <td class="tdr">5</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">5</td>
    <td class="tdr">5</td>
    <td class="tdr">3</td>
    <td class="tdr">5</td>
    <td class="tdr">5</td>
    <td class="tdr"> 66</td>
  </tr>
  <tr>
    <td class="tdp">Harlequin duck <em>(Histrionicus histrionicus)</em></td>
    <td class="tdr">3</td>
    <td class="tdr">5</td>
    <td class="tdr">1</td>
    <td class="tdr">5</td>
    <td class="tdr"> 1</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">5</td>
    <td class="tdr">5</td>
    <td class="tdr">5</td>
    <td class="tdr">5</td>
    <td class="tdr">5</td>
    <td class="tdr"> 60</td>
  </tr>
  <tr>
    <td class="tdp">Steller's eider <em>(Polysticta stelleri)</em></td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">5</td>
    <td class="tdr">5</td>
    <td class="tdr"> 1</td>
    <td class="tdr">3</td>
    <td class="tdr">5</td>
    <td class="tdr">5</td>
    <td class="tdr">5</td>
    <td class="tdr">5</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">5</td>
    <td class="tdr">5</td>
    <td class="tdr">3</td>
    <td class="tdr">5</td>
    <td class="tdr">5</td>
    <td class="tdr"> 72</td>
  </tr>
  <tr>
    <td class="tdp">Common eider <em>(Somateria mollissima)</em></td>
    <td class="tdr">3</td>
    <td class="tdr">5</td>
    <td class="tdr">3</td>
    <td class="tdr">5</td>
    <td class="tdr"> 1</td>
    <td class="tdr">3</td>
    <td class="tdr">5</td>
    <td class="tdr">5</td>
    <td class="tdr">5</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">5</td>
    <td class="tdr">5</td>
    <td class="tdr">3</td>
    <td class="tdr">5</td>
    <td class="tdr">5</td>
    <td class="tdr"> 68</td>
  </tr>
  <tr>
    <td class="tdp">King eider <em>(S. spectabilis)</em></td>
    <td class="tdr">3</td>
    <td class="tdr">5</td>
    <td class="tdr">3</td>
    <td class="tdr">5</td>
    <td class="tdr"> 1</td>
    <td class="tdr">3</td>
    <td class="tdr">5</td>
    <td class="tdr">5</td>
    <td class="tdr">5</td>
    <td class="tdr">5</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">5</td>
    <td class="tdr">5</td>
    <td class="tdr">3</td>
    <td class="tdr">5</td>
    <td class="tdr">5</td>
    <td class="tdr"> 70</td>
  </tr>
  <tr>
    <td class="tdp">Spectacled eider <em>(Lampronetta fisheri)</em></td>
    <td class="tdr">5</td>
    <td class="tdr">5</td>
    <td class="tdr">5</td>
    <td class="tdr">5</td>
    <td class="tdr"> 3</td>
    <td class="tdr">3</td>
    <td class="tdr">5</td>
    <td class="tdr">5</td>
    <td class="tdr">5</td>
    <td class="tdr">5</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">5</td>
    <td class="tdr">5</td>
    <td class="tdr">3</td>
    <td class="tdr">5</td>
    <td class="tdr">5</td>
    <td class="tdr"> 78</td>
  </tr>
  <tr>
    <td class="tdp">White-winged scoter <em>(Melanitta deglandi)</em></td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr"> 1</td>
    <td class="tdr">3</td>
    <td class="tdr">5</td>
    <td class="tdr">5</td>
    <td class="tdr">5</td>
    <td class="tdr">5</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">5</td>
    <td class="tdr">5</td>
    <td class="tdr">5</td>
    <td class="tdr">5</td>
    <td class="tdr">5</td>
    <td class="tdr"> 72</td>
  </tr>
  <tr>
    <td class="tdp">Surf scoter <em>(M. perspicillata)</em></td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr"> 1</td>
    <td class="tdr">3</td>
    <td class="tdr">5</td>
    <td class="tdr">5</td>
    <td class="tdr">5</td>
    <td class="tdr">5</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">5</td>
    <td class="tdr">5</td>
    <td class="tdr">5</td>
    <td class="tdr">5</td>
    <td class="tdr">5</td>
    <td class="tdr"> 72</td>
  </tr>
  <tr>
    <td class="tdp">Common scoter <em>(Oidemia nigra)</em></td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr"> 1</td>
    <td class="tdr">3</td>
    <td class="tdr">5</td>
    <td class="tdr">5</td>
    <td class="tdr">5</td>
    <td class="tdr">5</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">5</td>
    <td class="tdr">5</td>
    <td class="tdr">5</td>
    <td class="tdr">5</td>
    <td class="tdr">5</td>
    <td class="tdr"> 72</td>
  </tr>
  <tr>
    <td class="tdp">Ruddy duck <em>(Oxyura jamaicensis)</em></td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr"> 1</td>
    <td class="tdr">1</td>
    <td class="tdr">5</td>
    <td class="tdr">5</td>
    <td class="tdr">5</td>
    <td class="tdr">5</td>
    <td class="tdr">1</td>
    <td class="tdr">5</td>
    <td class="tdr">5</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">0</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr"> 55</td>
  </tr>
  <tr>
    <td class="tdp">Hooded merganser <em>(Laphodytes cucullatus)</em></td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr"> 3</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">5</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">0</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr"> 37</td>
  </tr>
  <tr>
    <td class="tdp">Common merganser <em>(Mergus merganser)</em></td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr"> 1</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">5</td>
    <td class="tdr">5</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr"> 56</td>
  </tr>
  <tr>
    <td class="tdp">Red-breasted merganser <em>(M. serrator)</em></td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr"> 1</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">5</td>
    <td class="tdr">5</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr"> 56</td>
  </tr>
  <tr>
    <td colspan="0">Accipitridae</td>
  </tr>
  <tr>
    <td class="tdp">Bald eagle <em>(Haliaeetus leucocephalus)</em></td>
    <td class="tdr">1</td>
    <td class="tdr">5</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr"> 5</td>
    <td class="tdr">5</td>
    <td class="tdr">0</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">0</td>
    <td class="tdr">1</td>
    <td class="tdr">5</td>
    <td class="tdr">0</td>
    <td class="tdr">0</td>
    <td class="tdr">5</td>
    <td class="tdr">3</td>
    <td class="tdr">5</td>
    <td class="tdr">5</td>
    <td class="tdr">5</td>
    <td class="tdr">5</td>
    <td class="tdr"> 58</td>
  </tr>
  <tr>
    <td class="tdp">Steller's sea eagle <em>(H. pelagicus)</em></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdr">1</td>
    <td></td>
    <td></td>
    <td class="tdr"> 1</td>
  </tr>
  <tr>
    <td class="tdp">Marsh hawk <em>(Circus cyaneus)</em></td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr"> 1</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">0</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">0</td>
    <td class="tdr">0</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">0</td>
    <td class="tdr">1</td>
    <td class="tdr">0</td>
    <td class="tdr"> 19</td>
  </tr>
  <tr>
    <td colspan="0">Pandionidae</td>
  </tr>
  <tr>
    <td class="tdp">Osprey <em>(Pandion haliaetus)</em></td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr"> 5</td>
    <td class="tdr">5</td>
    <td class="tdr">0</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">0</td>
    <td class="tdr">1</td>
    <td class="tdr">5</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">5</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">0</td>
    <td class="tdr"> 37</td>
  </tr>
  <tr>
    <td colspan="0">Falconidae</td>
  </tr>
  <tr>
    <td class="tdp">Peregrine falcon <em>(Falco peregrinus)</em></td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr"> 5</td>
    <td class="tdr">5</td>
    <td class="tdr">0</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">0</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">0</td>
    <td class="tdr">5</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr"> 41</td>
  </tr>
  <tr>
    <td colspan="0">Gruidae</td>
  </tr>
  <tr>
    <td class="tdp">Sandhill crane <em>(Grus canadensis)</em></td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr"> 1</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">0</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">0</td>
    <td class="tdr"> 24</td>
  </tr>
  <tr>
    <td colspan="0">Rallidae</td>
  </tr>
  <tr>
    <td class="tdp">American coot <em>(Fulica americana)</em></td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr"> 1</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">0</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr"> 33</td>
  </tr>
  <tr>
    <td colspan="0">Haematopodidae</td>
  </tr>
  <tr>
    <td class="tdp">Black oystercatcher <em>(Haematopus bachmani)</em></td>
    <td class="tdr">5</td>
    <td class="tdr">5</td>
    <td class="tdr">5</td>
    <td class="tdr">5</td>
    <td class="tdr"> 3</td>
    <td class="tdr">5</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">5</td>
    <td class="tdr">0</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">5</td>
    <td class="tdr">5</td>
    <td class="tdr">5</td>
    <td class="tdr">5</td>
    <td class="tdr"> 65</td>
  </tr>
  <tr>
    <td colspan="0">Charadriidae</td>
  </tr>
  <tr>
    <td class="tdp">Ringed plover <em>(Charadrius hiaticula)</em></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdr">1</td>
    <td></td>
    <td></td>
    <td class="tdr"> 1</td>
  </tr>
  <tr>
    <td class="tdp">Semipalmated plover <em>(C. semipalmatus)</em></td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr"> 1</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">0</td>
    <td class="tdr">5</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">0</td>
    <td class="tdr"> 28</td>
  </tr>
  <tr>
    <td class="tdp">Mongolian plover <em>(C. mongolus)</em></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdr">1</td>
    <td class="tdr"> 1</td>
  </tr>
  <tr>
    <td class="tdp">Killdeer <em>(C. vociferus)</em></td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr"> 1</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">0</td>
    <td class="tdr">5</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">0</td>
    <td class="tdr"> 26 <span class="pagenum"><a name="Page_232" id="Page_232">[Pg 232]</a></span></td>
  </tr>
  <tr>
    <td class="tdp">Dotterel <em>(Eudromias morinellus)</em></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdr">1</td>
    <td></td>
    <td></td>
    <td class="tdr"> 1</td>
  </tr>
  <tr>
    <td class="tdp">American golden plover <em>(Pluvialis dominica)</em></td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr"> 3</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">5</td>
    <td class="tdr">0</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">0</td>
    <td class="tdr">1</td>
    <td class="tdr">0</td>
    <td class="tdr"> 35</td>
  </tr>
  <tr>
    <td class="tdp">Black-bellied plover <em>(Squatarola squatarola)</em></td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">5</td>
    <td class="tdr"> 3</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">5</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">0</td>
    <td class="tdr"> 43</td>
  </tr>
  <tr>
    <td class="tdp">Surfbird <em>(Aphriza virgata)</em></td>
    <td class="tdr">5</td>
    <td class="tdr">1</td>
    <td class="tdr">5</td>
    <td class="tdr">5</td>
    <td class="tdr"> 3</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">0</td>
    <td class="tdr">5</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">5</td>
    <td class="tdr">0</td>
    <td class="tdr">5</td>
    <td class="tdr">3</td>
    <td class="tdr"> 54</td>
  </tr>
  <tr>
    <td class="tdp">Ruddy turnstone <em>(Arenaria interpres)</em></td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">5</td>
    <td class="tdr"> 3</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">0</td>
    <td class="tdr">5</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">0</td>
    <td class="tdr"> 44</td>
  </tr>
  <tr>
    <td class="tdp">Black turnstone <em>(A. melanocephala)</em></td>
    <td class="tdr">5</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">5</td>
    <td class="tdr"> 3</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">0</td>
    <td class="tdr">5</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">5</td>
    <td class="tdr">3</td>
    <td class="tdr">5</td>
    <td class="tdr">3</td>
    <td class="tdr"> 57</td>
  </tr>
  <tr>
    <td colspan="0">Scolopacidae</td>
  </tr>
  <tr>
    <td class="tdp">Common snipe <em>(Capella gallinago)</em></td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr"> 1</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">5</td>
    <td class="tdr">5</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">0</td>
    <td class="tdr"> 29</td>
  </tr>
  <tr>
    <td class="tdp">Eurasian curlew <em>(Numenius arquata)</em></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdr">1</td>
    <td></td>
    <td></td>
    <td class="tdr"> 1</td>
  </tr>
  <tr>
    <td class="tdp">Whimbrel <em>(N. phaeopus)</em></td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr"> 3</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">0</td>
    <td class="tdr"> 37</td>
  </tr>
  <tr>
    <td class="tdp">Bristle-thighed curlew <em>(N. tahitiensis)</em></td>
    <td class="tdr">5</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">5</td>
    <td class="tdr"> 5</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">0</td>
    <td class="tdr"> 45</td>
  </tr>
  <tr>
    <td class="tdp">Eskimo curlew <em>(N. borealis)</em></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdr">99</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdr">1</td>
    <td></td>
    <td></td>
    <td class="tdr">100</td>
  </tr>
  <tr>
    <td class="tdp">Upland plover <em>(Bartramia longicauda)</em></td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">0</td>
    <td class="tdr"> 5</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">0</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">0</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">0</td>
    <td class="tdr"> 26</td>
  </tr>
  <tr>
    <td class="tdp">Spotted sandpiper <em>(Actitis macularia)</em></td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr"> 1</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">0</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">0</td>
    <td class="tdr"> 24</td>
  </tr>
  <tr>
    <td class="tdp">Common sandpiper <em>(Tringa hypoleucos)</em></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdr">1</td>
    <td></td>
    <td></td>
    <td class="tdr"> 1</td>
  </tr>
  <tr>
    <td class="tdp">Solitary sandpiper <em>(T. solitaria)</em></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdr">1</td>
    <td></td>
    <td></td>
    <td class="tdr"> 1</td>
  </tr>
  <tr>
    <td class="tdp">Wood sandpiper <em>(T. glareola)</em></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdr">1</td>
    <td></td>
    <td></td>
    <td class="tdr"> 1</td>
  </tr>
  <tr>
    <td class="tdp">Wandering tattler <em>(Heteroscelus incanum)</em></td>
    <td class="tdr">5</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">5</td>
    <td class="tdr"> 5</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">5</td>
    <td class="tdr">0</td>
    <td class="tdr">5</td>
    <td class="tdr">0</td>
    <td class="tdr"> 48</td>
  </tr>
  <tr>
    <td class="tdp">Polynesian tattler <em>(H. brevipes)</em></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdr">1</td>
    <td></td>
    <td class="tdr"> 1</td>
  </tr>
  <tr>
    <td class="tdp">Willet <em>(Catoptrophorus semipalmatus)</em></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdr">1</td>
    <td></td>
    <td></td>
    <td class="tdr"> 1</td>
  </tr>
  <tr>
    <td class="tdp">Greater yellowlegs <em>(Totanus melanoleucus)</em></td>
    <td class="tdr">1</td>
    <td class="tdr">5</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr"> 3</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">0</td>
    <td class="tdr">1</td>
    <td class="tdr">0</td>
    <td class="tdr"> 30</td>
  </tr>
  <tr>
    <td class="tdp">Lesser yellowlegs <em>(T. flavipes)</em></td>
    <td class="tdr">1</td>
    <td class="tdr">5</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr"> 3</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">0</td>
    <td class="tdr">1</td>
    <td class="tdr">0</td>
    <td class="tdr"> 30</td>
  </tr>
  <tr>
    <td class="tdp">Spotted redshank <em>(T. totanus)</em></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdr">1</td>
    <td></td>
    <td></td>
    <td class="tdr"> 1</td>
  </tr>
  <tr>
    <td class="tdp">Greenshank <em>(Tringa nebularia)</em></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdr">1</td>
    <td></td>
    <td></td>
    <td class="tdr"> 1</td>
  </tr>
  <tr>
    <td class="tdp">Knot <em>(Calidris canutus)</em></td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">5</td>
    <td class="tdr"> 5</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">0</td>
    <td class="tdr"> 39</td>
  </tr>
  <tr>
    <td class="tdp">Great knot <em>(C. tenuirostris)</em></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdr">1</td>
    <td></td>
    <td></td>
    <td class="tdr"> 1</td>
  </tr>
  <tr>
    <td class="tdp">Rock sandpiper <em>(Erolia ptilocnemis)</em></td>
    <td class="tdr">5</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">5</td>
    <td class="tdr"> 3</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">0</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">5</td>
    <td class="tdr">5</td>
    <td class="tdr">5</td>
    <td class="tdr">5</td>
    <td class="tdr"> 59</td>
  </tr>
  <tr>
    <td class="tdp">Sharp-tailed sandpiper <em>(E. acuminata)</em></td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">5</td>
    <td class="tdr"> 3</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">0</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">0</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr"> 46</td>
  </tr>
  <tr>
    <td class="tdp">Pectoral sandpiper <em>(E. melanotos)</em></td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr"> 3</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">0</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">0</td>
    <td class="tdr"> 32</td>
  </tr>
  <tr>
    <td class="tdp">White-rumped sandpiper <em>(E. fuscicollis)</em></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdr">1</td>
    <td class="tdr"> 1</td>
  </tr>
  <tr>
    <td class="tdp">Baird sandpiper <em>(E. bairdii)</em></td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr"> 3</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">0</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">0</td>
    <td class="tdr"> 34</td>
  </tr>
  <tr>
    <td class="tdp">Least sandpiper <em>(E. minutilla)</em></td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr"> 1</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">0</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">0</td>
    <td class="tdr"> 34</td>
  </tr>
  <tr>
    <td class="tdp">Long-toed stint <em>(E. subminuta)</em></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdr">1</td>
    <td></td>
    <td></td>
    <td class="tdr"> 1</td>
  </tr>
  <tr>
    <td class="tdp">Temminck's stint <em>(Calidrus temminckii)</em></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdr">1</td>
    <td></td>
    <td></td>
    <td class="tdr"> 1</td>
  </tr>
  <tr>
    <td class="tdp">Rufous-necked sandpiper <em>(E. ruficollis)</em></td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">5</td>
    <td class="tdr"> 3</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">0</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">0</td>
    <td class="tdr"> 36</td>
  </tr>
  <tr>
    <td class="tdp">Curlew sandpiper <em>(E. ferruginea)</em></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdr">1</td>
    <td></td>
    <td></td>
    <td class="tdr"> 1 <span class="pagenum"><a name="Page_233" id="Page_233">[Pg 233]</a></span></td>
  </tr>
  <tr>
    <td class="tdp">Dunlin <em>(E. alpina)</em></td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">5</td>
    <td class="tdr"> 1</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">0</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr"> 41</td>
  </tr>
  <tr>
    <td class="tdp">Short-billed dowitcher <em>(Limnodromus griseus)</em></td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr"> 3</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">0</td>
    <td class="tdr"> 45</td>
  </tr>
  <tr>
    <td class="tdp">Long-billed dowitcher <em>(L. scolopaceus)</em></td>
    <td class="tdr">5</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr"> 3</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">0</td>
    <td class="tdr"> 47</td>
  </tr>
  <tr>
    <td class="tdp">Stilt sandpiper <em>(Micropalama himantopus)</em></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdr">1</td>
    <td></td>
    <td></td>
    <td class="tdr"> 1</td>
  </tr>
  <tr>
    <td class="tdp">Semipalmated sandpiper <em>(Ereunetes pusillus)</em></td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr"> 1</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">0</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">5</td>
    <td class="tdr">3</td>
    <td class="tdr">0</td>
    <td class="tdr"> 34</td>
  </tr>
  <tr>
    <td class="tdp">Western sandpiper <em>(E. mauri)</em></td>
    <td class="tdr">5</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">5</td>
    <td class="tdr"> 1</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">0</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">5</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr"> 47</td>
  </tr>
  <tr>
    <td class="tdp">Buff-breasted sandpiper <em>(Tryngites subruficollis)</em></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdr">1</td>
    <td></td>
    <td></td>
    <td class="tdr"> 1</td>
  </tr>
  <tr>
    <td class="tdp">Marbled godwit <em>(Limosa fedoa)</em></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdr">1</td>
    <td></td>
    <td></td>
    <td class="tdr"> 1</td>
  </tr>
  <tr>
    <td class="tdp">Bar-tailed godwit <em>(L. lapponica)</em></td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">5</td>
    <td class="tdr"> 3</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">5</td>
    <td class="tdr">5</td>
    <td class="tdr">3</td>
    <td class="tdr">0</td>
    <td class="tdr"> 49</td>
  </tr>
  <tr>
    <td class="tdp">Hudsonian godwit <em>(L. haemastica)</em></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdr">1</td>
    <td></td>
    <td></td>
    <td class="tdr"> 1</td>
  </tr>
  <tr>
    <td class="tdp">Black-tailed godwit <em>(L. limosa)</em></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdr">1</td>
    <td></td>
    <td></td>
    <td class="tdr"> 1</td>
  </tr>
  <tr>
    <td class="tdp">Ruff <em>(Philomachus pugnax)</em></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdr">1</td>
    <td></td>
    <td></td>
    <td class="tdr"> 1</td>
  </tr>
  <tr>
    <td class="tdp">Sanderling <em>(Crocethia alba)</em></td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">5</td>
    <td class="tdr"> 3</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">0</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr"> 45</td>
  </tr>
  <tr>
    <td class="tdp">Spoon-billed sandpiper <em>(Eurynorhynchus pygmeum)</em></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdr">1</td>
    <td></td>
    <td></td>
    <td class="tdr"> 1</td>
  </tr>
  <tr>
    <td colspan="0">Phalaropodidae</td>
  </tr>
  <tr>
    <td class="tdp">Red phalarope <em>(Phalaropus fulicarius)</em></td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">5</td>
    <td class="tdr"> 1</td>
    <td class="tdr">3</td>
    <td class="tdr">5</td>
    <td class="tdr">5</td>
    <td class="tdr">1</td>
    <td class="tdr">5</td>
    <td class="tdr">1</td>
    <td class="tdr">5</td>
    <td class="tdr">0</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">5</td>
    <td class="tdr">5</td>
    <td class="tdr">3</td>
    <td class="tdr">5</td>
    <td class="tdr">0</td>
    <td class="tdr"> 58</td>
  </tr>
  <tr>
    <td class="tdp">Wilson's phalarope <em>(Steganopus tricolor)</em></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdr">1</td>
    <td></td>
    <td class="tdr"> 1</td>
  </tr>
  <tr>
    <td class="tdp">Northern phalarope <em>(Lobipes lobatus)</em></td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">5</td>
    <td class="tdr"> 1</td>
    <td class="tdr">3</td>
    <td class="tdr">5</td>
    <td class="tdr">5</td>
    <td class="tdr">1</td>
    <td class="tdr">5</td>
    <td class="tdr">1</td>
    <td class="tdr">5</td>
    <td class="tdr">0</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">5</td>
    <td class="tdr">5</td>
    <td class="tdr">3</td>
    <td class="tdr">5</td>
    <td class="tdr">0</td>
    <td class="tdr"> 62</td>
  </tr>
  <tr>
    <td colspan="0">Stercorariidae</td>
  </tr>
  <tr>
    <td class="tdp">Pomarine jaeger <em>(Stercorarius pomarinus)</em></td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">5</td>
    <td class="tdr"> 1</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">0</td>
    <td class="tdr"> 41</td>
  </tr>
  <tr>
    <td class="tdp">Parasitic jaeger <em>(S. parasiticus)</em></td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">5</td>
    <td class="tdr"> 1</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">0</td>
    <td class="tdr"> 43</td>
  </tr>
  <tr>
    <td class="tdp">Long-tailed jaeger <em>(S. longicaudus)</em></td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr"> 1</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">0</td>
    <td class="tdr"> 39</td>
  </tr>
  <tr>
    <td class="tdp">Skua <em>(Catharacta skua)</em></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdr">1</td>
    <td class="tdr"> 1</td>
  </tr>
  <tr>
    <td colspan="0">Laridae</td>
  </tr>
  <tr>
    <td class="tdp">Glaucous gull <em>(Larus hyperboreus)</em></td>
    <td class="tdr">1</td>
    <td class="tdr">5</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr"> 1</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">0</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr"> 45</td>
  </tr>
  <tr>
    <td class="tdp">Glaucous-winged gull <em>(L. glaucescens)</em></td>
    <td class="tdr">5</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">5</td>
    <td class="tdr"> 1</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">5</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">5</td>
    <td class="tdr">5</td>
    <td class="tdr">5</td>
    <td class="tdr">3</td>
    <td class="tdr"> 56</td>
  </tr>
  <tr>
    <td class="tdp">Slaty-backed gull <em>(L. schistisagus)</em></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdr">1</td>
    <td class="tdr"> 1</td>
  </tr>
  <tr>
    <td class="tdp">Western gull <em>(L. occidentalis)</em></td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">5</td>
    <td class="tdr"> 1</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">5</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr"> 48</td>
  </tr>
  <tr>
    <td class="tdp">Herring gull <em>(L. argentatus)</em></td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr"> 1</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr"> 38</td>
  </tr>
  <tr>
    <td class="tdp">Thayer's gull <em>(L. thayeri)</em></td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">5</td>
    <td class="tdr">3</td>
    <td class="tdr"> 1</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr"> 42</td>
  </tr>
  <tr>
    <td class="tdp">California gull <em>(L. californicus)</em></td>
    <td class="tdr">3</td>
    <td class="tdr">5</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr"> 1</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr"> 38</td>
  </tr>
  <tr>
    <td class="tdp">Ring-billed gull <em>(L. delawarensis)</em></td>
    <td class="tdr">1</td>
    <td class="tdr">5</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr"> 1</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr"> 36</td>
  </tr>
  <tr>
    <td class="tdp">Mew gull <em>(L. canus)</em></td>
    <td class="tdr">1</td>
    <td class="tdr">5</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr"> 1</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">1</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr">3</td>
    <td class="tdr"> 44</td>
  </tr>
  <tr>
    <td class="tdp">Black-headed gull <em>(L. ridibundus)</em></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdr">1</td>
    <td></td>
    <td></td>
    <td class="tdr"> 1</td>
  </tr>
  <tr>
    <td class="tdp">Franklin's gull <em>(L. pipixcan)</em></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdr">1</td>
    <td></td>
    <td></td>
    <td class="tdr"> 1<span class="pagenum"><a name="Page_234" id="Page_234">[Pg 234]</a></span></td>
  </tr>
  <tr>
    <td class="tdp">Bonaparte's gull <em>(L. philadelphia)</em></td>
    <td class="tdr"> 1</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 3</td>
    <td class="tdr"> 3</td>
    <td class="tdr"> 1</td>
    <td class="tdr"> 3</td>
    <td class="tdr"> 3</td>
    <td class="tdr"> 3</td>
    <td class="tdr"> 1</td>
    <td class="tdr"> 3</td>
    <td class="tdr"> 1</td>
    <td class="tdr"> 1</td>
    <td class="tdr"> 1</td>
    <td class="tdr"> 1</td>
    <td class="tdr"> 1</td>
    <td class="tdr"> 1</td>
    <td class="tdr"> 3</td>
    <td class="tdr"> 1</td>
    <td class="tdr"> 3</td>
    <td class="tdr"> 1</td>
    <td class="tdr"> 40</td>
  </tr>
  <tr>
    <td class="tdp">Heerman's gull <em>(L. heermanni)</em></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdr"> 1</td>
    <td class="tdr"> 1</td>
  </tr>
  <tr>
    <td class="tdp">Ivory gull <em>(Pagophila eburnea)</em></td>
    <td class="tdr"> 1</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 3</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 3</td>
    <td class="tdr"> 3</td>
    <td class="tdr"> 3</td>
    <td class="tdr"> 3</td>
    <td class="tdr"> 1</td>
    <td class="tdr"> 3</td>
    <td class="tdr"> 1</td>
    <td class="tdr"> 3</td>
    <td class="tdr"> 0</td>
    <td class="tdr"> 1</td>
    <td class="tdr"> 1</td>
    <td class="tdr"> 1</td>
    <td class="tdr"> 1</td>
    <td class="tdr"> 1</td>
    <td class="tdr"> 1</td>
    <td class="tdr"> 3</td>
    <td class="tdr"> 43</td>
  </tr>
  <tr>
    <td class="tdp">Black-legged kittiwake <em>(Rissa tridactyla)</em></td>
    <td class="tdr"> 1</td>
    <td class="tdr"> 3</td>
    <td class="tdr"> 3</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 1</td>
    <td class="tdr"> 3</td>
    <td class="tdr"> 3</td>
    <td class="tdr"> 3</td>
    <td class="tdr"> 1</td>
    <td class="tdr"> 3</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 3</td>
    <td class="tdr"> 0</td>
    <td class="tdr"> 1</td>
    <td class="tdr"> 1</td>
    <td class="tdr"> 1</td>
    <td class="tdr"> 3</td>
    <td class="tdr"> 3</td>
    <td class="tdr"> 3</td>
    <td class="tdr"> 3</td>
    <td class="tdr"> 49</td>
  </tr>
  <tr>
    <td class="tdp">Red-legged kittiwake <em>(R. brevirostris)</em></td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 3</td>
    <td class="tdr"> 3</td>
    <td class="tdr"> 3</td>
    <td class="tdr"> 3</td>
    <td class="tdr"> 1</td>
    <td class="tdr"> 3</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 3</td>
    <td class="tdr"> 0</td>
    <td class="tdr"> 1</td>
    <td class="tdr"> 0</td>
    <td class="tdr"> 1</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 66</td>
  </tr>
  <tr>
    <td class="tdp">Ross' gull <em>(Rhodostethia rosea)</em></td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 3</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 3</td>
    <td class="tdr"> 3</td>
    <td class="tdr"> 3</td>
    <td class="tdr"> 3</td>
    <td class="tdr"> 1</td>
    <td class="tdr"> 3</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 0</td>
    <td class="tdr"> 1</td>
    <td class="tdr"> 0</td>
    <td class="tdr"> 1</td>
    <td class="tdr"> 3</td>
    <td class="tdr"> 1</td>
    <td class="tdr"> 3</td>
    <td class="tdr"> 3</td>
    <td class="tdr"> 56</td>
  </tr>
  <tr>
    <td class="tdp">Sabine's gull <em>(Xema sabini)</em></td>
    <td class="tdr"> 3</td>
    <td class="tdr"> 3</td>
    <td class="tdr"> 3</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 1</td>
    <td class="tdr"> 3</td>
    <td class="tdr"> 3</td>
    <td class="tdr"> 3</td>
    <td class="tdr"> 1</td>
    <td class="tdr"> 3</td>
    <td class="tdr"> 1</td>
    <td class="tdr"> 3</td>
    <td class="tdr"> 0</td>
    <td class="tdr"> 1</td>
    <td class="tdr"> 1</td>
    <td class="tdr"> 1</td>
    <td class="tdr"> 3</td>
    <td class="tdr"> 3</td>
    <td class="tdr"> 3</td>
    <td class="tdr"> 0</td>
    <td class="tdr"> 44</td>
  </tr>
  <tr>
    <td class="tdp">Common tern <em>(Sterna hirundo)</em></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdr"> 1</td>
    <td></td>
    <td class="tdr"> 1</td>
  </tr>
  <tr>
    <td class="tdp">Arctic tern <em>(S. paradisaea)</em></td>
    <td class="tdr"> 1</td>
    <td class="tdr"> 1</td>
    <td class="tdr"> 1</td>
    <td class="tdr"> 3</td>
    <td class="tdr"> 1</td>
    <td class="tdr"> 3</td>
    <td class="tdr"> 3</td>
    <td class="tdr"> 3</td>
    <td class="tdr"> 1</td>
    <td class="tdr"> 3</td>
    <td class="tdr"> 1</td>
    <td class="tdr"> 1</td>
    <td class="tdr"> 0</td>
    <td class="tdr"> 1</td>
    <td class="tdr"> 1</td>
    <td class="tdr"> 1</td>
    <td class="tdr"> 3</td>
    <td class="tdr"> 1</td>
    <td class="tdr"> 3</td>
    <td class="tdr"> 0</td>
    <td class="tdr"> 32</td>
  </tr>
  <tr>
    <td class="tdp">Aleutian tern <em>(S. aleutica)</em></td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 3</td>
    <td class="tdr"> 3</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 3</td>
    <td class="tdr"> 3</td>
    <td class="tdr"> 3</td>
    <td class="tdr"> 3</td>
    <td class="tdr"> 1</td>
    <td class="tdr"> 3</td>
    <td class="tdr"> 1</td>
    <td class="tdr"> 1</td>
    <td class="tdr"> 0</td>
    <td class="tdr"> 1</td>
    <td class="tdr"> 1</td>
    <td class="tdr"> 1</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 1</td>
    <td class="tdr"> 53</td>
  </tr>
  <tr>
    <td class="tdp">Caspian tern <em>(Hydroprogne caspia)</em></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdr"> 1</td>
    <td></td>
    <td class="tdr"> 1</td>
  </tr>
  <tr>
    <td class="tdp">Black tern <em>(Chlidonias niger)</em></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdr"> 1</td>
    <td></td>
    <td class="tdr"> 1</td>
  </tr>
  <tr>
    <td colspan="0">Alcidae</td>
  </tr>
  <tr>
    <td class="tdp">Common murre <em>(Uria aalge)</em></td>
    <td class="tdr"> 1</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 3</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 1</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 3</td>
    <td class="tdr"> 1</td>
    <td class="tdr"> 1</td>
    <td class="tdr"> 3</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 3</td>
    <td class="tdr"> 3</td>
    <td class="tdr"> 3</td>
    <td class="tdr"> 3</td>
    <td class="tdr"> 70</td>
  </tr>
  <tr>
    <td class="tdp">Thick-billed murre <em>(U. lomvia)</em></td>
    <td class="tdr"> 1</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 3</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 1</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 3</td>
    <td class="tdr"> 1</td>
    <td class="tdr"> 1</td>
    <td class="tdr"> 3</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 3</td>
    <td class="tdr"> 3</td>
    <td class="tdr"> 3</td>
    <td class="tdr"> 3</td>
    <td class="tdr"> 70</td>
  </tr>
  <tr>
    <td class="tdp">Dovekie <em>(Plautus alle)</em></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdr"> 1</td>
    <td></td>
    <td></td>
    <td class="tdr"> 1</td>
  </tr>
  <tr>
    <td class="tdp">Black guillemot <em>(Cepphus grylle)</em></td>
    <td class="tdr"> 1</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 3</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 3</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 3</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 1</td>
    <td class="tdr"> 1</td>
    <td class="tdr"> 1</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 3</td>
    <td class="tdr"> 3</td>
    <td class="tdr"> 3</td>
    <td class="tdr"> 3</td>
    <td class="tdr"> 70</td>
  </tr>
  <tr>
    <td class="tdp">Pigeon guillemot <em>(C. columba)</em></td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 3</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 3</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 3</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 1</td>
    <td class="tdr"> 1</td>
    <td class="tdr"> 3</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 3</td>
    <td class="tdr"> 82</td>
  </tr>
  <tr>
    <td class="tdp">Marbled murrelet <em>(Brachyramphus marmoratus)</em></td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 3</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 1</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 3</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 1</td>
    <td class="tdr"> 3</td>
    <td class="tdr"> 3</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 84</td>
  </tr>
  <tr>
    <td class="tdp">Kittlitz's murrelet <em>(B. brevirostris)</em></td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 1</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 1</td>
    <td class="tdr"> 3</td>
    <td class="tdr"> 3</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 88</td>
  </tr>
  <tr>
    <td class="tdp">Xantus' murrelet <em>(Endomychura hypoleuca)</em></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdr"> 1</td>
    <td class="tdr"> 1</td>
  </tr>
  <tr>
    <td class="tdp">Ancient murrelet <em>(Synthliboramphus antiquus)</em></td>
    <td class="tdr"> 3</td>
    <td class="tdr"> 3</td>
    <td class="tdr"> 3</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 1</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 1</td>
    <td class="tdr"> 3</td>
    <td class="tdr"> 3</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 3</td>
    <td class="tdr"> 3</td>
    <td class="tdr"> 3</td>
    <td class="tdr"> 3</td>
    <td class="tdr"> 74</td>
  </tr>
  <tr>
    <td class="tdp">Cassin's auklet <em>(Ptychoramphus aleutica)</em></td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 3</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 1</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 1</td>
    <td class="tdr"> 3</td>
    <td class="tdr"> 3</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 3</td>
    <td class="tdr"> 84</td>
  </tr>
  <tr>
    <td class="tdp">Parakeet auklet <em>(Cyclorrhynchus psittacula)</em></td>
    <td class="tdr"> 3</td>
    <td class="tdr"> 3</td>
    <td class="tdr"> 3</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 1</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 1</td>
    <td class="tdr"> 3</td>
    <td class="tdr"> 3</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 3</td>
    <td class="tdr"> 80</td>
  </tr>
  <tr>
    <td class="tdp">Crested auklet <em>(Aethia cristatella)</em></td>
    <td class="tdr"> 3</td>
    <td class="tdr"> 3</td>
    <td class="tdr"> 3</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 1</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 1</td>
    <td class="tdr"> 3</td>
    <td class="tdr"> 1</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 3</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 3</td>
    <td class="tdr"> 76</td>
  </tr>
  <tr>
    <td class="tdp">Least auklet <em>(A. pusilla)</em></td>
    <td class="tdr"> 3</td>
    <td class="tdr"> 3</td>
    <td class="tdr"> 3</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 1</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 1</td>
    <td class="tdr"> 3</td>
    <td class="tdr"> 3</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 3</td>
    <td class="tdr"> 80</td>
  </tr>
  <tr>
    <td class="tdp">Whiskered auklet <em>(A. pygmaea)</em></td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 1</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 1</td>
    <td class="tdr"> 3</td>
    <td class="tdr"> 3</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 88</td>
  </tr>
  <tr>
    <td class="tdp">Rhinoceros auklet <em>(Cerorhinca monocerata)</em></td>
    <td class="tdr"> 3</td>
    <td class="tdr"> 3</td>
    <td class="tdr"> 3</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 1</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 1</td>
    <td class="tdr"> 3</td>
    <td class="tdr"> 3</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 3</td>
    <td class="tdr"> 3</td>
    <td class="tdr"> 3</td>
    <td class="tdr"> 3</td>
    <td class="tdr"> 74</td>
  </tr>
  <tr>
    <td class="tdp">Horned puffin <em>(Fratercula corniculata)</em></td>
    <td class="tdr"> 3</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 3</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 1</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 3</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 1</td>
    <td class="tdr"> 3</td>
    <td class="tdr"> 1</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 3</td>
    <td class="tdr"> 3</td>
    <td class="tdr"> 3</td>
    <td class="tdr"> 3</td>
    <td class="tdr"> 72</td>
  </tr>
  <tr>
    <td class="tdp">Tufted puffin <em>(Lunda cirrhata)</em></td>
    <td class="tdr"> 3</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 3</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 1</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 3</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 1</td>
    <td class="tdr"> 3</td>
    <td class="tdr"> 1</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 3</td>
    <td class="tdr"> 3</td>
    <td class="tdr"> 3</td>
    <td class="tdr"> 3</td>
    <td class="tdr"> 72</td>
  </tr>
  <tr>
    <td colspan="0">Alcedinidae</td>
  </tr>
  <tr>
    <td class="tdp">Belted kingfisher <em>(Megaceryle alcyon)</em></td>
    <td class="tdr"> 1</td>
    <td class="tdr"> 1</td>
    <td class="tdr"> 3</td>
    <td class="tdr"> 1</td>
    <td class="tdr"> 1</td>
    <td class="tdr"> 1</td>
    <td class="tdr"> 1</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 1</td>
    <td class="tdr"> 0</td>
    <td class="tdr"> 1</td>
    <td class="tdr"> 3</td>
    <td class="tdr"> 0</td>
    <td class="tdr"> 3</td>
    <td class="tdr"> 1</td>
    <td class="tdr"> 1</td>
    <td class="tdr"> 1</td>
    <td class="tdr"> 1</td>
    <td class="tdr"> 1</td>
    <td class="tdr"> 1</td>
    <td class="tdr"> 28</td>
  </tr>
  <tr>
    <td colspan="0">Corvidae</td>
  </tr>
  <tr>
    <td class="tdp">Common raven <em>(Corvus corax)</em></td>
    <td class="tdr"> 1</td>
    <td class="tdr"> 1</td>
    <td class="tdr"> 1</td>
    <td class="tdr"> 1</td>
    <td class="tdr"> 1</td>
    <td class="tdr"> 1</td>
    <td class="tdr"> 1</td>
    <td class="tdr"> 1</td>
    <td class="tdr"> 1</td>
    <td class="tdr"> 0</td>
    <td class="tdr"> 1</td>
    <td class="tdr"> 1</td>
    <td class="tdr"> 3</td>
    <td class="tdr"> 1</td>
    <td class="tdr"> 1</td>
    <td class="tdr"> 1</td>
    <td class="tdr"> 1</td>
    <td class="tdr"> 1</td>
    <td class="tdr"> 1</td>
    <td class="tdr"> 1</td>
    <td class="tdr"> 21</td>
  </tr>
  <tr>
    <td class="tdp">Northwestern crow <em>(C. caurinus)</em></td>
    <td class="tdr"> 3</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 3</td>
    <td class="tdr"> 3</td>
    <td class="tdr"> 1</td>
    <td class="tdr"> 1</td>
    <td class="tdr"> 1</td>
    <td class="tdr"> 1</td>
    <td class="tdr"> 1</td>
    <td class="tdr"> 0</td>
    <td class="tdr"> 1</td>
    <td class="tdr"> 3</td>
    <td class="tdr"> 1</td>
    <td class="tdr"> 1</td>
    <td class="tdr"> 1</td>
    <td class="tdr"> 1</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 5</td>
    <td class="tdr"> 47</td>
  </tr>
</tbody>
</table><span class="pagenum"><a name="Page_235" id="Page_235">[Pg 235]</a></span></div>

<div class="center">
<table border="0" cellpadding="4" cellspacing="0" summary="Table 2. Criteria and points used in calculating Oil Vulnerability Index.">
    <caption>Table 2. <em>Criteria and points used in calculating Oil Vulnerability Index.</em></caption>
  <tr>
    <th rowspan="2"></th>
    <th colspan="3">Point assignment</th>
  </tr>
  <tr>
    <th>1</th>
    <th>3</th>
    <th>5</th>
  </tr>
  <tr>
    <td colspan="0">Range</td>
  </tr>
  <tr>
    <td class="tdp">Breeding</td>
    <td>Large</td>
    <td>Medium</td>
    <td>Small</td>
  </tr>
  <tr>
    <td class="tdp">Migration</td>
    <td>Long</td>
    <td>Medium</td>
    <td>Short</td>
  </tr>
  <tr>
    <td class="tdp">Winter</td>
    <td>Large</td>
    <td>Medium</td>
    <td>Small</td>
  </tr>
  <tr>
    <td class="tdp">Marine orientation</td>
    <td>Coastal zone</td>
    <td>Intertidal</td>
    <td>Open water</td>
  </tr>
  <tr>
    <td colspan="0">Population</td>
  </tr>
  <tr>
    <td class="tdp">Size</td>
    <td>Large</td>
    <td>Medium</td>
    <td>Small</td>
  </tr>
  <tr>
    <td class="tdp">Productivity</td>
    <td>Large</td>
    <td>Medium</td>
    <td>Small</td>
  </tr>
  <tr>
    <td colspan="0">Habits</td>
  </tr>
  <tr>
    <td class="tdp">Roosting</td>
    <td>Shore</td>
    <td>Drift</td>
    <td>Water</td>
  </tr>
  <tr>
    <td class="tdp">Foraging</td>
    <td>Walking</td>
    <td>Flying</td>
    <td>Swimming</td>
  </tr>
  <tr>
    <td class="tdp">Escape</td>
    <td>Leave area</td>
    <td>Fly</td>
    <td>Dive</td>
  </tr>
  <tr>
    <td class="tdp">Flocking</td>
    <td>Small</td>
    <td>Medium</td>
    <td>Large</td>
  </tr>
  <tr>
    <td class="tdp">Nesting density</td>
    <td>Low</td>
    <td>Medium</td>
    <td>High</td>
  </tr>
  <tr>
    <td class="tdp">Specialization</td>
    <td>Low</td>
    <td>Medium</td>
    <td>High</td>
  </tr>
  <tr>
    <td colspan="0">Mortality</td>
  </tr>
  <tr>
    <td class="tdp">Hunted by man</td>
    <td>Low</td>
    <td>Medium</td>
    <td>High</td>
  </tr>
  <tr>
    <td class="tdp">Animal depredations</td>
    <td>Low</td>
    <td>Medium</td>
    <td>High</td>
  </tr>
  <tr>
    <td class="tdp">Non-oil pollution</td>
    <td>Low</td>
    <td>Medium</td>
    <td>High</td>
  </tr>
  <tr>
    <td class="tdp">History of oiling</td>
    <td>Low</td>
    <td>Medium</td>
    <td>High</td>
  </tr>
  <tr>
    <td colspan="0">Exposure</td>
  </tr>
  <tr>
    <td class="tdp">Spring</td>
    <td>Low</td>
    <td>Medium</td>
    <td>High</td>
  </tr>
  <tr>
    <td class="tdp">Summer</td>
    <td>Low</td>
    <td>Medium</td>
    <td>High</td>
  </tr>
  <tr>
    <td class="tdp">Fall</td>
    <td>Low</td>
    <td>Medium</td>
    <td>High</td>
  </tr>
  <tr>
    <td class="tdp">Winter</td>
    <td>Low</td>
    <td>Medium</td>
    <td>High</td>
  </tr>
</table></div>

<p>With these points in mind it is immediately
obvious that Southeast Alaska (Table 4),
which has only 9 high-score birds, offers far
less potential for bird problems than does the
Aleutian area (Table 5), which has 24 high-score
species. The planning agency could
make some immediate decisions on site priorities
and research funding based on such information.</p>


<h4>Discussion</h4>

<p>We are convinced that the OVI principle expressed
here will become a useful management
tool with all sorts of possible applications.
We recognize some difficulties with the
present version, but believe it is timely to present
the system so that a broader range of
thought, improvements, and application can
be applied to it.</p>

<p>Of prime importance is the system's simplicity.
The use of four levels of value for each
factor, instead of five or more, is an attempt
to simplify. Ian McHarg (1969) has shown
that extremely complex land-use values can
be graphically compared and displayed by
using three levels in a way that is useful to
decision makers. The difficulty of using more
levels of value was indicated by Sparrowe and
Wight (1975) who used up to 10 levels, enormously
complicating the problem of dealing
with low-quality information, which is often
all that is available. The use of scores of 0, 1,
3, 5 instead of 0, 1, 2, 3 for 20 factors enabled
us to use the convenient 100 points instead of
60 points as the maximum potential total
score for any species.</p>

<p>The 20 factors that were evaluated are admittedly
arbitrary; with refinement and more
detailed data they could be adjusted to show
better separation between affected species.
The decision to use 20 factors instead of more
or less again relates to simplicity. This appears
to be the minimum number that will assure
species separation and that can be neatly
displayed.</p>

<p><span class="pagenum"><a name="Page_236" id="Page_236">[Pg 236]</a></span></p>

<div class="center">
<table border="0" cellpadding="4" cellspacing="0" summary="Table 3.
Oil Vulnerability Index (OVI) for families of birds of the Northeast
Pacific marine habitats, excluding rare and endangered species in the
scoring.">
    <caption>Table 3. <em>Oil Vulnerability Index (OVI) for families of
    birds of the Northeast Pacific marine habitats, excluding rare
    and endangered species in the scoring.</em></caption>
  <tr>
    <th rowspan="2">Family</th>
    <th rowspan="2">Number of species</th>
    <th rowspan="2">Total OVI</th>
    <th colspan="0">OVI per species</th>
  </tr>
  <tr>
    <th>Average</th>
    <th>Range</th>
  </tr>
  <tr>
    <td>Loons&mdash;Gaviidae</td>
    <td class="tdn">4</td>
    <td class="tdn">219</td>
    <td class="tdn">55</td>
    <td class="tdc">47-65</td>
  </tr>
  <tr>
    <td>Grebes&mdash;Podicipedidae</td>
    <td class="tdn">3</td>
    <td class="tdn">148</td>
    <td class="tdn">49</td>
    <td class="tdc">44-56</td>
  </tr>
  <tr>
    <td>Albatrosses&mdash;Diomedeidae</td>
    <td class="tdn">2</td>
    <td class="tdn">102</td>
    <td class="tdn">51</td>
    <td class="tdc">50-52</td>
  </tr>
  <tr>
    <td>Shearwaters&mdash;Procellaridae</td>
    <td class="tdn">4</td>
    <td class="tdn">208</td>
    <td class="tdn">52</td>
    <td class="tdc">47-57</td>
  </tr>
  <tr>
    <td>Storm-petrels&mdash;Hydrobatidae</td>
    <td class="tdn">2</td>
    <td class="tdn">130</td>
    <td class="tdn">65</td>
    <td class="tdc">63-67</td>
  </tr>
  <tr>
    <td>Cormorants&mdash;Phalacrocoracid</td>
    <td class="tdn">4</td>
    <td class="tdn">235</td>
    <td class="tdn">59</td>
    <td class="tdc">52-63</td>
  </tr>
  <tr>
    <td>Herons&mdash;Ardeidae</td>
    <td class="tdn">1</td>
    <td class="tdn">29</td>
    <td class="tdn">29</td>
    <td class="tdc">29</td>
  </tr>
  <tr>
    <td>Waterfowl&mdash;Anatidae</td>
    <td class="tdn">33</td>
    <td class="tdn">1,765</td>
    <td class="tdn">53</td>
    <td class="tdc">32-78</td>
  </tr>
  <tr>
    <td>Eagles and Hawks&mdash;Accipitridae</td>
    <td class="tdn">2</td>
    <td class="tdn">77</td>
    <td class="tdn">39</td>
    <td class="tdc">19-58</td>
  </tr>
  <tr>
    <td>Ospreys&mdash;Pandionidae</td>
    <td class="tdn">1</td>
    <td class="tdn">37</td>
    <td class="tdn">37</td>
    <td class="tdc">37</td>
  </tr>
  <tr>
    <td>Falcons&mdash;Falconidae</td>
    <td class="tdn">1</td>
    <td class="tdn">41</td>
    <td class="tdn">41</td>
    <td class="tdc">41</td>
  </tr>
  <tr>
    <td>Cranes&mdash;Gruidae</td>
    <td class="tdn">1</td>
    <td class="tdn">24</td>
    <td class="tdn">24</td>
    <td class="tdc">24</td>
  </tr>
  <tr>
    <td>Rails and Coots&mdash;Rallidae</td>
    <td class="tdn">1</td>
    <td class="tdn">33</td>
    <td class="tdn">33</td>
    <td class="tdc">33</td>
  </tr>
  <tr>
    <td>Oystercatchers&mdash;Haematopodidae</td>
    <td class="tdn">1</td>
    <td class="tdn">65</td>
    <td class="tdn">65</td>
    <td class="tdc">65</td>
  </tr>
  <tr>
    <td>Plovers&mdash;Charadriidae</td>
    <td class="tdn">7</td>
    <td class="tdn">287</td>
    <td class="tdn">41</td>
    <td class="tdc">26-57</td>
  </tr>
  <tr>
    <td>Sandpipers&mdash;Scolopacidae</td>
    <td class="tdn">22</td>
    <td class="tdn">857</td>
    <td class="tdn">39</td>
    <td class="tdc">24-59</td>
  </tr>
  <tr>
    <td>Phalaropes&mdash;Phalaropodidae</td>
    <td class="tdn">2</td>
    <td class="tdn">120</td>
    <td class="tdn">60</td>
    <td class="tdc">58-62</td>
  </tr>
  <tr>
    <td>Jaegers and Skuas&mdash;Stercorariidae</td>
    <td class="tdn">3</td>
    <td class="tdn">123</td>
    <td class="tdn">41</td>
    <td class="tdc">39-43</td>
  </tr>
  <tr>
    <td>Gulls and Terns&mdash;Laridae</td>
    <td class="tdn">16</td>
    <td class="tdn">730</td>
    <td class="tdn">46</td>
    <td class="tdc">32-66</td>
  </tr>
  <tr>
    <td>Auks&mdash;Alcidae</td>
    <td class="tdn">15</td>
    <td class="tdn">1,164</td>
    <td class="tdn">78</td>
    <td class="tdc">70-88</td>
  </tr>
  <tr>
    <td>Kingfishers&mdash;Alcedinidae</td>
    <td class="tdn">1</td>
    <td class="tdn">28</td>
    <td class="tdn">28</td>
    <td class="tdc">28</td>
  </tr>
  <tr>
    <td>Crows&mdash;Corvidae</td>
    <td class="tdn">2</td>
    <td class="tdn">68</td>
    <td class="tdn">34</td>
    <td class="tdc">21-47</td>
  </tr>
  <tr>
    <td class="tdp">Total and Mean</td>
    <td class="tdn">128</td>
    <td class="tdn">6,490</td>
    <td class="tdn">51</td>
    <td class="tdc">19-88</td>
  </tr>
</table></div>

<p>The system will be much more useful when
it is expanded to the subspecific level. Many
Holarctic species are represented in the
Northeast Pacific by a single race that would
have a much higher OVI than the species as a
whole. For example, the OVI for the Peale's
peregrine falcon <em>(Falco peregrinus pealei)</em> confined
to marine habitats within the Pacific region
would be high; and the endangered Aleutian
Canada goose <em>(Branta canadensis leucopareia)</em>
would score 100 points instead of the
34 we show for Canada geese <em>(B. c.)</em>. If Tables
4 and 5 showed subspecies, the differences in
value would be more marked.</p>

<p>Tables 4 and 5 are for broad geographical
areas. A comparison between smaller areas
would probably show more dramatic differences.</p>

<p>Because the dearth of easily available, applicable
information poses a problem in
evaluating the various factors, our scoring
was conservative. Experts on the various
avian families can doubtless refine the scoring.
If this system proves useful, investigators
will begin to acquire the information
needed for more precise evaluations. Ultimate
perfection may never be achieved; however, as
with the field guides, the fact of minor professional
disagreement should not destroy the
system's utility.</p>

<p>We believe re-scoring of all birds on the basis
of various projects should be avoided because
a standard against which individual projects
can be measured is needed. If everyone did
their own scoring, there would be no standard,
and projects evaluated by different investigators
would not be comparable. If a species list
for the project area and standard point scores
are used, the level of involvement for many
species and perhaps for most species will be
properly identified. As with any system, there
will be exceptions and the assessor will need
to deal with these as appropriate. The result
will still be to focus attention on those species
and impacting factors where it is most
needed.</p>

<p><span class="pagenum"><a name="Page_237" id="Page_237">[Pg 237]</a></span></p>

<div class="center">
<table border="0" class="tdt" cellpadding="4" cellspacing="0" summary="Table 4. Oil
Vulnerability Index for 109 species of birds of Southeast Alaska">
    <caption>Table 4. <em>Oil Vulnerability Index for
    109 species of birds of Southeast Alaska (Total
    Points&mdash;2,678).</em></caption>
<col width="2em"></col>
<col width="1em"></col>
<col width="2em"></col>
<col width="1em"></col>
<col width="2em"></col>
<col width="1em"></col>
<col width="2em"></col>
<col width="1em"></col>
<col width="2em"></col>
<col width="1em"></col>
  <tr>
    <th colspan="2" class="br">OVI 1-20</th>
    <th colspan="2" class="br">OVI 21-40</th>
    <th colspan="2" class="br">OVI 41-60</th>
    <th colspan="2" class="br">OVI 61-80</th>
    <th colspan="2">OVI 81-100</th>
  </tr>
  <tr>
    <td>Marsh hawk</td>
    <td class="tdn">19</td>
    <td>Great blue heron</td>
    <td class="tdn">29</td>
    <td>Common loon</td>
    <td class="tdn">47</td>
    <td>Pelagic cormorant</td>
    <td class="tdn">63</td>
    <td>Pigeon guillemot </td>
    <td class="tdn">82</td>
  </tr>
  <tr>
    <td rowspan="25" class="tdt">52 species, rare or occasional (one point each) </td>
    <td rowspan="25" class="tdn tdt">52</td>
    <td>Canada goose</td>
    <td class="tdn">34</td>
    <td>Arctic loon</td>
    <td class="tdn">58</td>
    <td>Oldsquaw</td>
    <td class="tdn">66</td>
    <td rowspan="0" class="tdt">Marbled murrelet </td>
    <td rowspan="25" class="tdn tdt">84</td>
  </tr>
  <tr>
    <td>White-fronted goose </td>
    <td class="tdn">36</td>
    <td>Red-throated loon</td>
    <td class="tdn">49</td>
    <td>White-winged scoter </td>
    <td class="tdn">72</td>
  </tr>
  <tr>
    <td>Snow goose</td>
    <td class="tdn">32</td>
    <td>Red-necked grebe</td>
    <td class="tdn">44</td>
    <td>Surf scoter</td>
    <td class="tdn">72</td>
  </tr>
  <tr>
    <td>Mallard</td>
    <td class="tdn">36</td>
    <td>Horned grebe</td>
    <td class="tdn">48</td>
    <td>Black oystercatcher </td>
    <td class="tdn">65</td>
  </tr>
  <tr>
    <td>Pintail</td>
    <td class="tdn">36</td>
    <td>Whistling swan</td>
    <td class="tdn">50</td>
    <td>Northern phalarope</td>
    <td class="tdn">62</td>
  </tr>
  <tr>
    <td>Green-winged teal</td>
    <td class="tdn">34</td>
    <td>Trumpeter swan</td>
    <td class="tdn">63</td>
    <td rowspan="0" class="tdt">Common murre</td>
    <td rowspan="20" class="tdn tdt">70</td>
  </tr>
  <tr>
    <td>American wigeon</td>
    <td class="tdn">36</td>
    <td>Greater scaup</td>
    <td class="tdn">52</td>
  </tr>
  <tr>
    <td>Semipalmated plover </td>
    <td class="tdn">28</td>
    <td>Lesser scaup</td>
    <td class="tdn">52</td>
  </tr>
  <tr>
    <td>Killdeer</td>
    <td class="tdn">26</td>
    <td>Common goldeneye</td>
    <td class="tdn">48</td>
  </tr>
  <tr>
    <td>Common snipe</td>
    <td class="tdn">29</td>
    <td>Barrow's goldeneye</td>
    <td class="tdn">56</td>
  </tr>
  <tr>
    <td>Spotted sandpiper</td>
    <td class="tdn">24</td>
    <td>Bufflehead</td>
    <td class="tdn">52</td>
  </tr>
  <tr>
    <td>Greater yellowlegs</td>
    <td class="tdn">30</td>
    <td>Harlequin duck</td>
    <td class="tdn">60</td>
  </tr>
  <tr>
    <td>Lesser yellowlegs</td>
    <td class="tdn">30</td>
    <td>Common merganser</td>
    <td class="tdn">56</td>
  </tr>
  <tr>
    <td>Pectoral sandpiper</td>
    <td class="tdn">32</td>
    <td>Red-breasted merganser </td>
    <td class="tdn">56</td>
  </tr>
  <tr>
    <td>Least sandpiper</td>
    <td class="tdn">34</td>
    <td>Bald eagle</td>
    <td class="tdn">58</td>
  </tr>
  <tr>
    <td>Herring gull</td>
    <td class="tdn">38</td>
    <td>Peregrine falcon</td>
    <td class="tdn">41</td>
  </tr>
  <tr>
    <td>Bonaparte's gull</td>
    <td class="tdn">40</td>
    <td>Black turnstone</td>
    <td class="tdn">57</td>
  </tr>
  <tr>
    <td>Arctic tern</td>
    <td class="tdn">32</td>
    <td>Rock sandpiper</td>
    <td class="tdn">59</td>
  </tr>
  <tr>
    <td>Belted kingfisher</td>
    <td class="tdn">28</td>
    <td>Dunlin</td>
    <td class="tdn">41</td>
  </tr>
  <tr>
    <td rowspan="0" class="tdt">Common raven</td>
    <td rowspan="6" class="tdn tdt">21</td>
    <td>Short-billed dowitcher </td>
    <td class="tdn">41</td>
  </tr>
  <tr>
    <td>Western sandpiper</td>
    <td class="tdn">47</td>
  </tr>
  <tr>
    <td>Glaucous-winged gull</td>
    <td class="tdn">56</td>
  </tr>
  <tr>
    <td>Thayer's gull</td>
    <td class="tdn">42</td>
  </tr>
  <tr>
    <td>Mew gull</td>
    <td class="tdn">44</td>
  </tr>
  <tr>
    <td rowspan="0" class="tdt">Northwestern crow</td>
    <td class="tdn">47</td>
  </tr>
  <tr>
    <td>Totals</td>
    <td class="tdn">71</td>
    <td class="tdn">665</td>
    <td class="tdn">1,324</td>
    <td class="tdn">470</td>
    <td class="tdn">166</td>
  </tr>
</table><span class="pagenum"><a name="Page_238" id="Page_238">[Pg 238]</a></span></div>

<div class="center">
<table border="0" cellpadding="4" cellspacing="0" summary="Table 5. Oil
Vulnerability Index for 123 species of birds of the Aleutian Islands">
    <caption>Table 5. <em>Oil Vulnerability Index for 123 species of birds
    of the Aleutian Islands</em> (Total Points&mdash;2,689).</caption>
<col width="2em"></col>
<col width="1em"></col>
<col width="2em"></col>
<col width="1em"></col>
<col width="2em"></col>
<col width="1em"></col>
<col width="2em"></col>
<col width="1em"></col>
<col width="2em"></col>
<col width="1em"></col>
  <tr>
    <th colspan="2" class="br">OVI 1-20</th>
    <th colspan="2" class="br">OVI 21-40</th>
    <th colspan="2" class="br">OVI 41-60</th>
    <th colspan="2" class="br">OVI 61-80</th>
    <th colspan="2">OVI 81-100</th>
  </tr>
  <tr>
    <td rowspan="22" class="tdt">80 species, rare or occasional (one point each) </td>
    <td rowspan="22" class="tdt tdn">80</td>
    <td>Canada goose</td>
    <td class="tdn">34</td>
    <td>Fulmar</td>
    <td class="tdn">57</td>
    <td>Fork-tailed storm-petrel </td>
    <td class="tdn">67</td>
    <td>Pigeon guillemot</td>
    <td class="tdn">82</td>
  </tr>
  <tr>
    <td>Least sandpiper </td>
    <td class="tdn">34</td>
    <td>Slender-billed shearwater </td>
    <td class="tdn">53</td>
    <td>Leach's storm-petrel</td>
    <td class="tdn">63</td>
    <td rowspan="0" class="tdt">Whiskered auklet</td>
    <td rowspan="21" class="tdn tdt">88</td>
  </tr>
  <tr>
    <td>Arctic tern</td>
    <td class="tdn">32</td>
    <td>Greater scaup</td>
    <td class="tdn">52</td>
    <td>Pelagic cormorant</td>
    <td class="tdn">63</td>
  </tr>
  <tr>
    <td rowspan="0" class="tdt">Common raven</td>
    <td rowspan="19" class="tdt tdn">21</td>
    <td>Common goldeneye</td>
    <td class="tdn">48</td>
    <td>Red-faced cormorant</td>
    <td class="tdn">63</td>
  </tr>
  <tr>
    <td>Bufflehead</td>
    <td class="tdn">52</td>
    <td>Black Brant</td>
    <td class="tdn">70</td>
  </tr>
  <tr>
    <td>Harlequin duck</td>
    <td class="tdn">60</td>
    <td>Emperor goose</td>
    <td class="tdn">70</td>
  </tr>
  <tr>
    <td>Bald eagle</td>
    <td class="tdn">58</td>
    <td>Oldsquaw</td>
    <td class="tdn">66</td>
  </tr>
  <tr>
    <td>Peregrine falcon</td>
    <td class="tdn">41</td>
    <td>Steller's eider</td>
    <td class="tdn">72</td>
  </tr>
  <tr>
    <td>Ruddy turnstone</td>
    <td class="tdn">44</td>
    <td>Common eider</td>
    <td class="tdn">68</td>
  </tr>
  <tr>
    <td>Rock sandpiper</td>
    <td class="tdn">59</td>
    <td>King eider</td>
    <td class="tdn">70</td>
  </tr>
  <tr>
    <td>Western sandpiper</td>
    <td class="tdn">47</td>
    <td>White-winged scoter</td>
    <td class="tdn">72</td>
  </tr>
  <tr>
    <td>Red phalarope</td>
    <td class="tdn">58</td>
    <td>Common scoter</td>
    <td class="tdn">72</td>
  </tr>
  <tr>
    <td>Parasitic jaeger</td>
    <td class="tdn">43</td>
    <td>Black oystercatcher</td>
    <td class="tdn">65</td>
  </tr>
  <tr>
    <td>Glaucous-winged gull</td>
    <td class="tdn">56</td>
    <td>Red-legged kittiwake</td>
    <td class="tdn">66</td>
  </tr>
  <tr>
    <td>Black-legged kittiwake</td>
    <td class="tdn">49</td>
    <td rowspan="0" class="tdt">Common murre</td>
    <td rowspan="8" class="tdn tdt">70</td>
  </tr>
  <tr>
    <td>Thick-billed murre</td>
    <td class="tdn">70</td>
  </tr>
  <tr>
    <td>Ancient murrelet</td>
    <td class="tdn">74</td>
  </tr>
  <tr>
    <td>Parakeet auklet</td>
    <td class="tdn">80</td>
  </tr>
  <tr>
    <td>Crested auklet</td>
    <td class="tdn">76</td>
  </tr>
  <tr>
    <td>Least auklet</td>
    <td class="tdn">80</td>
  </tr>
  <tr>
    <td>Horned puffin</td>
    <td class="tdn">72</td>
  </tr>
  <tr>
    <td rowspan="0" class="tdt">Tufted puffin</td>
    <td class="tdn">72</td>
  </tr>
  <tr>
    <td>Totals</td>
    <td class="tdn">80</td>
    <td class="tdn">121</td>
    <td class="tdn">777</td>
    <td class="tdn">1,541</td>
    <td class="tdn">170</td>
  </tr>
</table><span class="pagenum"><a name="Page_239" id="Page_239">[Pg 239]</a></span></div>

<p>We have used our OVI system to show the
vulnerability of birds to oil, but it seems likely
that the vulnerability index could be applied
on a much broader scale to help make decisions
in other areas of human activity and resource
development. The vulnerability index
system could be applied to terrestrial as well
as aquatic species by adding or subtracting
impacting factors, as appropriate. Indexes relating
the impact of man upon each North
American species could have broad uses in the
field of conservation. Population explosions,
as well as declines, might be predictable.
Human activity could be better adjusted to
favor or depress wildlife populations, as
appropriate.</p>

<p>We believe that this vulnerability index system
has promise for aiding in the decision-making
processes upon which future bird conservation
will depend.</p>


<h4>References</h4>

<div class="hangindent">

<p>American Ornithologists' Union. 1957. Check-list of
North American birds. 5th ed. Lord Baltimore
Press, Baltimore, Maryland.</p>

<p>Fay, F. H., and T. J. Cade. 1959. An ecological
analysis of the avifauna of St. Lawrence Island,
Alaska. Univ. Calif. Publ. Zool. 63(2):73-150.</p>

<p>Gabrielson, J. N., and F. C. Lincoln. 1959. The birds
of Alaska. The Stackpole Company, Harrisburg,
Pa., and Wildlife Management Institute, Washington,
D.C. 922 pp.</p>

<p>Gibson, D. D. 1970. Check-list of the birds of
Alaska. Univ. of Alaska, Fairbanks. 2 pp.</p>

<p>Isleib, M. E., and B. Kessel. 1973. Birds of the
North Gulf Coast-Prince William Sound region,
Alaska. Univ. of Alaska Biol. Pap. 14. 149 pp.</p>

<p>Kortright, F. H. 1942. The ducks, geese and swans
of North America. Wildlife Management Institute,
Washington, D.C.</p>

<p>McHarg, I. L. 1969. Design with nature. Natural
History Press, Garden City, N. Y. 197 pp.</p>

<p>Murie, O. J. 1959. Fauna of the Aleutian Islands
and Alaska Peninsula. U.S. Fish Wildl. Serv., N.
Am. Fauna 61:1-364.</p>

<p>Palmer, R. S. 1962. Handbook of North American
birds. Vol. 1. Yale University Press, New Haven,
Conn. 567 pp.</p>

<p>Peterson, R. T., G. Montfort, and P. H. D. Hallom.
1967. A field guide to the birds of Britain and
Europe. Houghton Mifflin Co., New York.</p>

<p>Robbins, D. S., B. Bruun, and H. S. Zimm. 1966.
Birds of North America, a guide to field identification.
Golden Press, New York.</p>

<p>Sanger, G. A. 1972. Preliminary standing stock and
biomass estimates of seabirds in the subarctic
Pacific region. Pages 589-611 <em>in</em> A. Y. Takenouti
et al., eds. Biological oceanography of the North
Pacific Ocean. Idemitsy Shoten, Tokyo.</p>

<p>Sparrowe, R. D., and H. M. Wight. 1975. Setting
priorities for the endangered species program.
Proc. N. Am. Wildl. Nat. Resourc. Conf. 40:142-156.</p>

<p>Stout, G. D., P. Matthiessen, V. R. Clem, and R. S.
Palmer. 1967. The shorebirds of North America.
Viking Press, New York. 270 pp.</p>

<p>U.S. Fish and Wildlife Service. 1974. Birds of the
Aleutian Islands National Wildlife Refuge. Washington,
D.C.</p>

<p>U.S. Forest Service and Alaska Department of
Fish and Game. 1970. Birds of Southeast Alaska,
a check-list. The agencies, Juneau, Alaska. 12 pp.</p></div>

<p><span class="pagenum"><a name="Page_240" id="Page_240">[Pg 240]</a><br /><a name="Page_241" id="Page_241">[Pg 241]</a></span></p>



<p><span class="pagenum"><a name="Page_242" id="Page_242">[Pg 242]</a><br /><a name="Page_243" id="Page_243">[Pg 243]</a></span></p>
<div class="chapter"></div>
<hr class="chap" />




<h2><a name="PROGRAMS_AND_AUTHORITIES_RELATED_TO" id="PROGRAMS_AND_AUTHORITIES_RELATED_TO">PROGRAMS AND AUTHORITIES RELATED TO
MARINE BIRD CONSERVATION</a></h2>
<div class="chapter"></div>
<hr class="chap" />




<h3><a name="Programs_and_Authorities_Related_to_Marine_Bird" id="Programs_and_Authorities_Related_to_Marine_Bird">Programs and Authorities Related to Marine Bird
Conservation in Washington State</a></h3>


<p class="center">by</p>

<p class="center">Ralph W. Larson</p>

<p class="center">Washington Department of Game<br />
600 North Capitol Way<br />
Olympia, Washington 98504
</p>


<h4>Abstract</h4>

<div class="blockquot">

<p>Seabirds are one of the most visible biological components of ecosystems, and
yet little is known about them. They could readily be used as an index of marine
environmental quality if adequate studies were conducted to determine populations,
habitat needs, and causes of fluctuations in abundance. The lack of adequate
funding at the State level has precluded necessary studies to make these
determinations and to provide habitat protection and preservation.</p>

<p>The State of Washington has developed a funding source for protection,
preservation, and enhancement of nongame wildlife, which includes seabirds.
The sale of personalized license plates for vehicles is now providing some funds
for nongame wildlife management&mdash;funds which should increase as the popularity
of the licensing program increases. Outdoor Recreation Bonds are providing
funding for habitat preservation. Authorities provided the Washington
Game Department are adequate to manage and protect seabird species. Other
State laws offer additional protection to their habitat&mdash;specifically the Shoreline
Management Act of 1971 and the State Environmental Act.</p></div>

<p>It has been often stated that seabirds are
one of the most visible biological components
of ecosystems, and yet little is known about
them. Most studies to date have been on fish,
and because of their recreational and commercial
value, the concern for maintaining the
marine environment has been primarily a result
of the concern for maintaining the fishery
resource. The visible knowledge of the fishery
resource, however, becomes an "after-the-fact"
knowledge since the status of the stocks
relates to the value and amount of the
fishery&mdash;a fishery resulting from survival
under the surface in the marine environment
that can be very secretive about its quality
until it is too late to do something about it.
Seabirds, however, are visible above the surface,
in numbers that can reflect changes in
the marine environment that occur below the
surface, since many depend on the subsurface
quality that reflects populations of fish.</p>

<p>Studies in Oregon have indicated that consumption
of pelagic fish by murres (<em>Uria</em> spp.),
cormorants (<em>Phalacrocorax</em> spp.), storm-petrels
(<em>Oceanodroma</em> spp.), and shearwaters
(<em>Puffinus</em> spp.) account for about 22% of the
annual production of various species of these
fish. A decline in this food source will reflect a
decline in the seabird population. Why then
should it be necessary to use only fish populations
as an index of marine environmental
quality, when seabirds can more readily be observed
and can reflect the same changes that
occur?</p>

<p>As a public wildlife agency, the Washington
Department of Game is often attempting to
justify the value of seabirds, and sometimes
that is not easy. When fishermen complain
that the seabirds are eating all of the food of
our mighty salmon, and hunters indicate little
compassion because the birds have no value
to sport hunting, one has to think a little to<span class="pagenum"><a name="Page_244" id="Page_244">[Pg 244]</a></span>
explain their value. However, rhinoceros auklets
<em>(Cerorhinca monocerata)</em> do drive herring
into ball-shaped schools, which attracts salmon
in search of food&mdash;which in turn provides
a signal to fishermen that salmon may soon be
in the area. Explaining value to the hunter is a
bit more difficult, but anyone who has taken
the time to go out on our marine waters and
observe the many species of seabirds and
watch them flying and feeding cannot help
but be fascinated by them. The flight of thousands
of murres skimming over the water surface
and somehow managing not to dash
headlong into a wave is a fascinating sight.</p>

<p>We who are in fish and wildlife work have
had to readjust our thinking and values during
recent years. Our primary programs and
concerns for many years were with the fish,
birds, and animals that were of value to fishermen
and hunters. Species of wildlife that we
now classify as nongame received incidental
benefit from programs related to game fish,
game birds, and game animals, but we did not
do badly in maintaining and enhancing these
incidental wildlife species, mostly by indirection.
However, in the last few years our Department,
at least, has taken on a new responsibility
and a new look as related to nongame
wildlife.</p>

<p>Our first positive step in this direction was
to develop a funding source for nongame wildlife
programs. Our funding attempt charted
its way through stormy waters, but finally
ended up being voted on by the citizens of the
State. Our citizens passed Referendum 33,
which provided funds to the Department for
nongame wildlife programs from the sale of
personalized license plates. Although the
funds have not been adequate, they are a step
in the right direction and have permitted the
Department to engage in a modest program of
research and management. We have placed
one person in charge of our program to do the
planning and programming so necessary for
developing an effective, growing program.
During the 1st year of operation, we contracted
studies on the rhinoceros auklet, the
tufted puffin <em>(Lunda cirrhata)</em>, and the black
oystercatcher <em>(Haematopus bachmani)</em>. These
studies have provided a basic knowledge of
some of the problems facing these seabird
species. As funds increase, additional studies
will be made to provide more information on
these birds and others.</p>

<p>During the 1975 legislative session we were
successful in amending the personalized
license program to include automobiles other
than passenger cars&mdash;a step which should
further enhance our funding. We anticipate
that funding will increase from the sale of
these license plates each year. They serve as
their own advertisement, and as more plates
are sold, the exposure to the public increases.
We anticipate that within the next few years
the funding should reach $150,000 per year&mdash;a
modest sum to be sure, but nevertheless adequate
to establish a viable program.</p>

<p>We have been involved in studies funded
through other agencies that involve seabirds.
The principal reasons for the studies are not
seabirds, but they become an integral part of
any analysis that must be made of our saltwater
environs. One such study involves a
comprehensive status survey of the marine
shoreline fauna of Washington. The Department
of Ecology has provided the funding as
a part of their analysis of resources that may
be adversely affected by oil spills and economic
development of our shorelines. This
study will be the first one designed to comprehensively
identify wildlife species associated
with our shorelines and will determine the
species, their status, location, and habitat.
This study will provide a basis for readily
identifying visually the results of oil spills and
of the economic development of critical habitat
areas, and provide sound basic data for
use in combating destructive projects in the
marine environment.</p>

<p>We are finding that you cannot separate
functions of other governmental agencies that
deal with marine waters from seabird analysis.
Pollution responsibilities, shoreline management,
coastal zone management, clam
dredging, channel dredging, erosion control,
housing development, industrial expansion,
shipping port development&mdash;to name a few&mdash;all
must have some effect on our seabird
species. Therefore, we must concentrate on
obtaining an adequate data base to insure the
perpetuation of these valuable marine species.</p>

<p>As I indicated earlier, the Department of
Game has not had a special program to manage
seabirds in the past, but this should not
indicate that we have not assisted in maintaining
the seabird resource. Our basic land<span class="pagenum"><a name="Page_245" id="Page_245">[Pg 245]</a></span>
acquisition program designed for waterfowl
enhancement has benefited seabirds. We now
own some 15,500 acres of lands, tideland, and
marshes bordering the marine waters (including
our Skagit and Nisqually holdings) which
provide habitat and protection for many seabirds.
We also recently acquired 48 acres on
Protection Island, designed to protect the
nesting area of the rhinoceros auklet. This
purchase was an excellent example of how
combined efforts of several groups accomplished
a nearly impossible goal.</p>

<p>Protection Island had been subdivided for
summer home development and many lots
had been sold. The developer, however, got
caught in the requirements of our Shoreline
Management Act with his last subdivision.
The uproar caused due to the use of this subdivision
by auklets created an atmosphere
that made subdivision a real conservation
issue. The outspoken critics of the project
from the Audubon Society, Fish and Wildlife
Service, independent conservationists, and
our Department enlisted the aid of Nature
Conservancy to negotiate for purchase of this
subdivision, and after lengthy negotiations
the option was obtained, and the Department
purchased the land from the Nature Conservancy
with funds provided by the Interagency
Committee for Outdoor Recreation.
The area now is destined to be a seabird sanctuary,
with limited public viewing and incidental
recreation use. This project is an excellent
example of the power of cooperative efforts
by conservationists to protect a
resource.</p>

<p>The State of Washington now has a reasonably
good legislative base to insure constructive
programs for management of our seabird
resource. Our legislative authority lies in
State statutes under Title 77. These authorities
first provide that the wild birds, wild animals,
and game fish of the State are the property
of the State and that they shall be preserved,
protected, and perpetuated. Any regulations
for taking shall be designed so as to
not "impair the supply thereof."</p>

<p>The commission also has the authority to
classify wild birds. Seabirds, other than
hunted species, fall into the category of nongame
birds. We also have the authority to
regulate the propagation and protection of
wild birds, develop rules and regulations for
taking them (or to prohibit taking them), and
to create game reserves and closed areas
where necessary to protect various species.
Our authorities also include the obligation to
enforce the laws, rules, and regulations pertaining
to the protection of all wild birds.</p>

<p>The Department may also acquire land for
habitat and for sanctuaries for nongame birds
and may exchange lands for these purposes.
We may also enter into agreements with the
Federal Government, persons, and municipal
subdivisions of the State for all matters relating
to propagation, protection, and conservation
of all wild birds, and may lease State
lands for this purpose.</p>

<p>We believe our authorities are now totally
adequate to satisfactorily manage the State's
marine bird resources.</p>

<p>In addition to our personalized license plate
legislation, which earmarks funds for nongame
wildlife, other State laws and programs
assist in protection of this resource. One program
that has assisted materially in providing
funds for habitat acquisition is a bond
issue passed by citizens of the State designed
to acquire and develop recreational land in the
State for public use. Our Interagency Committee
for Outdoor Recreation provides the
necessary mechanism for funding of projects,
using these bond monies to match Federal
funds. Although recreation is a key factor in
obtaining funding, it is still possible to acquire
key habitat for wildlife and develop a
people-use program around the primary purpose
for acquisition.</p>

<p>The purchase of a portion of Protection Island
was accomplished by use of these funds,
as I indicated earlier, and we are now working
again with Nature Conservancy to acquire
key bald eagle habitat on the Skagit River in
northwestern Washington. The bond issues
total some $50 million, of which this Department
receives about 15%. The State now is in
its third bond issue, and we hope the citizens
will continue to support this program.</p>

<p>One of the newer laws is the Shoreline Management
Act of 1971. This act provides for development
of comprehensive shoreline management
programs designed to control the development
of these areas to insure protection
of the public interest, while still recognizing
and protecting private property rights consistent
with this public interest. These plans<span class="pagenum"><a name="Page_246" id="Page_246">[Pg 246]</a></span>
must be developed with citizen involvement.
Shoreline classification generally falls into
four categories&mdash;natural, conservancy, rural,
and urban. The natural classification can accomplish
the most substantial benefit for
marine birds. Provisions are also made for
protection of "shorelines of statewide significance."
Plans for these areas must give preference
to uses favoring the public and long-range
goals. These shorelines cover the areas
between low and high tide levels on inland
waters and high water and the western boundary
of the State on our Pacific Ocean coast.</p>

<p>Our State Environmental Policy Act, which
requires that environmental impact statements
be prepared for various programs and
developments, gives our Department an opportunity
to insure that our valuable wildlife
resources are given consideration during the
planning phase of the proposed project.</p>

<p>The Department feels that our authorities
at this time are adequate to protect marine
bird populations and their habitat. The one
lacking factor, as usual, is the funding for
both adequate management programs and
habitat protection. Our marine habitat is
rapidly being developed for recreational
homesites and public use which can eliminate
key habitat use. A greater public awareness of
the needs of marine birds can be a help in preventing
destruction of their habitat; however,
money talks the loudest. The acquisition of
these key habitats is the most positive means
of insuring their retention. We have no solution
at this time to the funding problem and
only hope that someone smarter than we are
can provide an acceptable solution before all
of our efforts become too little and too late.</p>

<p><span class="pagenum"><a name="Page_247" id="Page_247">[Pg 247]</a></span></p>
<div class="chapter"></div>
<hr class="chap" />




<h3><a name="Programs_and_Authorities_of_the_Province_of" id="Programs_and_Authorities_of_the_Province_of">Programs and Authorities of the Province of
British Columbia Related to Marine Bird Conservation</a></h3>


<p class="center">by</p>

<p class="center">W. T. Munro</p>

<p class="center">British Columbia Fish and Wildlife Branch<br />
300-1019 Wharf Street<br />
Victoria, British Columbia, Canada V8W 2Z1</p>

<p class="center">and</p>

<p class="center">R. Wayne Campbell</p>

<p class="center">British Columbia Provincial Museum<br />
Victoria, British Columbia, Canada
</p>


<h4>Abstract</h4>

<div class="blockquot">

<p>British Columbia Provincial agencies are given authority for protecting marine
birds and their habitats by the Provincial Wildlife Act, the Parks Act, and the
Ecological Reserves Act. The Provincial Museum Act accommodates research on
marine birds. The Fish and Wildlife branch has protected over 30,000 ha of intertidal
estuarine habitat in the form of reserves and has conducted limited inventories
of birds on the Queen Charlotte Islands and northern mainland coast. The
Provincial Museum has conducted inventories and life-history studies of marine
birds and maintains a repository for information on seabirds, including a catalog
of colonies. Pollution from oil and chemicals, improper logging practices, and disturbance
by boating recreationists are the most apparent threats to the well-being
of birds. Additional inventories and the determination of seasonal distribution
are among the information needed to better protect the marine birds of
British Columbia.</p></div>

<p>Most marine-associated birds in Canada are
covered by the Migratory Birds Convention
Act and are therefore federally protected. In
British Columbia additional protection is provided
by the Provincial Wildlife Act. Several
other provincial acts provide authorities related
to seabirds. The Provincial Museum Act
permits research related to natural history;
the Parks Act and Ecological Reserves Act
provide for the protection of habitat and prohibit
harassment of wildlife within park and
reserve boundaries; and the Firearms Act permits
the closure of areas frequented by selected
wildlife to the discharge of firearms.
The fact that several authorities for the protection
and conservation of marine birds are
available does not mean that they have been
used to full advantage.</p>

<p>British Columbia's irregular shores provide
thousands of kilometers of coastline, much of
which is used by marine birds for nesting and
wintering as well as during migration.
Through legislation of different types, some of
the more ecologically important and unique
sites have been protected. Twelve "ecological
reserves," which are basically inviolate preserve
areas, provide habitat for and protection
to a number of major breeding colonies.
Over 30,000 ha of intertidal estuarine habitat
has been protected by the provincial Fish and
Wildlife Branch in the form of reserves. Less
than half of the total area is in Order-in-Council
reserves (passed by the Provincial Cabinet),
which afford strong protection; the rest
is in departmental map reserves, which
merely means other agencies must inform the<span class="pagenum"><a name="Page_248" id="Page_248">[Pg 248]</a></span>
branch before they disturb them; they are
hardly secure. Provincial Parks Branch protects
other areas used by marine birds by incorporating
them within parks.</p>

<p>Research and conservation of seabirds in
British Columbia have not been a high
priority in the Fish and Wildlife Branch, basically
because seabirds are not consumed by
people. Our primary interest in seabirds has
been in their role as a life support system for
the peregrine falcon <em>(Falco peregrinus)</em>. Most
Fish and Wildlife Branch reserves have been
established to protect estuarine habitat for
fishes, waterfowl, and shorebirds rather than
for true seabirds. That situation is not likely
to change in the near future unless additional
funds become available to the Branch. About
the most we can expect to do is designate key
areas as sanctuaries or wildlife management
reserves. Under the folio and referral systems
now operational among resource agencies in
British Columbia, we have the opportunity to
advise other disciplines against approving
practices that would adversely affect wildlife.
By those methods we are attempting to protect
critical seabird habitat. It must be
stressed, however, that we can only advise; we
cannot force other agencies to follow procedures
we suggest.</p>

<p>The only significant work relating to seabirds
in which the Fish and Wildlife Branch is
presently engaged involves inventory of
specific sites on the Queen Charlotte Islands
and the northwest mainland coast. Those
areas are ones on which we expect to find seabird
colonies and where applications for logging
are pending. To enable us to advise the
Forest Service on the wildlife values of those
sites, we began field work in the summer of
1975.</p>

<p>The Federal Government, in comparison to
what it has done on the east coast and in the
north of Canada, has been negligent in its support
of seabird conservation on the west
coast. By far the most seabird research by a
government agency in British Columbia has
been accomplished by the staff at the Provincial
Museum in Victoria. In the past, beginning
in the 1940's, museum personnel (mainly
C. J. Guiguet) explored and inventoried seabird
colonies along the British Columbia
coast. Most work then was exploratory, and
little quantitative information was gathered.
More recently, precise counts have been obtained
of seabirds nesting in the Strait of
Georgia, Juan de Fuca Strait, the central west
coast of Vancouver Island, the northern mainland
coast, and the east coast of the Queen
Charlotte Islands. That information, along
with quantitative data gathered in the summer
of 1975, will be used to update the "Catalogue
of British Columbia Seabird Colonies"
published in 1961 by the museum.</p>

<p>The museum has a number of programs
under way.</p>

<p>&#8226; A cooperative survey with Washington
State of colonies of the double-crested cormorant
<em>(Phalacrocorax auritus)</em> in the Pacific
Northwest. To limit disturbance, that survey
is to be conducted at 5-year intervals beginning
in the summer of 1975.</p>

<p>&#8226; A survey of all islands, whether or not
they are supporting seabirds, in the Strait of
Georgia and Juan de Fuca Strait in 1980, to
detect changes in populations after 1974.</p>

<p>&#8226; Monitoring changes in seabird populations
along the west coast of Vancouver Island,
gathering data for all islands there. Permanent
quadrats will be established on ecological
reserves in the area to help detect such
changes. As a result of such quadrats having
been set up in 1967 on Cleland Island and
being re-examined in 1974, we can document a
significant decrease in Leach's storm-petrel
<em>(Oceanodroma leucorhoa)</em> and a corresponding
increase in rhinoceros auklet <em>(Cerorhinca
monocerata)</em>.</p>

<p>&#8226; Mapping vegetation substrate as it relates
to seabird populations on selected islands in
the Province.</p>

<p>&#8226; Investigating differences in eggshell thickness
between eggs within clutches of glaucous-winged
gulls <em>(Larus glaucescens)</em> near
Victoria.</p>

<p>&#8226; A saturation banding program for cormorants
(<em>Phalacrocorax penicillatus</em>, <em>P. pelagicus</em>,
and <em>P. auritus</em>) on south-coast colonies.</p>

<p>&#8226; Continued banding of select colonies of
glaucous-winged gulls which began in the
1960's. Life tables, survivorship curves, and
dispersal patterns should result.</p>

<p>The museum also acts as a repository for information
on seabirds in British Columbia
and maintains files on the history of seabird
islands as well as references to literature published
on all seabirds in the Province. The ref<span class="pagenum"><a name="Page_249" id="Page_249">[Pg 249]</a></span>erences
include unpublished theses and reports.
This information is easily retrievable&mdash;not
a small contribution in today's paper-producing
society.</p>

<p>Future programs planned by the Provincial
Museum, in addition to the continuance of
some of those already mentioned, include a
system of monitoring colonies every 5 to 10
years, depending on the sensitivity of the
species involved, to detect changes in population
numbers and distribution. It is also
hoped that the first complete provincial census,
with cooperation from Federal and provincial
agencies, naturalist groups, and the
like, can be budgeted and arranged for in the
summer of 1980. That census could conceivably
be expanded to include the entire Pacific
coast of North America.</p>

<p>Some research on the breeding biology of
seabirds has been conducted by universities,
notably the University of British Columbia
under the guidance of R. H. Drent and M. Udvardy.
We expect that graduates returning to
coastal universities will continue that work.
The section of government dealing with ecological
reserves has just recently received
funding to permit field studies on reserves
harboring marine birds. J. B. Foster, Coordinator
of Ecological Reserves, emphasizes that
research by other agencies is encouraged
under permit on ecological reserves.</p>

<p>There are a number of threats to seabirds in
British Columbia. Along with the chemical
pollutants in their environment and food, logging,
and the specter of huge oil tankers plying
the west coast, we are greatly concerned
by the potential threat of boating enthusiasts
and recreationists. Well-meaning but uninformed
vacationers and boaters stopping to
visit or picnic at seabird islands can do serious
damage to nesting seabirds. The possibility of
loss of habitat to seabirds from people searching
for island summer homes poses a threat,
and indeed some seabird islands have already
been lost to speculators. With increased
leisure time and travel the potential of unintentionally
introducing predators, such as
rats (<em>Rattus</em> spp.) and snakes, to seabird islands
is great. Intentional or accidental introduction
of mammals, such as mink <em>(Mustella
vison)</em>, rabbit (<em>Sylvilagus</em> spp.), fox (<em>Vulpes
fulva)</em>, and raccoon <em>(Procyon lotor)</em>, to islands
is another serious threat to the future
existence of seabird populations. The recent
unauthorized and apparently unsuccessful introduction
of mink on the Queen Charlotte Islands
could have resulted in the eventual
devastation of seabird colonies there and on
adjacent islands. The destruction of habitat
by logging near colonies on large islands and
complete logging on small offshore islands
will no doubt adversely affect some seabird
populations. Competition between increasing
numbers of gulls (<em>Larus</em> spp.) and certain
species of seabirds (e.g., storm-petrels and
cormorants) may result in reduced numbers of
the seabirds.</p>

<p>What types of programs are needed? About
80% of all known seabird colonies in British
Columbia have been investigated to date, and
a modest program to monitor changes has
been established. We do, however, require exploratory
work along the west coast of the
Queen Charlotte Islands and northern mainland
coast. We need to know more about the
breeding biology and reproductive potential
of each of the species nesting in the Province,
as well as about their adaptability to different
habitats. Will some burrow-nesting alcids use
man-made tubes erected in otherwise marginal
habitat? Can and should more man-made
habitat be created for cormorants that
have been displaced from ancestral breeding
grounds?</p>

<p>Of immediate urgency is exploratory work
involving seasonal distribution, abundance,
and flight lanes of pelagic seabirds along the
coast of British Columbia&mdash;especially the
northern portion. We lack the base-line data
which could help influence routes of oil
tankers to lessen the potential danger of spills
to marine birds. We know little about the winter
distribution of marine birds, especially
alcids.</p>

<p>As a general rule, offshore islands of less
than 100 ha should be protected completely
from logging, and the larger ones supporting
major seabird colonies should have some protection
from development. We must also consider
the possibility of preserving some islands
which may act as buffer areas and provide
potential alternate habitat to seabirds.</p>

<p>Another concern is the effect of commercial
and sport fishing in the Province on food supplies
for seabirds, and what damage, if any,
gillnetting may have on diving seabirds. Per<span class="pagenum"><a name="Page_250" id="Page_250">[Pg 250]</a></span>haps
we should discourage fishing by nets in
areas where large numbers of seabirds aggregate
to feed.</p>

<p>We also need to know more about the effects
of chemical pollutants on individual
species and on their reproduction. Of paramount
importance, and one which biologists
tend to neglect, is communication among all
disciplines interested in seabirds. For
example, a comprehensive file of the history
of seabird colonies in British Columbia is established
at the Provincial Museum. It would
be a waste of time and money to duplicate
that file and have three or four scattered
across the country. We would be better advised
to tackle another phase of work yet to
be accomplished. Communication assures
that seabirds benefit and are not unduly
harassed.</p>

<p>Annual meetings, both local and international,
of persons interested in marine birds
should be arranged so that problems relating
to seabirds can be discussed. For example,
populations of glaucous-winged gulls in
British Columbia have increased exponentially
in the past 10 years. If they are a threat
to the existence of other seabirds (e.g.,
Leach's storm-petrel, double-crested cormorants),
should they be controlled, and, if so,
how? Such meetings would also help develop a
pattern of universal census methods and techniques
that could be put to use along the
Pacific Coast to provide comparable data
from different areas.</p>

<p>Finally, in today's world, natural resource
agencies must operate on limited funding.
How can one convince administrators to divert
a significant portion of those funds to the
investigation of species that are widely regarded
as having little social importance?</p>

<p>A detailed bibliography of seabirds of
British Columbia is available from either of
us.</p>

<p>We thank D. F. Hatler, J. B. Foster, and
A. L. Allen for comments on the manuscript.</p>

<p><span class="pagenum"><a name="Page_251" id="Page_251">[Pg 251]</a></span></p>
<div class="chapter"></div>
<hr class="chap" />




<h3>Petroleum Industry's Role in Marine Bird Conservation</h3>


<p class="center">by</p>

<p class="center">Keith G. Hay</p>

<p class="center">American Petroleum Institute<br />
2101 L Street NW<br />
Washington, D.C. 20037
</p>


<h4>Abstract</h4>

<div class="blockquot">

<p>Despite improved safety practices, engineering, and navigational skills, marine
tanker transportation will not be 100% accident free. The industry seeks to mitigate
wildlife losses through improved technology, research in the rehabilitation
of species exposed to oil, and the development of oil spill/wildlife contingency
plans.</p></div>

<p>Oil spills and marine birds not only constitute
a deadly mix but have proved to be one of
our toughest environmental problems to
solve. The rehabilitation of these tragic victims
is plagued with controversy, emotion,
apathy, and biological unknowns. The costs
have been high and the survival rates low.
During the last 10 years, a few dedicated
people working here and in Europe have reversed
this trend. They have, in addition,
taken steps to develop contingency plans and
conducted research to reduce seabird mortalities
from oil spills. I present a brief status report
on their progress and the melange of
problems involved.</p>

<p>The unfortunate encounter between spilled
oil and marine birds is not new. It goes back
at least to the turn of the century, when coal-burning
steamships and sailing clippers were
replaced by oil-fueled vessels. Since then thousands
of marine birds have succumbed to
floating oil, especially during World Wars I
and II (Blanks 1942) and in recent spills here
and off the coast of Europe (Clark 1969).</p>

<p>With the current and projected demands for
energy in the United States and with expanded
tanker traffic and accelerated development
of offshore petroleum reserves, the
oil-contaminated ("oiled") bird is not going to
go away. Periodically, this ugly problem will
arise, despite the efforts of the petroleum industry
to improve its safety practices, engineering,
and navigational skills. Unfortunately,
the problem is the product of the inherent
fallibility of man and his imperfect
machines.</p>

<p>We cannot ignore the situation. We must
here, as elsewhere, improve our technology
and mitigate the impact.</p>

<p>A study of more than 100 spills that occurred
throughout the world between 1960
and 1971 revealed that about 1 in 5 spills
(20%) involved 50 or more birds (Ottway
1971). Nearshore spills have a far greater effect
on waterfowl than do spills occurring
several miles or more offshore.</p>

<p>In the 1967 <em>Torrey Canyon</em> tanker spill,
some 8,000 oiled birds were rescued. About
6,000 were picked up alive in England and
about 2,000 in France, at a cost estimated at
$160,000 (Clark 1969; Bourne 1970). Less
than 5% of those treated by British authorities
survived for release some months later.
The survival rate of those rescued in France is
unknown.</p>

<p>In 1969 the Santa Barbara spill resulted in
the treatment of 1,575 marine birds, of which
169 were eventually released. Many of those
released were found dead within a short time
(Smail 1971).</p>

<p>In 1970 the tanker <em>Delian Apollon</em> was responsible
for a spill in Tampa Bay, Florida.
Thousands of seabirds were lost. No exact
count was taken, but hundreds of birds were<span class="pagenum"><a name="Page_252" id="Page_252">[Pg 252]</a></span>
cleaned and farmed out for rehabilitation. Reports
show that many of the birds were returned
dead within a few days (Smithsonian
Institution 1971).</p>

<p>In 1971, when two tankers collided under
the Golden Gate Bridge at the mouth of San
Francisco Bay, the resulting spill involved
some 4,686 oiled birds taken to cleaning centers
(Lassen 1972). Eight months later the
last of 200 survivors (less than 5%) were released
at a cost estimated at $900 per bird
(Smith 1975).</p>

<p>The most vulnerable species involved in
spills have been the oceanic birds such as the
alcids&mdash;murres (<em>Uria</em> spp.), auks (<em>Pinguinus</em>
spp., <em>Alca</em> spp.), puffins (<em>Fratercula</em> spp.,
<em>Lunda</em> spp.), and guillemots (<em>Cepphus</em> spp.).
Other species less affected included ruddy
ducks <em>(Oxyura jamaicensis)</em>, scaup (<em>Aythya
marila</em>, <em>A. affinis</em>), scoters (<em>Melanitta</em> spp.),
mergansers (<em>Lophodytes</em> spp.), oldsquaws
(<em>Clangula</em> spp.), and goldeneyes (<em>Bucephala</em>
spp.). Grebes (<em>Podiceps</em> spp.), eiders (<em>Polysticta</em>
spp.), loons (<em>Gavia</em> spp.), and cormorants
(<em>Phalacrocorax</em> spp.) are also frequently
involved. Ruddy ducks and scaup are
particularly vulnerable during winter on large
river systems with heavy oil transport traffic.
Fortunately, none of the above species have
been reported in jeopardy as a result of spills
in American waters.</p>

<p>In Europe and South Africa, however, it is
believed that oil pollution is responsible for a
steady decline in seabird colonies. For
example, in known oil-dumping areas in the
Baltic Sea, where some mortality of oldsquaws
has been associated with surface oil,
their population has dropped to about one-tenth
of the pre-World War II level (Bergman
1961). Other authors report that oil spills
have reduced the number of scoters in the Baltic
and off southeast England (Atkinson-Willes
1963). The auk populations off the
coast of England have been reported to be
substantially decreased by oil pollution (Parslow
1967). Tankers traversing South Africa's
Cape of Good Hope are said to be responsible
for the reduction of jackass penguins,
<em>Spheniscus demersus</em> (Rowan 1968). Oil pollution,
especially sustained pollution, has thus
been cited as a limiting factor on certain seabird
populations.</p>

<p>Estimates of seabird mortalities from an oil
spill are imprecise; they may differ by thousands
of birds. It is believed that only a small
fraction of the birds killed in a spill wash up
on the shore. Some authors have even speculated
that the death rate at sea could range
from 6 to 25 times the number washed ashore
(Tanis and M&ouml;rzer-Bruyns 1968).</p>

<p>In contrast to terrestrial birds and semiaquatic
species (e.g., ducks; geese; coots,
<em>Fulica</em> spp.; or gulls, <em>Larus</em> spp.), totally seaborne
species have a restricted reproductive
potential. Many, such as the alcids, do not
breed until they are 3 or more years old, and
lay only one egg per year. Only one in five survives
to go to sea.</p>

<p>Until about 5 years ago we knew little about
seabirds. They are not game species (they
taste fishy) and thus do not constitute an important
economic resource. They have never
been the subject of intensive waterfowl management
or research by either State or
Federal governments.</p>

<p>During the last 5 years a small group of
people here and in England have been studying
marine birds&mdash;their distribution, population
status, physiology, diseases, and husbandry
in captivity. Four organizations have
primarily been involved: The American Petroleum
Institute (API); the Wildlife Rehabilitation
Center at Upton, Massachusetts; England's
Advisory Committee on Oil Pollution
of the Sea; and the International Bird Rescue
Research Center in Berkeley, California. They
have encountered many common biological
and people problems, some of which I discuss
here.</p>


<h4>Biological Problems</h4>

<p>The recuperation record for oiled seabirds in
the past has admittedly been dismal. A few
birds have been returned to nature, but only
after a long and costly period of care. In the
process, semidomestication often takes place.
The percentage of cleaned birds that actually
survive after release is even smaller. One
should not infer from this small percentage
that rehabilitated birds cannot readjust to life
in the wild. Several successful reintroductions
have been documented. U.S. Fish and Wildlife
Service bands were returned from two western
grebes that were cleaned and released<span class="pagenum"><a name="Page_253" id="Page_253">[Pg 253]</a></span>
after the 1971 San Francisco spill. One bird
was picked up a year later near Treasure Island,
California, and the second after almost 2
years, in the State of Washington (Fletcher
1973).</p>

<p>Survival rates have zoomed with recent
strides in cleaning technology and husbandry.
The International Bird Rescue Research Center
reported a survival rate of 41%, based on
hundreds of birds and about 20 different
species over a 2-year period (Smith 1975). In
South Africa, where powdered clay was used
as a cleaning agent on jackass penguins,
nearly 50% survived, although exact percentages
have not been published (Edwards 1963;
Holmes 1973). Rapid retrieval, the relatively
small groups of birds treated, and expert
cleaning and husbandry techniques are
largely responsible for high success ratios. Rehabilitation
success is measured not only in
terms of percent survival but also in terms of
median length of captivity and average cost
per bird.</p>

<p>Rescued oiled birds arrive at cleaning centers
under a wide range of physical conditions.
Before capture they may have spent hours or
days in water, during which their energy has
been continuously drained. The oil destroys
the bird's protective insulation, and metabolic
rate must be increased to sustain body temperature.
Constant preening also takes
energy. Food demands increase, but feeding
attempts, especially for diving birds, are
thwarted by oil-fouled plumage. A bird may
arrive at the cleaning center under stress,
chilled, exhausted, dehydrated, starved, and
ill from ingested oil. Cold weather accentuates
these conditions. Often such birds are jammed
together with other species, hauled long
distances, and immediately put through a
series of cleaning processes that would leave
even a healthy bird weak and in a state of
shock. One marvels at the stamina of the
survivors.</p>

<p>In most past spills, every bird found was
routinely cleaned regardless of its condition.
Instead of attempting to reclaim all birds, a
selective judgment should be made. If a bird's
physical condition makes its chances of survival
nearly impossible, it should be humanely
killed (except for rare or endangered species).
This would enable workers to devote more
time and care to birds having a reasonable
chance at survival.</p>

<p>Fletcher (1973) stated that many variables
affect bird survival: weather conditions, the
type and amount of oil in and on the bird, the
species, the distance of the spill from the
shore, the time lag from initial fouling until
initial treatment, the degree of stress a bird is
subjected to, the husbandry techniques used,
the time of release (the sooner released, the
higher the apparent survival), the number of
birds being cared for (the fewer birds being
handled, the higher the survival rate), the
quality of the facilities available, and the
training and experience of the people handling
the birds.</p>

<p>Many of the above biological problems are
under study here and in Europe, including the
following.</p>

<p>&#8226; The effect of ingested oil on the mucosal
transport mechanism of marine birds. To use
seawater, birds must be able to transport
sodium ions through the gut and expel the excess
salt through the nasal passages. Oil can
block the mucosal ion transport mechanism,
resulting in dehydration and eventual death.</p>

<p>&#8226; The development of a successful program
of hormonal and electrolyte therapy to restore
osmotic balance and the functioning of the
salt glands in contaminated seabirds.</p>

<p>&#8226; Treatment and prevention of aspergillosis
(fungus infection); septic arthritis or "bumble-foot"
(joint capsule infections); breast sores
(especially in seabirds confined on hard surfaces);
eye lesions (caused by ammonia fumes
from unsanitary pens); dehydration and hypoglycemia;
lipid pneumonia; and bacterial
infections.</p>

<p>&#8226; Treatment of stress after capture, including
perfection of handling and cleaning techniques,
administration of proper steroids,
crowding, light, temperature, noise levels, and
so on.</p>

<p>&#8226; Development of proper nutritional regimes
for certain species and feeding techniques
to eliminate forced feeding.</p>

<p>&#8226; The establishment of criteria for confident
recognition of terminal pathological conditions
in oiled birds.</p>

<p>&#8226; Determination of optimum density of confined
birds to insure healthy conditions and
adequate room for preening.</p>

<p>&#8226; Determination of proper time and conditions
for reintroduction of the birds into their
native habitat.</p>

<p><span class="pagenum"><a name="Page_254" id="Page_254">[Pg 254]</a></span></p>


<h4>People Problems</h4>

<p>Handling an over-responsive and emotional
army of bird-cleaning volunteers and training
them to play constructive roles is a major
undertaking. Planning, cooperation, understanding,
patience, and clear direction must
be developed. In the absence of these virtues,
chaos can and has prevailed.</p>

<p>The San Francisco Bay oil spill of 1971 was
a classic example. There was virtually no
State or Federal coordination. Splinter
groups of volunteers established their own
"treatment centers" and jealously guarded
their patients. Some actually absconded with
their pet patients to seek better care elsewhere.
Long hours, fatigue, and frustrations
led to dissension and bitter quarrels. Antiestablishment
sentiment was rampant.</p>

<p>Instant experts on bird cleaning, avian
medicine, and nutrition appeared or developed
overnight. Veterinarians volunteered their
services, but their knowledge of oiled-bird
treatment was limited. A wide variety of food
(from canned dog food to live shrimp) was
given the birds. Forced feeding was routine.
Medications and vitamins of all kinds were
also administered. Needless to say, the states
of the art in treating oiled birds and handling
volunteers were both in their infancy. For
both, the success ratio was near zero.</p>

<p>To prevent such fruitless efforts and the
frantic, unorganized response that prevailed,
a well-designed contingency plan for wildlife
involved in an oil spill is needed.</p>


<h4>Contingency Planning</h4>

<p>It is only prudent to take reasonable measures
to prepare for oiled-bird emergencies.
This is especially true in regions where bird
concentrations and oil shipment traffic converge.
Almost equal attention must be devoted
to handling volunteers as to handling
birds. Safety is a major consideration. The
sharp beaks of birds can be very dangerous.</p>

<p>A model State contingency plan should include
the following:</p>

<p>&#8226; A list of State and Federal agencies to be
alerted, including 24-h, 7-day-a-week telephone
numbers, and names of individuals to contact.</p>

<p>&#8226; Clarification of the roles of State and
Federal agencies under the Regional Response
Plan of the National Oil and Hazardous
Substances Pollution Contingency Plan.</p>

<p>&#8226; A list of State and Federal laws pertaining
to possession of birds and mammals.</p>

<p>&#8226; An updated roster should be maintained of
team members, assignments, and responsibilities
for inland and marine spills, including discovery
and notification, record keeping, public
information, containment and counter-measures,
wildlife protection, and cleanup,
restoration and evaluation of effects on the
biota.</p>

<p>&#8226; A list of individuals or organizations that
possess skills and experience in treatment of
oiled birds (locally and nationally).</p>

<p>&#8226; Location of emergency wildlife reception
and treatment centers.</p>

<p>&#8226; A list of the necessary supplies, equipment,
and holding facilities for cleaning, treating,
drying, and post-care operations. Such information
can be obtained from:</p>

<div class="hangindent">

<p>&mdash;California Department of Fish and
Game, Oil and Hazardous Materials
Contingency Plan (July 1974)</p>

<p>&mdash;International Bird Rescue Research
Center, Aquatic Park, Berkeley, California
94710</p>

<p>&mdash;American Petroleum Institute, 2101 L
Street, Northwest, Washington, D.C.
20037</p>

<p>&mdash;Wildlife Rehabilitation Center, 84
Grove Street, Upton, Massachusetts
01568</p></div>

<p>&#8226; An organizational plan which includes assignments
of duties and responsibilities for
personnel manning a bird-cleaning center. In
addition to bird cleaning and husbandry, assignments
must be made for record keeping,
internal communications, public relations,
logistics (supplies), security, sanitation,
safety, and meals.</p>

<p>&#8226; A slide lecture or film to instruct volunteers
in the correct techniques for handling,
cleaning, and post-care of oiled birds.</p>

<p>&#8226; A selected bibliography of key references
on oiled-bird cleaning and care.</p>

<p>&#8226; Appendices to the plan should include
maps of the major coastal oil terminals, bays,
and estuarine areas with heavy oil transport<span class="pagenum"><a name="Page_255" id="Page_255">[Pg 255]</a></span>
traffic. Map overlays would depict the location
of resident species and the migratory patterns,
species composition, relative abundance,
and winter concentration areas of migrants.
Additional overlays would locate commercially
important demersal seafood areas
(e.g., oyster and abalone beds, lobster and
crabbing locales) and marine mammal habitats.
Further refinement of an atlas could include
information on tides, prevailing winds,
ocean currents, and water mass movements to
assist in predicting the path of spilled oil.</p>


<h4>What Has Been Accomplished</h4>

<p>The petroleum industry, through the API,
took prompt steps to mitigate the problem
after the first seabird mortalities were reported
from Santa Barbara in 1969. They
commissioned a young aviculturist, Philip
Stanton, who has extensive experience working
with wild waterfowl, to start a research
program on cleaning and caring for oiled
birds. At his Wildlife Rehabilitation Center at
Upton, Massachusetts. Stanton, with the
help of API, has been conducting research on
oiled birds for 7 years. He is also an assistant
professor of biology at nearby Framingham
State College. Stanton's studies (unpublished)
include investigations on food shape and color
preferences in wild ducks, the effects of
lengthened photoperiods on breeding of arctic
geese, and the effects of diets of varying protein
concentrations on growth and development
of the common eider duck.</p>

<p>As a result of his research on cleaning techniques
and agents, Stanton has recommended
a nontoxic liquid cleaner called Polycomplex
A-11. Although not perfect, it is one of several
cleaning agents being successfully used
today. He has authored a "how to" guide for
oiled-bird treatment entitled "Operation Rescue"
and prepared a companion bibliography
(Stanton 1972). These booklets have been distributed
throughout the United States to
State and Federal agencies and conservation
organizations. He has provided consulting
services at numerous spills and has worked to
establish oiled-wildlife treatment centers in
coastal States.</p>

<p>Since 1972 the API has sponsored an avian
physiology study at the University of California
at Santa Barbara. Under the direction
of W. N. Holmes, the studies are directed at
the effects of ingested crude oil and petroleum
products on marine birds. Holmes has revealed
that small quantities of crude oil introduced
into the gut of a saltwater-adapted bird
can affect the mucosal transport and extra-renal
excretory mechanisms, resulting in
acute dehydration and eventual death. Dr.
Holmes is also examining the effects of the
various distillation fractions derived from
crude oil and the long-term effects of ingested
oil in mature birds. Incidentally, Alaska
North Slope oil was found to be almost innocuous
when administered to ducklings in
amounts similar to the effective doses of other
oils (Holmes and Cronshaw 1975).</p>

<p>Refined products (diesel oil, No. 2 fuel oil,
and Bunker "C") are known to be more toxic
than crude oil. For example, the relatively
small spills of Bunker "C" at Tampa, Florida,
in 1970 and in San Francisco in 1971 caused
approximate mortalities of 90 and 20 birds
per ton of spilled product, respectively. The
crude oil spills of the <em>Torrey Canyon</em> and at
Santa Barbara, however, resulted in mortalities
of only 0.5 and 0.6 bird per ton of oil
(Clark 1973).</p>

<p>Dr. Holmes is now testing measured
amounts of the above refined oils on adult
birds. He is determining the degree of dehydration
incurred, the resulting pathological
changes, and the replacement (hormonal and
electrolyte) therapy necessary to rehabilitate
the birds.</p>

<p>It is obviously important to keep as many
birds away from an oil slick as possible. This
was the objective of an API contract with the
Av-Alarm Corporation of Santa Maria, California.
Their objective was to determine the
feasibility of repelling aquatic birds from an
area by using an acoustical jamming device as
the stimulus.</p>

<p>The flocking instinct in birds provides
mutual protection through their almost constant
communication with one another. When
this (audio) communication is prevented by
jamming with high-frequency sounds, the
birds immediately leave the area to seek relief.
This harmless technique has been used successfully
for years to repel agricultural pest
birds.</p>

<p><span class="pagenum"><a name="Page_256" id="Page_256">[Pg 256]</a></span></p>

<p>The Av-Alarm device was tested on waterfowl
at the Grizzly Island Game Refuge some
48 km north of San Francisco Bay and in the
bay itself over a 2-year period (1972-73). Using
a single, fixed-location system covering a
three-quarter square mile (1.21 km<sup>2</sup>) area
Crummett (1973) repelled 82% of the ducks
and 92% of the shorebirds on the Refuge. The
intrepid coot, however, was found to be relatively
indifferent to the sounds. Immediately
upon activation, there was a sudden drop in
the bird count, which was followed by a continual
decline in numbers.</p>

<p>In tests of the device from a cruising boat in
ocean and bay waters, the degree of effectiveness
varied by species. Ducks were repelled
100%; pelicans (<em>Pelecanus</em> spp.) 92%; great
egrets <em>(Casmerodius albus)</em> 85%; gulls 42%;
cormorants 75%; shearwaters (<em>Adamastor</em>
spp.) 29%; and murres, 51%.</p>

<p>Grebes and murres dived away from the
stimulus, then surfaced and dived again if the
threat was still present. To prevent driving
the diving species deeper into the center of a
slick, investigators recommended that
buoyed repelling equipment be placed within
the spill area. When the alarm system was
used in conjunction with the occasional firing
of a rocket or shellcracker, an even greater
percentage of birds was repelled.</p>

<p>The International Bird Rescue Research
Center, a nonprofit corporation in Berkeley,
California, was an outgrowth of the Richmond
Bird Care Center that played an active role in
the 1971 San Francisco Bay spill. Since that
time, a small group of individuals has continued
research on bird-cleaning techniques,
testing cleaning agents, perfecting husbandry
methods, and alleviating stress. Their 41%
survival rate speaks for itself. A paper describing
their work is being presented at this
conference (Smith 1975).</p>

<p>Under a grant from the API, the center is
currently evaluating various cleaning agents,
and testing the pressurized jet versus serial
baths and the re-establishment of feather
waterproofing. The center is also perfecting
an audio-visual slide presentation that will
illustrate how to select the proper cleaning
agent, together with the latest bird-cleaning
and care procedures.</p>

<p>About 5 years ago, England's Advisory
Committee on Oil Pollution of the Sea established
a research unit in the Department of
Zoology at the University of Newcastle-Upon-Tyne.
It was funded by a grant from the
Royal Society for Prevention of Cruelty to
Animals, the Royal Society for the Preservation
of Birds, the World Wildlife Fund Seabird
Appeal, and the British Institute of
Petroleum.</p>

<p>Their efforts have also led to high survival
rates. Focusing primarily on the efficiency of
various detergents, they have found that the
loss of waterproofing is largely due to soap
and oil residues and the disturbance of the
feather structure in the cleaning process. Consequently,
they have devoted their efforts to
selecting detergents that can be completely
removed with a minimum disturbance of
plumage (Seabird Research Unit 1971).</p>

<p>In May 1974, the API in cooperation with
the U.S. Fish and Wildlife Service convened a
seminar on Oil Spill Wildlife Response Planning.
The 2-day workshop was held at the
Patuxent Wildlife Research Center at Laurel,
Maryland. Some 70 State and Federal government
personnel in charge of oil spill response
plans involving wildlife participated. The program
addressed itself to fish and wildlife considerations
and the role of regional response
teams under the National Oil and Hazardous
Substances Pollution Contingency Plan. The
actions of State wildlife departments, U.S.
Fish and Wildlife Service, Environmental
Protection Agency, U.S. Coast Guard, and
the oil industry in handling spills involving
wildlife were examined. The latest oil spill
cleanup technology was reviewed, and the
workshop ended with demonstrations of the
cleaning of oiled waterfowl. Similar seminars
were planned for the Gulf of Mexico and the
West Coast.</p>

<p>It was obvious from this seminar that the
most comprehensive wildlife oil spill contingency
plan had been developed by the State of
California. Copies of this plan (Oil and
Hazardous Materials Contingency Plan, California
Department of Fish and Game, July
1974) were later distributed to all coastal
States as a prototype or model plan by API.</p>

<p>The U.S. Fish and Wildlife Service has been
conducting experiments on various bird-cleaning
agents and techniques at its Migratory<span class="pagenum"><a name="Page_257" id="Page_257">[Pg 257]</a></span>
Bird and Habitat Research Laboratory near
Laurel, Maryland. The Fish and Wildlife Service
is also working with the API in developing
information on migratory patterns and
winter waterfowl concentration areas on the
East Coast as they relate to petroleum transport
traffic and oil terminals.</p>

<p>In Canada, the Petroleum Association for
Conservation of the Canadian Environment
(PACCE) employed the services of a consulting
firm to make a comprehensive review of
dispersal and rehabilitation of waterfowl associated
with oil spills. The resulting PACCE report
(LGL Ltd. 1974) codified what was
known about the problem, identified research
needs, and developed effective wildlife oil-spill
contingency plans for critical areas on
Canada's east and west coasts, the Great
Lakes, and the Arctic.</p>

<p>The Florida Game and Fresh Water Fish
Commission has initiated a program for the
rehabilitation and treatment of oiled birds. It
is being organized by veterinarian Harold F.
Albers of St. Petersburg. He is working in cooperation
with the Florida Associated Marine
Institutes, the Shell Oil Company, Clean Gulf
Associates, and the API.</p>

<p>The Standard Oil Company of California
provided a grant to James Naviaux of
Pleasant Hill, California, to develop bird-cleaning
technology, including the testing of
various cleaners. Dr. Naviaux had treated
birds from the 1971 San Francisco spill. A
publication on the after-care of oil-covered
birds (Naviaux 1972) resulted from the collaboration
with Alan Pittman, research
chemist of the U.S. Department of Agriculture's
Western Research Laboratory.</p>

<p>In 1971, the API in cooperation with the
National Wildlife Federation (NWF) initiated
an NWF/API Fellowship program. One of the
first grants under this program was to
Charles W. Kirkpatrick, Professor of Wildlife
Management at Purdue University. He and
assistants studied for 4 years the nesting ecology
and productivity of the emperor goose
<em>(Philacte canagica)</em> in the Igiak Bay area of
the Yukon Delta in Alaska (Eisenhauer and
Kirkpatrick 1977).</p>

<p>An extensive program of marine bird research
was initiated on the North Slope of
Alaska by the Atlantic Richfield Company in
1969. It has been continued ever since and includes
the acquisition of extensive base-line
data on all waterfowl, including June surveys
of breeding pair counts and August surveys
for brood counts. The results of these surveys
for 1969-73 are presented by Gavin (1975).</p>

<p>Base-line data on marine birds of the Gulf of
Alaska are currently being collected and compiled
through grants to various universities
and institutions by the American petroleum
industry. These data will constitute elements
of a report on the environmental status of the
Gulf of Alaska. Such information is essential
prior to development of the Gulf's offshore
petroleum resources.</p>


<h4>Marine Mammals</h4>

<p>Most sea mammals are relatively resistant
to oil slicks and tend to avoid contaminated
waters. As a result, little research has been
conducted on cleaning and treatment techniques
except for experiments on live beavers
and on the carcasses and pelts of sea otters
and beavers.</p>

<p>No sea otter or seal has ever been oiled and
subsequently cleaned in an oil spill situation.
It is possible, however, that a spill could have
significant adverse effects on sea otters and
fur seals, especially at a rookery during the
pupping season. These animals depend on an
air blanket trapped in their dense underfur for
warmth and buoyancy. Any form of pollutant,
especially oil, could penetrate the outer guard
hairs and underfur and allow water to reach
the skin, with disastrous effects.</p>

<p>Seals and otters are powerful animals, and
the larger males and females can be quite aggressive
and dangerous. Only professional
wildlife specialists and consulting veterinarians
should be permitted to handle and
treat them. A guide to cleaning and care of
oiled sea otters can be found in the California
Oil and Hazardous Materials Contingency
Plan.</p>


<h4>Conclusions</h4>

<p>This status report has revealed that substantial
efforts and progress have been made
in oiled-wildlife research. New techniques
being developed are leading to higher survival
rates. Preventive measures are being devised
to keep birds from entering a spill area. Wild<span class="pagenum"><a name="Page_258" id="Page_258">[Pg 258]</a></span>
life contingency plans are being developed
and materials to handle future emergencies
are being stockpiled. Basic research is being
continued on the difficult problems inherent
in achieving high survival levels and a rapid
return to the wild, at a reasonable cost.</p>

<p>Much more must be done, but these pioneering
efforts both within and outside of industry
reflect a difficult problem yielding to the
time and attention of dedicated men and
women.</p>


<h4>References</h4>

<div class="hangindent">

<p>Advisory Committee on Oil Pollution of the Sea.
1972. Research unit on the rehabilitation of oiled
seabirds. Committee, Dep. Zool., Univ. of Newcastle-Upon-Tyne,
England, Annu. Rep. 2. 33 pp.</p>

<p>Atkinson-Willes, C. 1963. Wildfowl in Great
Britain. Nat. Conserv. Monogr. 3. 368 pp.</p>

<p>Bergman, G. 1961. The migrating populations of
the long-tailed duck <em>(Clangula hyemalis)</em> and the
common scoter <em>(Melanitta nigra)</em> in the spring,
1960. Suomen Riista 14:69-74.</p>

<p>Blanks, D. W. 1942. Birds in the war zone. Gull
24(4):11.</p>

<p>Bourne, W. R. P. 1970. Special review&mdash;after the
<em>Torrey Canyon</em> disaster. Ibis 112(1):124.</p>

<p>Clark, R. B. 1969. Oil pollution and the conservation
of seabirds. Proc. Int. Conf. on Oil Pollut. of
the Sea 1968:76-112.</p>

<p>Clark, R. B. 1973. Impact of chronic and acute oil
pollution of seabirds. Page 634 <em>in</em> Background
papers for a workshop on inputs, fates and effects
of petroleum in the marine environment, Vol. 11.
Ocean Board National Academy of Science,
Washington, D.C.</p>

<p>Crummett, J. G. 1973. Bird dispersal techniques for
use in oil spills. Final report. American Petroleum
Institute, Washington, D.C.</p>

<p>Edwards, R. A. 1963. Treatment of washed up penguins.
Bokmakierie 15(1):8.</p>

<p>Eisenhauer, D. I., and C. M. Kirkpatrick. 1977.
Ecology of the emperor goose in Alaska. Wildl.
Monogr. 57. 62 pp.</p>

<p>Fletcher, A. 1973. Why save oiled birds? University
of California, College of Forestry, Berkeley.
26 pp. (Unpublished report)</p>

<p>Gavin, A. 1975. Wildlife of the North Slope, a five
year study, 1969-73. Atlantic Richfield Co.,
Anchorage, Alaska. 63 pp.</p>

<p>Holmes, M. 1973. Oil and penguins don't mix. Natl.
Geogr. Mag. 143(3):384-397.</p>

<p>Holmes, W. N., and J. Cronshaw. 1975. Final progress
report on studies completed, 1972-75, on the
effects of petroleum on marine birds. American
Petroleum Institute, Washington, D.C. 77 pp.
(Unpublished report)</p>

<p>LGL Limited. 1974. Review of current knowledge
on reducing bird mortalities associated with oil
spills. Petroleum Association for Conservation of
the Canadian Environment Rep. 75-4. 50 pp.</p>

<p>Lassen, R. W. 1972. Waterbirds and the San Francisco
oil spill. Proc. Cal-Nev Wildl. 1972:20-24.</p>

<p>Naviaux, J. L. 1972. Aftercare of oil covered birds.
National Wildlife Health Foundation, Pleasant
Hill, Calif. 52 pp.</p>

<p>Ottway, S. M. 1971. A review of world oil spillages,
1960-1971. Oil Pollut. Res. Unit, Orielton Field
Centre, Pembroke, Wales.</p>

<p>Parslow, J. L. F. 1967. Changes in status among
breeding birds in Britain and Ireland. Br. Birds
60:2-47, 97-122, 177-202.</p>

<p>Rowan, M. K. 1968. Oiling of marine birds in South
Africa. Pages 121-124 <em>in</em> Proc. Int. Conf. on Oil
Pollut. of the Sea.</p>

<p>Seabird Research Unit. 1972. Advisory Committee
on Oil Pollution of the Sea Research Unit on the
Rehabilitation of Oiled Seabirds. Second annual
report. Dep. Zool., Univ. of Newcastle-Upon-Tyne,
England. 33 pp.</p>

<p>Smail, John. 1971. The oil spill in retrospect. Point
Reyes Bird Observatory News 18:1-2.</p>

<p>Smith, D.C. 1975. Rehabilitating oiled aquatic
birds. Pages 241-247 <em>in</em> Proc. 1975 Conf. Prevention
and Control of Oil Pollution.</p>

<p>Smithsonian Institution. 1971. Annual report of
the Center for Short-lived Phenomena, 1970.
Cambridge, Massachusetts.</p>

<p>Stanton, P. B. 1972. Operation rescue. American
Petroleum Institute, Washington, D.C. 32 pp.</p>

<p>Tanis, J. J. C., and M. F. M&ouml;rzer-Bruyns. 1968. The
impact of oil-pollution on seabirds in Europe.
Proc. Int. Conf. on Oil Pollut. of the Sea. 1968:67-76.</p></div>

<p><span class="pagenum"><a name="Page_259" id="Page_259">[Pg 259]</a></span></p>



<p><span class="pagenum"><a name="Page_260" id="Page_260">[Pg 260]</a><br /><a name="Page_261" id="Page_261">[Pg 261]</a></span></p>
<div class="chapter"></div>
<hr class="chap" />




<h2><a name="CONSERVATION_OF_MARINE_BIRDS_IN" id="CONSERVATION_OF_MARINE_BIRDS_IN">CONSERVATION OF MARINE BIRDS IN
OTHER LANDS</a></h2>
<div class="chapter"></div>
<hr class="chap" />




<h3>Conservation of Marine Birds in New Zealand</h3>


<p class="center">by</p>

<p class="center">Gordon R. Williams</p>

<p class="center">New Zealand Wildlife Service<br />
Department of Internal Affairs<br />
Wellington, New Zealand
</p>


<h4>Abstract</h4>

<div class="blockquot">

<p>Marine species (pelagic birds and those of exposed coasts) make up about 48%
of New Zealand's native avifauna, excluding stragglers and antarctic species.
The biological history that has led to the present status of marine birds in this
archipelago of some 700 islands is outlined, methods of conservation are briefly
described, and some illustrative case histories of management programs are
given. In spite of the major environmental changes that have occurred in New
Zealand during 200 years of European occupation, only one marine species has
become extinct, although five such endemic species are currently regarded as
threatened as are a few subspecies of widely distributed forms.</p></div>

<p>New Zealand, which lies some 2,000 km
southeast of Australia, has been a changing
archipelago for many millions of years. It has
been separated from any major landmass
(first, Gondwanaland and later, Australia) for
at least 80 million years.</p>

<p>Before the arrival of man, probably between
1,000 and 1,500 years ago, New Zealand was
free of any land mammals except two species
of bats, and there were few avian predators.
These, among a number of other biological
peculiarities, reflect the archipelago's considerable
and long-standing isolation.</p>

<p>There are nearly 700 islands 0.5 ha or more
in area in the New Zealand region; and, if
North, South, and Stewart islands are regarded
collectively as the mainland, about
650 of these islands lie within 50 km of the
coast and 30 beyond that limit, to about
850 km offshore (Atkinson and Bell 1973).
The archipelago extends from about 30&deg; to
52&deg;S lat. (over a distance of about 2,400 km)&mdash;that
is, from the subtropical to the subAntarctic&mdash;and
from about 166&deg; to 176&deg;W
long. (Fig. 1).</p>

<p>Pelagic and coastal birds must obviously be
an important part of the avifauna and, in fact,
aside from stragglers, antarctic species, and
established introduced species, they make up
about 48% of the 173 in the New Zealand
Checklist (Kinsky 1970). Of the 83 species I
have regarded as marine, 48 (28%) are pelagic
and 35 (20%) shorebirds of exposed coasts.
Ten of the 48 pelagics (21%) and 12 (34%) of
the 35 shorebirds are endemic.</p>

<p>More than a thousand years of occupation
by Polynesian man with his commensal Polynesian
rats <em>(Rattus exulans)</em> and a peculiar
breed of domesticated and feral dog (now extinct),
did little damage to pelagic and open
coast species, even though many, if not most,
were used as food&mdash;especially the petrels, and
particularly those belonging to the genera
<em>Puffinus</em>, <em>Procellaria</em>, and <em>Pterodroma</em>. However,
the Europeans, who arrived about 200
years ago, brought with them a menagerie of
mammals and birds, and 33 species of each
have become established and are now feral
(Gibb and Flux 1973; Williams 1973). They
also put into practice, on a large scale, European
methods of land use that had unfortunate
effects on almost the entire native avifauna.
Although terrestrial, freshwater, and
estuarine species suffered most, marine
species suffered also. However, reduction in
numbers and range rather than extinction
was the rule, except locally.</p>

<p>Apart from habitat destruction by man and<span class="pagenum"><a name="Page_262" id="Page_262">[Pg 262]</a></span>
the various mammalian browsers and grazers,
the most inimical agents have been black rats
<em>(Rattus rattus)</em>, Norway rats <em>(R. norvegicus)</em>,
feral cats, and feral pigs. One would expect
the inhospitality or inaccessibility of an island
to be a marine species' best protection, and so
it has generally proved-the greatest losses
have occurred on the two major mainland islands
(North Island and South Island).
Bourne (1967) suggested that Polynesians in
pre-European times may have caused the extinction
of numerous petrels in the Chatham
Islands. There are still a few islands on which
no exotic mammals occur, but modern transport,
allied with human curiosity and
cupidity, are stripping all but the most wild
and remote of these of the protection against
invasion they have had so far. Cruises by
nature-hungry but sometimes environmentally
illiterate tourists are beginning to be
a local problem.</p>

<div class="figcenter" style="width: 491px;">
<img src="images/p_262.jpg" width="491" height="700" alt="" />
<div class="caption"><p>Fig. 1. New Zealand and its main offshore and outlying
islands (from Atkinson and Bell 1973).</p></div>
</div>

<p>The matter of conservation of marine
species in New Zealand has stemmed mainly
from the recognition of the value of certain islands
as refuges for whole ecosystems, as convenient
areas for study, and as arks for the
rescue of the threatened species that can be
successfully established on them&mdash;an often
highly hazardous and uncomfortable procedure
for men as well as birds.</p>


<h4>Conservation Measures</h4>

<p>By statute, all feral species of birds in New
Zealand are automatically protected unless
specifically legislated for otherwise. (About
50 of our grand total of 285 species have been
so legislated for.) One fortunate consequence
of this provision is that all new arrivals&mdash;vagrants
or new discoveries&mdash;are also fully protected.
The legislation also states that it is
illegal to have in one's possession the nests,
eggs, feathers, skins, or bones of any fully
protected species unless one has been issued a
permit for this purpose. This restriction may
apply to institutions as well as to persons.</p>

<p>After this good start and the setting aside
of conservation reserves of various kinds, active
conservation measures depend on making
careful and comprehensive surveys of the
species and its ecosystem&mdash;often none too
easy a task in the New Zealand region because
of the rough seas, the relative inaccessibility
of many of the important islands and their
ruggedness, and the near-impenetrability of
some of the vegetation types they support.
Having decided that positive action is necessary,
the next step is to use all available
media to inform the public (local as well as national,
if the island is inhabited) of the situation
and the proposals for remedying it. As in
most other countries, uninformed emotionalism
is one of the most pervasive and serious
obstacles to effective conservation because of
the political pressure it can generate.</p>

<p>Apart from formal ecological studies, the
New Zealand Wildlife Service uses three main
methods to support threatened species (other
than the attempts we are making to breed certain
freshwater and terrestrial species in
captivity):</p>

<p>&#8226; The translocation and founding of new
colonies in promising or unmodified habitat.
Such habitats are not common in New Zealand
because of the ubiquity of the introduced<span class="pagenum"><a name="Page_263" id="Page_263">[Pg 263]</a></span>
mammalian browsers, grazers, and predators
(Williams 1977).</p>

<p>&#8226; The destruction, or at least the reduction,
of such browsers, grazers, and predators by
physical, chemical, or biological methods, or
combinations of these.</p>

<p>&#8226; The exertion of social influences to promote
changes in methods of land use or in traditional
harvest for food (the latter can be particularly
important as far as the Polynesian
[Maori] population is concerned, as nowadays
the taking of birds for food is predominantly a
cultural rather than an economic matter).</p>

<p>Translocation has been a valuable technique
for increasing the numbers and ranges of a
few threatened terrestrial species. The very
nature of most marine species, however,
limits its application as far as they are concerned.
Nevertheless, we have considered it
worth trying for one nonmigrant wader; and
no doubt it could be tried under similar circumstances
elsewhere.</p>

<p>Convincing local experiences have shown
that predator or competitor destruction is
likely to be practical only on small, not-too-rugged
islands, usually no larger than about
500 ha. However, special circumstances have
prompted us to attempt destruction, or at
least control, on much larger and more difficult
islands. It is implicit that the predators
or competitors are exotic, not indigenous. Recently,
on those rare islands that are inhabited
but still free of either black or Norway
rats, we have set up permanent bait stations
(at which sodium fluoroacetate&mdash;"1080"&mdash;is
used as the poison) on wharves and jetties in
the hope that such a precaution will, with the
addition of a propaganda campaign calling for
the regular fumigation of visiting vessels, prolong
the charmed lives that these fortunate islands
have so far enjoyed. It goes without
saying that we ask that the greatest care be
taken when expeditions land stores on uninhabited,
rat-free islands which, if by "rat-free"
we mean also free of <em>R. exulans</em>, are even
rarer in our seas.</p>

<p>The sociolegal approach is effective only
when ecosystems or communities have not
been seriously modified, otherwise it is no
substitute for either of the other two measures
discussed.</p>


<h5>Some Case Histories</h5>


<h6>Translocation</h6>

<p>Last century, an endemic monotypic genus
of wader&mdash;the New Zealand shore plover <em>(Thinornis
novaeseelandiae)</em>&mdash;was widespread and
occasionally very common around the coasts
of the North and South islands and the Chatham
Islands. As a result of European settlement
and the accompanying predation by
feral cats and rats, the species now occurs
only on South East Island in the Chatham
group (860 km east of the mainland), where it
at present seems safe, since there are no rats
on the island and it is now a reserve. However,
the population numbers only about 120 individuals.
Because calamities can always occur
(for example, ship rats recently reached shore
on three important islets off the southwest
coast of Stewart Island), the Wildlife Service
is anxious to spread the shore plover to other
suitable islands, if they can be found. The
species is not a migrant and is rather sedentary.
The first translocation attempts failed,
probably because mainly adult birds were
used, and we are now continuing our studies
of the species with the thought in mind,
among others, that success may come if
young birds are used instead; the question
is&mdash;how young?</p>

<p>As is widely known, the New Zealand Wildlife
Service has been remarkably successful in
recent years in translocating one species of
the endemic wattlebird family&mdash;the forest-dwelling
saddleback <em>(Philesturnus carunculatus)</em>&mdash;to
other islands than the four small
ones it had been reduced to by the early
1960's; three of these islands were the ones recently
invaded by ship rats, referred to above.</p>


<h6>Predator Control</h6>

<p>Some 25 km off the North Island's east
coast lies the 3,000-ha, very rugged and
forested Little Barrier Island, which has now
been a reserve for the protection of flora and
fauna for about 80 years. Before that, it had
been almost continually occupied by Maoris
since their arrival in New Zealand, and about
one-third of its forest was felled or burnt, especially
after European settlement of the adjoining
New Zealand mainland began.</p>

<p><span class="pagenum"><a name="Page_264" id="Page_264">[Pg 264]</a></span></p>

<p>Most unusually, Little Barrier is now free of
any grazing or browsing mammals, and has
only the Polynesian rat (a reminder of the
Maori occupation) and feral cats (a European
legacy) to impair its extreme importance as a
reserve. The rats have been unmolested by
man because, rightly or wrongly, they are considered
ineffective predators generally; however,
their impact has probably been under-rated.
More than half a century of trapping
and hunting of cats by successive caretakers
on the island has not effectively reduced that
population.</p>

<p>Among its other important attributes,
Little Barrier supports two birds endemic to
New Zealand&mdash;the rare black petrel <em>(Procellaria
parkinsoni)</em>, and one endemic honey-eater,
the stitchbird <em>(Notiomystis cincta)</em>,
which was once widespread on the North Island
but is now found only on Little Barrier in
moderate numbers, and apparently in no immediate
danger. The impact of feral cats on
stitchbirds has not been determined, but it is
known that cats are seriously affecting the
black petrel especially: they kill at least 90%
of the chicks and some adults annually. Their
impact on a locally remnant population of
Cook's petrels <em>(P. cookii)</em> is apparently less
severe.</p>

<p>In 1968-69 the Wildlife Service, with veterinary
advice and assistance, added an attempt
at biological control to the campaign of poisoning
("1080" in fish was the poison and bait
used), trapping, and shooting. The very
specific viral disease&mdash;feline enteritis&mdash;was introduced
by trapping island cats, infecting
them, and then releasing them. Some estimates
of the resulting mortality from the
combined techniques were as high as 90%; but
there has been a recovery since, and the campaign
is expensive in both time and man-power.
And, oddly enough, the control effort
has met with some opposition. Nevertheless,
another campaign is planned.</p>


<h6>Habitat Rehabilitation by
Destruction of Mammals</h6>

<p>The Kermadecs are a group of small islands
about 800 km north-northeast of the North
Island. Their biological significance, insofar
as this symposium is concerned, is that they
are the southernmost breeding area in New
Zealand seas for many elements of the Pacific
tropic and subtropical marine avifauna. Unfortunately,
goats were liberated on the two
largest islands&mdash;Raoul (3,000 ha) and
Macauley (300 ha)&mdash;almost 150 years ago and
Macauley Island was burnt over; such forest
cover as it had was severely damaged or destroyed,
probably at about the same time. The
goats were to be an emergency food supply for
whalers and shipwrecked mariners. Cats, too,
became feral on Raoul Island during one of its
fitful periods of occupation. The New Zealand
Wildlife Service, in spite of the distance and
difficulties involved, has undertaken pest destruction
campaigns on both islands, but I
offer here only an account of the simpler, and
more successful, Macauley operation.</p>

<p>In 1966, a 5-week expedition to this waterless
and almost treeless island resulted in the
shooting of what was then thought to be all of
its 3,000-odd goats (a density of about 15/ha).
Four years later, a follow-up expedition found
and destroyed another 17 goats (a later brief
inspection suggested that these were indeed
the last), and rehabilitation of the island is
well under way. Now that the short turf is disappearing,
erosion of the soft volcanic soils is
reduced. With compaction no longer occurring,
it will be interesting to see what the effect
will be on birds breeding on the island&mdash;six
breeding species of petrels, three breeding
species of terns, and other marine species.</p>


<h6>Sociolegal Conservation</h6>

<p>The taking of petrels and other procellariiform
birds for food has always been part of
the Polynesian economy and culture throughout
the Pacific. In New Zealand, the practice
now has only minor economic importance, but
it is still an essential part of Maori culture and
tradition. The most commonly taken species
are the sooty shearwater <em>(Puffinus griseus)</em>
and, until recently, the gray-faced petrel
<em>(Pterodroma macroptera)</em>. Although no formal
study of the impact of the annual harvest of
chicks on the population has yet been made,
all the indications are that it is not significant.
Nevertheless, the Maoris willingly accepted
the limited amount of legislation that
has been passed to afford the two principal exploited
species at least token protection. However,
on the Chatham Islands, where there is a
strong tradition of taking some of the albatrosses,
this tradition has persisted, even<span class="pagenum"><a name="Page_265" id="Page_265">[Pg 265]</a></span>
though all albatrosses are fully protected
throughout New Zealand.</p>

<p>Enforcement of legislation in small and isolated
communities is not always easy and
sometimes may not be wholly politic. However,
the Maoris of the Chathams have been
specially informed of the conservation issues
at stake, and a "gentleman's agreement" has
been reached: If a planned survey shows that
full protection of albatrosses in the Chathams
is indeed essential, the Maoris will honor the
legislation to the letter; on the other hand, if
limited exploitation seems justified, the Wildlife
Service has agreed that it will be allowed.</p>


<h4>Conclusions</h4>

<p>Insofar as conservation measures of a passive
type are concerned, it is fortunate that
the offshore and outlying islands not yet occupied,
farmed, or set aside as reserves, are
likely to remain unexploited, either because
they are too remote for exploitation to be economical
or because they are too inhospitable,
or both. In any event, public opinion is now
such that unmodified or otherwise biologically
important islands not already reserved
would be proclaimed as reserves if threat of
exploitation arose unexpectedly, unless they
were found to be major sites for oil or
minerals. Even so, legislation exists that
offers the possibility of protection even from
this threat, and has already been used to
exempt some important mainland areas from
prospecting and the granting of mining
rights.</p>

<p>It is gratifying to realize that, although
some endemic marine subspecies (generally
not very different from neighboring subspecies)
are endangered to varying degrees,
there are very few whose disappearance would
result in the disappearance of the species itself
from the New Zealand area. Only one endemic
marine species has become extinct in recent
times, the Auckland Island merganser
<em>(Mergus australis)</em> in about 1905, and only six
are currently in any real danger: the Chatham
Island taiko <em>(Pterodroma magentae)</em>, the
black petrel, Hutton's shearwater <em>(Puffinus
huttoni)</em>, the Westland black petrel (<em>Procellaria
westlandica)</em>, the shore plover, and the
Chatham Island oystercatcher <em>(Haematopus
chathamensis)</em>. However, a list of this kind is
often a matter of some controversy. Something
is at present being done to help all but
the first and last of these. The Chatham Island
taiko had not been positively identified
for about 50 years, until 1977 when this
species was "rediscovered" on the main island
of the Chatham group; though its numerical
status is unknown, it is rare. The Chatham Island
oystercatcher, although certainly
"threatened" (only about 50 are known to
exist), does occur on four islands, two of which
are reserves. Although this species has not
been actively studied until now, it is soon to
be the subject of a full ecological survey.</p>

<p>A few words about the hunting of marine
species: Mutton-birding aside&mdash;that is, apart
from the taking by Maoris of the young of the
sooty shearwater and the gray-faced petrel&mdash;there
has been no legal hunting of any marine
birds in New Zealand for 35 years now, nor is
there likely to be. This situation reflects the
consistently increasing weight of informed
public opinion in favor of, let alone scientific
concern for, transoceanic migrants. The pro-hunting
lobby for some species of waders, in
particular the eastern bar-tailed godwit
<em>(Limosa lapponica baueri)</em>, is a small one, the
numbers of which decrease yearly. However,
small-scale poaching occasionally occurs; it is
punished when discovered.</p>

<p>Protection for marine species extends only
to the 3-mile limit of New Zealand's territorial
waters, but it would be extended further
should New Zealand follow the present trend
of including as territorial waters all those that
cover the continental shelf or beyond. [This
extension occurred in 1977; the marine fishing
zone for New Zealand waters has been extended
to 200 miles (360 km) around all
coasts.]</p>

<p>Only three marine species are not afforded
full protection under the Wildlife Act: two,
the black-backed or Dominican gull <em>(Larus
dominicanus)</em> and the black shag <em>(Phalacrocorax
carbo)</em>, are totally unprotected&mdash;the
first because of its predation on some rare
shorebirds during the breeding season and for
its attacks on sheep and lambs at a similar
time, and the second because of its depredations
(seldom serious) on the introduced trout
and salmon, mainly in fresh waters&mdash;the third<span class="pagenum"><a name="Page_266" id="Page_266">[Pg 266]</a></span>
species, the southern skua <em>(Stercorarius skua
lonnbergi)</em>, may be destroyed only when it is
actually attacking sheep or lambs, an occasional
event confined to the Chatham Islands.
Destruction of these three common species is
not encouraged by the Wildlife Service except
when black-backed gulls become too active
among colonies of, say, the fairy tern <em>(Sterna
nereis)</em>, which is very rare in New Zealand but
not elsewhere in its range. Otherwise, control
of the species is left in the hands of those most
affected by their depredations but whose
judgment is usually reasonable.</p>

<p>Marine birds, therefore, are generally satisfactorily
protected by law or managed for conservation
in New Zealand&mdash;especially when
one considers the remarkable changes that
have occurred in the New Zealand archipelago
over the last 200 years. Although the situation
could be better, it would certainly have
been worse if the Wildlife Service (and other
conservation organizations) had not been untiring
in keeping the general public and the
legislature aware of the issues at stake and
seen to it that as much as possible of the necessary
conservation work was done&mdash;and
done before it was too late.</p>


<h4>Acknowledgments</h4>

<p>I thank my Wildlife Service colleagues,
B. D. Bell, M. J. Imber, D. V. Merton, and
C. J. R. Robertson, for valuable comments
and advice on the preparation of this paper.</p>


<h4>References</h4>

<div class="hangindent">

<p>Atkinson, I. A. E., and B. D. Bell. 1973. Offshore
and outlying islands. Pages 372-392 <em>in</em> G. R.
Williams, ed. The natural history of New Zealand.
A. H. &amp; A. W. Reed, Wellington.</p>

<p>Bourne, W. R. P. 1967. Subfossil petrel bones from
the Chatham Islands. Ibis 109:1-7.</p>

<p>Kinsky, F. C. 1970. Annotated checklist of the birds
of New Zealand. A. H. &amp; A. W. Reed, Wellington.</p>

<p>Gibb, J. A., and J. E. C. Flux. 1973. Mammals.
Pages 334-371 <em>in</em> G. R. Williams, ed. The natural
history of New Zealand. A. H. &amp; A. W. Reed,
Wellington.</p>

<p>Williams, G. R. 1973. Birds. Pages 304-333 <em>in</em> G. R.
Williams, ed. The natural history of New Zealand.
A. H. &amp; A. W. Reed, Wellington.</p>

<p>Williams, G. R. 1977. Marooning&mdash;a technique for
saving threatened species from extinction. Int.
Zoo Yearb. 17:102-106.</p></div>

<p><span class="pagenum"><a name="Page_267" id="Page_267">[Pg 267]</a></span></p>
<div class="chapter"></div>
<hr class="chap" />




<h3><a name="Marine_Birds_in_the_Danish_Monarchy_and" id="Marine_Birds_in_the_Danish_Monarchy_and">Marine Birds in the Danish Monarchy and
Their Conservation</a></h3>


<p class="center">by</p>

<p class="center">Finn Salomonsen</p>

<p class="center">Chief Curator of Birds<br />
The Zoological Museum<br />
University of Copenhagen<br />
Copenhagen, Denmark
</p>


<h4>Abstract</h4>

<div class="blockquot">

<p>Most species of seabirds that regularly breed in Denmark are declining, for a
variety of reasons: shooting; oil pollution; toxic chemicals; reclamation of land;
collecting of eggs; disturbance at breeding sites by visitors, motorboats, camping,
etc.; destruction by predators; and others. On the other hand, the numbers of
certain other species are increasing as a result of climatic changes (six species),
protection (three species), and increase in food supply (three species of gulls). In
addition to breeding birds, a total of about 3 million birds occur in Danish waters
as passage migrants or winter visitors. More than half of the European winter
populations of a number of marine waterfowl species winter in Denmark. Large
numbers of seabirds spend the summer in Danish waters, including several hundred
thousand immature gulls and just as many molting waterfowl.</p>

<p>The seabird fauna of the Faroe Islands is very rich, the immense number of
birds being attracted by the local abundance of macroplankton and fish. The seabirds
are harvested by man, formerly by fowling (capturing and shooting), now
primarily by shooting. Until about 1910, more than 400,000 birds were taken
annually by fowling. The Faroese game act is now very restrictive, and most seabird
populations appear to be almost stable. However, a census in 1972 indicated
that common murres <em>(Uria aalge)</em> have declined by about 20% to a population of
about 600,000. Shooting and snaring appear to be the primary causes of the
decline; oil pollution and toxic chemicals do not seem to be contributing to the
population decrease.</p>

<p>In Greenland seabirds provide an important source of human food; however,
because of the increase in human population and in the use of guns and speedboats
for hunting, and the absence of a game act, serious overshooting of seabirds
is taking place. A new game act passed in 1977 should largely alleviate this
overharvest. Oil pollution and toxic chemicals do not yet play an important part
in influencing the number of seabirds, though offshore oil drilling is being
initiated in West Greenland. A recently established gigantic national park,
covering 200,000 km<sup>2</sup> of ice-free land, is the largest nature reserve in the world.</p></div>

<p><span class="pagenum"><a name="Page_268" id="Page_268">[Pg 268]</a></span></p>

<p>The Danish Monarchy consists of three
parts far removed from each other, scattered
in the North Atlantic&mdash;namely Denmark
proper, the Faroe Islands, and Greenland.
They differ so much from each other in climate
and in bird life that they must be treated
separately in this paper. The Faroes possess a
provincial government and also a sort of home
rule. Greenland also has a provincial government,
but all statutory provisions, including
acts concerning hunting or wildlife protection,
must be passed by Danish authorities, usually
by the Ministry of Greenland.</p>

<p>Insofar as seabirds are concerned, it is important
that Greenland is an arctic country,
whereas the Faroes and Denmark are boreal.
In both Greenland and the Faroes the breeding
birds are most significant, from an ecological
point of view, whereas in Denmark the
passage migrants and winter visitors are far
more important.</p>

<p>There are other differences as well. In
Greenland and the Faroes the seabirds mostly
breed in colonies on high and steep cliffs, and
the structure of these breeding places is not
disturbed by man. In Denmark, on the other
hand, the seabirds usually breed on glacial deposits,
now forming meadows, low islets, salt
marshes, etc., and these habitats have unfortunately
been largely changed in the last hundred
years by draining and reclamation. This
practice has taken place in Denmark on a
much larger scale than in most other countries
and has, therefore, to a high degree
diminished the life conditions of seabirds.</p>


<h4>Seabirds in Denmark</h4>

<p>Denmark is situated on the continental
shelf of western Europe; all seas surrounding
the country are shallow (less than 100 m
deep), apart from the Skagerrak, north of Jutland,
which is much deeper. The shallow
depth, combined with the rapid flow of water
between the Baltic and the North seas causes
much upwelling, which forms excellent life
conditions for plants and animals. It is well
known that the fishery in Danish waters, especially
in the North Sea, is very rich. This
richness of the seas provides suitable conditions
for a high diversity of seabirds and ecological
types.</p>

<p>Seabirds regularly breeding in Denmark include
five species of terns (common tern,
<em>Sterna hirundo</em>; arctic tern, <em>S. paradisaea</em>;
least tern, <em>S. albifrons</em>; Sandwich tern,
<em>S. sandvicensis</em>; and gull-billed tern, <em>Gelochelidon
nilota</em>); seven species of gulls (black-headed
gull, <em>Larus ridibundus</em>; herring gull,
<em>L. argentatus</em>; lesser black-backed gull, <em>L. fuscus</em>;
great black-backed gull, <em>L. marinus</em>; mew
gull, <em>L. canus</em>; little gull, <em>L. minutus</em>; and
black-legged kittiwake, <em>Rissa tridactyla</em>); four
species of geese, swans, and ducks (mute
swan, <em>Cygnus olor</em>; greylag goose, <em>Anser
anser</em>; common eider, <em>Somateria mollissima</em>;
common merganser, <em>Mergus merganser</em>; and
red-breasted merganser, <em>M. serrator</em>); three
species of auks (black guillemot, <em>Cepphus
grylle</em>; common murre, <em>Uria aalge</em>; and razorbill,
<em>Alca torda</em>); and one species of cormorant
(great cormorant, <em>Phalacrocorax carbo</em>).
Shorebirds have not been included in this review.
Some of the species mentioned are
partly freshwater birds&mdash;for example, the
black-headed gull, little gull, mute swan, greylag
goose, and the two species of mergansers.
The gull-billed tern forages in terrestrial habitats,
but nests along the coast with the other
seabirds. It is often difficult, therefore, to
make a clear-cut distinction between seabirds
and freshwater birds.</p>

<p>Among the auks, the black guillemot
breeds in the Cattegat area in the huge heaps
of boulders on small raised islets, or in holes
(mostly formed by starlings, <em>Sturnus vulgaris</em>)
on steep clayey slopes or promontories.
The common murre and razorbill are restricted
to the islet Graesholm in the Christians&oslash;
Archipelago, about 24 km east of Bornholm
Island in the Baltic, where they breed on
small cliffs of Precambrian granite rock.</p>

<p>The estimated number of seabirds of different
species that breed in Denmark is shown
in Table 1. Species like the mergansers, mute
swan, and greylag goose, which breed partly
or mostly in freshwater localities, are not included.
Overall, the number of breeding seabirds
is slowly declining, probably due to
many factors which are discussed below.
There are two exceptions, however, to this
general decrease&mdash;the herring gull (and to a
lesser degree the other big gull species) and
common eider. Both species have increased
during the last 50 years. Since they breed in
the same habitat, usually mixed together, the
eider is probably dependent on herring gulls
for protection against predators. When the
ducklings are fledged, the herring gull acts as
a successful predator itself, but the eider
nevertheless maintains a close association
with herring gulls.</p>

<p><span class="pagenum"><a name="Page_269" id="Page_269">[Pg 269]</a></span></p>

<div class="center">
<table border="0" cellpadding="4" cellspacing="0" summary="Table 1.
Estimated average number of breeding pairs of seabirds in Denmark,
based on a census in 1970-72.">
    <caption>Table 1. <em>Estimated average number of breeding pairs of
    seabirds in Denmark, based on a census in 1970-72.</em> (Data for
    terns from Mardal 1974, and for other species from Sten Asbirk
    and N. O. Preuss, personal communications.)</caption>
  <tr>
    <th>Species</th>
    <th>Number of breeding pairs</th>
  </tr>
  <tr>
    <td><em>Sterna paradisaea</em></td>
    <td class="tdr">5,750</td>
  </tr>
  <tr>
    <td><em>S. hirundo</em></td>
    <td class="tdr">900</td>
  </tr>
  <tr>
    <td><em>S. sandvicensis</em></td>
    <td class="tdr">4,000</td>
  </tr>
  <tr>
    <td><em>S. albifrons</em></td>
    <td class="tdr">600</td>
  </tr>
  <tr>
    <td><em>Gelochelidon nilotica</em></td>
    <td class="tdr">105</td>
  </tr>
  <tr>
    <td><em>Larus marinus</em></td>
    <td class="tdr">300</td>
  </tr>
  <tr>
    <td><em>L. argentatus</em></td>
    <td class="tdr">60,000</td>
  </tr>
  <tr>
    <td><em>L. fuscus</em></td>
    <td class="tdr">2,000</td>
  </tr>
  <tr>
    <td><em>L. canus</em></td>
    <td class="tdr">28,500</td>
  </tr>
  <tr>
    <td><em>L. ridibundus</em></td>
    <td class="tdr">135,000</td>
  </tr>
  <tr>
    <td><em>L. minutus</em></td>
    <td class="tdr">25</td>
  </tr>
  <tr>
    <td><em>Rissa tridactyla</em></td>
    <td class="tdr">125</td>
  </tr>
  <tr>
    <td><em>Phalacrocorax carbo</em></td>
    <td class="tdr">600</td>
  </tr>
  <tr>
    <td><em>Somateria mollissima</em></td>
    <td class="tdr">3,800</td>
  </tr>
  <tr>
    <td><em>Cepphus grylle</em></td>
    <td class="tdr">325</td>
  </tr>
  <tr>
    <td><em>Alca torda</em></td>
    <td class="tdr">400</td>
  </tr>
  <tr>
    <td><em>Uria aalge</em></td>
    <td class="tdr">1,100</td>
  </tr>
  <tr>
    <td class="tdp">Total</td>
    <td class="tdr">243,530</td>
  </tr>
</table></div>

<p>More than 90% of the herring gull population
breeds on small islands, and a large proportion
occurs in a few large colonies. It never
breeds in freshwater localities, but is exclusively
found as a breeding bird in coastal habitats.
The population has particularly increased
in the last 5 decades, some colonies
reaching their maximum size in the 1960's.
Others are still expanding and occupying new
breeding grounds. Today the largest colonies
are found on the following islands: Saltholm,
20,000-40,000 pairs; Christians&oslash; 9,000 pairs;
Hirsholmene, 2,500 pairs; Jordsand, 1,800
pairs; Sams&oslash;, 2,000 pairs; Hjelm, 1,500 pairs;
and the archipelago south of Funen, a total of
3,500 pairs in several colonies.</p>

<p>Attempts have been made to reduce the
breeding population of herring gulls at
Hirsholmene and Christians&oslash; sanctuaries (in
1973 and 1974, respectively), to improve conditions
for other nesting seabirds. In 1969 the
Bird Strike Committee of the Royal Danish
Airforce also initiated a program to reduce
the number of herring gulls breeding on
Saltholm Island, which is near the Kastrup
airport in Copenhagen. Nests were sprayed
with a formaldehyde oil dye, which resulted in
a 33% reduction in population. In Christians&oslash;
and Hirsholmene, where the adult breeding
birds were poisoned, the effect is not yet
known.</p>

<p>The total number of seabirds occurring in
the Danish waters as passage migrants and
winter visitors is substantially larger than the
breeding population, because Denmark is
situated on a very important fall migration
route for seabirds from Scandinavia, the Baltic
countries, northern Russia, and northwestern
Siberia. Furthermore, the shallow waters
of the Danish seas (less than 10 m deep) that
occupy extensive regions bordering the coasts
are important feeding grounds for diving
ducks. Birds frequenting the seas outside the
breeding season include hundreds of thousands,
or probably millions, of gulls;
numerous ducks (especially diving ducks);
swans and brants, <em>Branta bernicla</em>; jaegers,
<em>Stercorarius</em> spp. (four species); loons, <em>Gavia</em>
spp. (four species); grebes, <em>Podiceps</em> spp. (four
or five species); gannet, <em>Morus bassanus</em>;
great cormorant; northern fulmar, <em>Fulmarus
glacialis</em>; common murre; razorbill; and other
species of alcids. To these should be added a
number of species of various seabirds, especially
gulls, tubenoses, phalaropes, and others
which appear as casual or accidental visitors
and which are not further mentioned in this
paper.</p>

<p><span class="pagenum"><a name="Page_270" id="Page_270">[Pg 270]</a></span></p>

<p>Table 2. <em>Total numbers of ducks, swans, and coots recorded in Denmark during a winter census
in January 1973 (based on ground counts and aerial surveys), compared with estimated
flyway populations wintering in western Europe and annual bird harvest in Denmark (after
Joensen 1974:23, 155, 168).</em></p>

<div class="center">
<table border="0" cellpadding="4" cellspacing="0" summary="Table 2.
Total numbers of ducks, swans, and coots recorded in Denmark during a
winter census in January 1973">
    <caption>Table 2. <em>Total numbers of ducks, swans, and coots recorded
    in Denmark during a winter census in January 1973 (based on
    ground counts and aerial surveys), compared with estimated flyway
    populations wintering in western Europe and annual bird harvest
    in Denmark (after Joensen 1974:23, 155, 168).</em></caption>
  <tr>
    <th>Species</th>
    <th>Census, January 1973</th>
    <th>Estimated winter populations of the Western Europe Flyway</th>
    <th>Average annual bag in Denmark</th>
  </tr>
  <tr>
    <td><em>Anas platyrhynchos</em></td>
    <td class="tdn">127,000</td>
    <td class="tdn">1,550,000</td>
    <td class="tdn">380,000</td>
  </tr>
  <tr>
    <td><em>A. crecca</em></td>
    <td class="tdn">500</td>
    <td class="tdn">260,000</td>
    <td class="tdn">76,000</td>
  </tr>
  <tr>
    <td><em>A. querquedula</em></td>
    <td class="tdn">11</td>
    <td class="tdc"><a name="FNanchor_67_67" id="FNanchor_67_67"></a><a href="#Footnote_67_67" class="fnanchor">[67]</a></td>
    <td class="tdc"><a name="FNanchor_68_68" id="FNanchor_68_68"></a><a href="#Footnote_68_68" class="fnanchor">[68]</a></td>
  </tr>
  <tr>
    <td><em>A. acuta</em></td>
    <td class="tdn">100</td>
    <td class="tdn">70,000</td>
    <td class="tdn">13,000</td>
  </tr>
  <tr>
    <td><em>A. strepera</em></td>
    <td class="tdn">5</td>
    <td class="tdc"><a href="#Footnote_67_67" class="fnanchor">[67]</a></td>
    <td class="tdc"><a name="FNanchor_69_69" id="FNanchor_69_69"></a><a href="#Footnote_69_69" class="fnanchor">[69]</a></td>
  </tr>
  <tr>
    <td><em>A. penelope</em></td>
    <td class="tdn">3,000</td>
    <td class="tdn">485,000</td>
    <td class="tdn">44,000</td>
  </tr>
  <tr>
    <td><em>A. clypeata</em></td>
    <td class="tdn">17</td>
    <td class="tdn">63,000</td>
    <td class="tdn">9,000</td>
  </tr>
  <tr>
    <td><em>Tadorna tadorna</em></td>
    <td class="tdn">13,000</td>
    <td class="tdn">105,000</td>
    <td class="tdc"><a href="#Footnote_69_69" class="fnanchor">[69]</a></td>
  </tr>
  <tr>
    <td><em>Aythya ferina</em></td>
    <td class="tdn">7,100</td>
    <td class="tdn">235,000</td>
    <td class="tdn">5,000</td>
  </tr>
  <tr>
    <td><em>A. fuligula</em></td>
    <td class="tdn">94,700</td>
    <td class="tdn">530,000</td>
    <td class="tdn">35,000</td>
  </tr>
  <tr>
    <td><em>A. marila</em></td>
    <td class="tdn">80,900</td>
    <td class="tdn">145,000</td>
    <td class="tdn">8,000</td>
  </tr>
  <tr>
    <td><em>Clangula hyemalis</em></td>
    <td class="tdn">11,000</td>
    <td class="tdc"><a href="#Footnote_67_67" class="fnanchor">[67]</a></td>
    <td class="tdn">11,000</td>
  </tr>
  <tr>
    <td><em>Melanitta nigra</em></td>
    <td class="tdn">148,100</td>
    <td class="tdc"><a href="#Footnote_67_67" class="fnanchor">[67]</a></td>
    <td class="tdn">18,000</td>
  </tr>
  <tr>
    <td><em>M. fusca</em></td>
    <td class="tdn">6,700</td>
    <td class="tdc"><a href="#Footnote_67_67" class="fnanchor">[67]</a></td>
    <td class="tdn">9,000</td>
  </tr>
  <tr>
    <td><em>Somateria mollissima</em></td>
    <td class="tdn">450,800</td>
    <td class="tdc"><a href="#Footnote_67_67" class="fnanchor">[67]</a></td>
    <td class="tdn">136,000</td>
  </tr>
  <tr>
    <td><em>Bucephala clangula</em></td>
    <td class="tdn">67,000</td>
    <td class="tdn">142,000</td>
    <td class="tdn">25,000</td>
  </tr>
  <tr>
    <td><em>Mergus serrator</em></td>
    <td class="tdn">11,700</td>
    <td class="tdn">40,000</td>
    <td class="tdn">8,000</td>
  </tr>
  <tr>
    <td><em>M. merganser</em></td>
    <td class="tdn">23,200</td>
    <td class="tdn">75,000</td>
    <td class="tdn">6,000</td>
  </tr>
  <tr>
    <td><em>M. albellus</em></td>
    <td class="tdn">206</td>
    <td class="tdn">5,000</td>
    <td class="tdc"><a href="#Footnote_69_69" class="fnanchor">[69]</a></td>
  </tr>
  <tr>
    <td><em>Cygnus olor</em></td>
    <td class="tdn">48,900</td>
    <td class="tdn">120,000</td>
    <td class="tdc"><a href="#Footnote_69_69" class="fnanchor">[69]</a></td>
  </tr>
  <tr>
    <td><em>C. cygnus</em></td>
    <td class="tdn">5,700</td>
    <td class="tdn">17,000</td>
    <td class="tdc"><a href="#Footnote_69_69" class="fnanchor">[69]</a></td>
  </tr>
  <tr>
    <td><em>C. bewickii</em></td>
    <td class="tdn">1,113</td>
    <td class="tdn">6,000</td>
    <td class="tdc"><a href="#Footnote_69_69" class="fnanchor">[69]</a></td>
  </tr>
  <tr>
    <td><em>Fulica atra</em></td>
    <td class="tdn">142,500</td>
    <td class="tdc"><a href="#Footnote_67_67" class="fnanchor">[67]</a></td>
    <td class="tdn">70,000</td>
  </tr>
  <tr>
    <td class="tdp">Totals</td>
    <td class="tdn">1,243,252</td>
    <td class="tdn">3,848,000</td>
    <td class="tdn">853,000</td>
  </tr>
</table></div>

<p>A comprehensive investigation of the nonbreeding
waterfowl in Danish waters was recently
undertaken by the Game Biology Station
Kal&oslash; (Joensen 1974). Aerial surveys of
marine ducks indicate that a large percentage
of the ducks that winter in European waters
do so in the shallow areas of the Danish seas.
A census in January 1973 indicated a total of
more than 1.2 million birds (Table 2). In a
number of other countrywide surveys, undertaken
in all winters since 1967, usually 1.0-1.5
million birds have been recorded. Since such
censuses usually give minimum numbers, and
certain species-especially marine ducks&mdash;generally
go unrecorded, the normal winter
population (November to February) of ducks,
swans, and coots in Danish waters can
scarcely be less than 2 million birds (Joensen
1974:156). In Table 2, bird numbers in Denmark
are compared with the estimated winter
populations in western Europe, based on the
investigation of Atkinson-Willes (1972).
When all the winter censuses in Denmark are
compared with those for Europe, as was done
by Joensen (1974:156), it is evident that
Danish waters support about half of all
greater scaup <em>(Aythya marila)</em>, common
goldeneye <em>(Bucephala clangula)</em>, red-breasted
merganser, mute, whooper <em>(Cygnus cygnus)</em>,
and tundra swans <em>(C. bewickii)</em> wintering in
Europe; about one-third of the population of
tufted duck <em>(Aythya fuligula)</em> and common
merganser; and probably also one-third of the
population of common eider and coot <em>(Fulica
atra)</em>.</p>

<p>The wintering population of common eider
is very large. According to banding records it
makes up the greater part of Baltic breeding
birds; however, it is not possible to calculate
its percentage contribution to the total European
winter population since its size is unknown
in most European countries. Although
most of the surface-feeding ducks disappear
from Denmark waters in winter, extremely
large numbers occur there during the fall migration
period. For example, it has been estimated
that for species like common teal <em>(Anas
crecca)</em> and wigeon <em>(A. penelope)</em> about one<span class="pagenum"><a name="Page_271" id="Page_271">[Pg 271]</a></span>-third
of the West European Flyway population
passes Denmark in the fall. Possibly
some of the surface-feeding ducks listed in
Table 2 for January 1973 were recorded in
fresh water and not from the seas, but at the
time the census was taken most freshwater
lakes were frozen and, therefore, unavailable
for water birds.</p>

<p>These breeding seabirds and the off-season
visitors do not constitute the total population
in Danish waters. Large numbers also occur in
summer as nonbreeding birds; most are in two
categories: (1) several hundred thousand pre-adult
(up to 4-5 years of age) gulls (mostly
great black-backed, herring, and lesser black-backed
gulls), which feed inshore or at the
coast, and (2) large concentrations of waterfowl
that carry out a molt migration in
Danish waters, particularly in shallow areas.
Black scoter <em>(Melanitta nigra)</em>, velvet scoter
<em>(M. fusca)</em>, common eider, and whooper swan
are especially numerous, totaling hundreds of
thousands of individuals, and probably constituting
the majority of the European molting
populations of these species. Less
numerous, but still totaling thousands of
molting birds, are sheld-duck <em>(Tadorna
tadorna)</em>, common goldeneye, red-breasted
merganser, and possibly some other diving
ducks. About 3,000 surface-feeding ducks of
various species, most of which undoubtedly
are local breeding birds undergo wing molt in
Danish waters. Comprehensive descriptions
of the molt migration, particularly in Denmark,
were published by Salomonsen (1968)
and Joensen (1973<em>a</em>, 1974).</p>

<p>It may then be concluded that very large
numbers of seabirds are found in Danish
waters in all periods of the year; most feed in
the inshore zone and some offshore, but none
in the pelagic zone.</p>


<h5><em>Increase of Seabirds</em></h5>

<p>Seabirds are affected by several factors related
to human activities, most of which pose
a threat to them and will eventually reduce
their numbers. Some factors, however, tend to
increase bird numbers, like climatic changes
which, as reported by Salomonsen (1963), have
given rise to the immigration to Denmark of
great cormorant (in 1938); eared grebe, <em>Podiceps
nigricollis</em> (about 1870); red-crested
pochard, <em>Netta rufina</em> (1940); common
pochard, <em>Aythya ferina</em> (about 1860); tufted
duck (about 1900); and common murre (1929).
They all still breed in Denmark, having more
or less increased in number.</p>

<p>Another reason for increases of certain
species is legal protection. Among protected
seabirds are the sheld-duck, which has been
completely protected since 1931, and particularly
the mute swan, of which only 2 or 3 pairs
were breeding in Denmark when the species
was completely protected in 1926. Since then,
mute swans have increased enormously,
reaching at least 2,740 pairs in 1966 (Bloch
1971:43), of which large numbers were breeding
colonially on small islets of boulders or on
sand reefs off the coast (Bloch 1970:152). The
gannet has also increased considerably as a
fall visitor since about 1945, apparently due
to protection in England and other countries.</p>

<p>Finally, some gull populations have increased
in size because of an increase in the
food supply, consisting especially of wastes
from commercial fisheries and garbage
dumps. In Denmark, this unnatural food
source has caused an enormous increase since
about 1925 in herring gulls (from less than
500 pairs to 60,000 pairs), lesser black-backed
gulls (all three subspecies, <em>fuscus</em>, <em>intermedius</em>,
and <em>graelsii</em> have immigrated to Denmark),
and great black-backed gulls (immigrated
to Denmark in 1930). Improved waste
disposal practices in recent years have not yet
offset the rate of growth of these gull populations.
The increase of common eiders, which
also started in about 1925, is probably related
to the increases in the larger gulls.</p>


<h5><em>Decrease of Seabirds</em></h5>

<p>A variety of factors tend to reduce the numbers
of seabirds. The most important ones are
outlined below, with comments on what has
been done or what is expected to be done to reduce
the impact of these activities on seabirds
and protect this endangered resource.</p>


<h6>Shooting of Seabirds</h6>

<p>The shooting of seabirds in Denmark is considerable,
because the seabirds are extraordinarily
numerous, and the number of sports<span class="pagenum"><a name="Page_272" id="Page_272">[Pg 272]</a></span>men
is very large, amounting to about
135,000 (a larger number per capita than in
any other country).</p>

<p>The Danish game statistics are excellent&mdash;well
known to be much more accurate than in
most other countries (see Salomonsen 1954;
Strandgaard 1964). According to Danish bag
records, almost one million ducks, geese, and
coots (Joensen 1974:31) and about 100,000-200,000
gulls (Salomonsen 1954:125) are shot
each year. The average annual bag of each
species of wildfowl is given in Table 2 and the
open season for each species of seabirds in
Table 3. The open season for dabbling ducks
is long, extending from 16 August to 31 December,
which means that local birds are persecuted
almost as soon as birds-of-the-year
are able to fly. This has resulted in a dabbling
duck breeding population that is much
smaller than what the available food supply
could support, and in the large-scale development
of artificial rearing of mallards for later
shooting. A 5-month hunting season on
specialized birds like loons, grebes, and
various auks is not good management practice
and should be carefully reviewed.</p>

<p>Four other important facts about the shooting
of seabirds in Denmark merit inclusion
here: (1) there is no bag-limit for any species;
(2) in general, all marine areas within territorial
limits are open to all Danish sportsmen,
and the admission is free; (3) motorboats with
a maximum speed of 10 knots are allowed for
shooting in the period 1 October-30 April; and
(4) the shooting of seabirds is permissible
from 1.5 h before sunrise to 1.5 h (in December
1 h) after sunset, whereas for most other
birds shooting is prohibited between sunset
and sunrise.</p>

<p>Shooting is a national tradition in Denmark,
and the large number of sportsmen has
considerable political power. Too much influence
is given to the representatives of the
hunters' organizations, which have the decisive
force in game committees dealing with
protective measures. It is difficult, therefore,
to change the existing system.</p>

<ul>
<li class="caption">Table 3. <em>Open hunting seasons for seabirds
in Denmark, according to the Game Act
of 1967. Species not given in the table are
fully protected.</em></li>

<li class="ifrst">Hunting period and species</li>

<li class="indx">1 August-31 December</li>
<li class="isub1"><em>Anser anser</em></li>
<li class="isub1"><em>A. fabalis</em></li>
<li class="isub1"><em>A. brachyrhynchus</em></li>
<li class="isub1"><em>A. albifrons</em></li>
<li class="isub1"><em>Branta bernicla</em><a name="FNanchor_70_70" id="FNanchor_70_70"></a><a href="#Footnote_70_70" class="fnanchor">[70]</a></li>
<li class="isub1"><em>B. canadensis</em></li>

<li class="indx">1 August-30 April</li>
<li class="isub1"><em>Phalacrocorax carbo</em></li>

<li class="indx">16 August-31 December</li>
<li class="isub1"><em>Anas platyrhynchos</em></li>
<li class="isub1"><em>A. crecca</em></li>
<li class="isub1"><em>A. querquedula</em></li>
<li class="isub1"><em>A. acuta</em></li>
<li class="isub1"><em>A. penelope</em></li>
<li class="isub1"><em>A. clypeata</em></li>

<li class="indx">16 August-29 February</li>
<li class="isub1"><em>Aythya ferina</em></li>
<li class="isub1"><em>Fulica atra</em></li>
<li class="isub1"><em>Larus ridibundus</em></li>
<li class="isub1"><em>L. canus</em></li>

<li class="indx">16 August-30 April</li>
<li class="isub1"><em>L. fuscus</em></li>
<li class="isub1"><em>L. argentatus</em></li>
<li class="isub1"><em>L. marinus</em></li>

<li class="indx">1 October-29 February</li>
<li class="isub1"><em>Aythya fuligula</em></li>
<li class="isub1"><em>A. marila</em></li>
<li class="isub1"><em>Clangula hyemalis</em></li>
<li class="isub1"><em>Melanitta nigra</em></li>
<li class="isub1"><em>M. fusca</em></li>
<li class="isub1"><em>Somateria mollissima</em></li>
<li class="isub1"><em>Bucephala clangula</em></li>
<li class="isub1"><em>Mergus serrator</em></li>
<li class="isub1"><em>M. merganser</em></li>
<li class="isub1"><em>Gavia stellata</em></li>
<li class="isub1"><em>G. arctica</em></li>
<li class="isub1"><em>G. immer</em></li>
<li class="isub1"><em>Podiceps cristatus</em></li>
<li class="isub1"><em>Uria aalge</em></li>
<li class="isub1"><em>U. lomvia</em></li>
<li class="isub1"><em>Alca torda</em></li>
</ul>
<p><span class="pagenum"><a name="Page_273" id="Page_273">[Pg 273]</a></span></p>
<p>Shooting of seabirds, especially various
waterfowl, is popular and intensive. The number
of ducks taken by Danish sportsmen is
probably in the order of 10-15% of the total
kill on the West European Flyway (Joensen
1974:171). Excessive duck shooting can, in
some cases, be controlled by banding in the
breeding areas; the ensuing results then give
rise to strong protests from the Scandinavian
countries against the extensive persecution.
As stated above, Denmark has (in relation to
its size) the largest number of sportsmen of
any nation in the world and the most intensive
shooting. The number of sportsmen
shooting ducks and shorebirds per 100 km<sup>2</sup> is
278 in Denmark, 28 in Sweden, 37 in Finland,
10 in Poland, 83 in Holland, 164 in Britain,
and 129 in Western Germany; the number of
ducks shot per 100 km<sup>2</sup> is 1,856 in Denmark,
39 in Sweden, 68 in Finland, and 129 in Western
Germany (Nowak 1973). This shooting is
undoubtedly of importance to dabbling duck
populations, which are popular as shooting
objects everywhere in Europe.</p>

<p>Insofar as marine ducks are concerned, it
can be seen in Table 2 that appreciable numbers
are shot in Denmark. The same is true for
other Scandinavian countries, whereas shooting
on the high seas is rather modest in most
other European countries. The Danish bag undoubtedly
makes up a significant proportion
of the total number of marine ducks killed
each year, but when the total number of ducks
in European waters is considered, the shooting
pressure in Denmark appears to be of only
minor importance. However, the shooting,
particularly when undertaken from motorboats,
is so noisy and makes such a disturbance
over large areas that the time for
seabirds to rest and forage is significantly reduced.
It must also be noted that the number
of pleasure craft is steadily increasing in the
present period of prosperity, and that increasing
numbers of sportsmen will probably make
use of the free shooting in territorial waters,
since it is becoming more and more expensive
to lease hunting areas.</p>

<p>To restrict seabird shooting, the Danish
Ornithological Society has recently (1975)
submitted a proposal to the Danish Government,
of which the following points are
relevant:</p>

<p>&#8226; The open season for dabbling ducks and
geese should begin 15 September except for
pintail <em>(Anas strepera)</em>, shoveler <em>(A. clypeata)</em>,
wigeon, and pochard&mdash;species which should
not be hunted until 1 October;</p>

<p>&#8226; the open season for all diving ducks, as
well as for coot, should end 31 December;</p>

<p>&#8226; the open season for the great cormorant
should be restricted to the period between 15
September and 31 October;</p>

<p>&#8226; murres, razorbill, great-crested grebe
<em>(Podiceps cristatus)</em>, and all species of loon
should be fully protected;</p>

<p>&#8226; it should be prohibited to shoot from
motorboats less than 1 km from the shoreline,
as well as in certain narrow sounds and fjords;</p>

<p>&#8226; it should be prohibited to shoot from
shooting-punts less than 100 m from the
shoreline;</p>

<p>&#8226; it should be prohibited to sell waterfowl
and shorebirds shot, except for eider ducks
and mallards <em>(Anas platyrhynchos)</em>; and</p>

<p>&#8226; no shooting should be allowed between
sunset and one hour before sunrise.</p>

<p>Oil Pollution</p>

<p>Oil pollution incidents constitute one of the
greatest dangers to seabird populations in
Danish waters. The enormous masses of seabirds
present in these waters throughout the
year, combined with the fact that Danish
waters contain some of the heaviest shipping
traffic in the world would give rise to anxiety
for oil disasters. The majority of all tanker
traffic from the Atlantic and the North Sea to
the Baltic passes through the Cattegat and
the narrow straits of the Sound, the Great
Belt, and the Little Belt, to supply a population
of about 100 million people. Up to
100,000 ships pass through these waters each
year, half through the Sound.</p>

<p>There have been severe oil pollution disasters
every year since about 1935, accompanied
by enormous mortalities of seabirds,
particularly marine ducks. The Danish Game
Biology Station, which has studied these
disasters (Joensen 1972<em>a</em>, 1972<em>b</em>, 1973<em>b</em>), has
noticed that the number of seabirds involved
has increased in recent years, in spite of increased
control by Danish authorities.</p>

<p>Unfortunately, it appears that small
amounts of oil in the sea, originating from
cleaning the tanks of vessels, or from the release
of a few tons of oil, are enough to create
mass mortality of seabirds when large concentrations
of birds are present in the vicinity.
Such incidents have passed unnoticed in spite
of control measures. In no case has the source
of the pollution been traced (Joensen
1972<em>b</em>:27). There has not yet been a real "oil
disaster" in the Danish waters similar to the
<em>Torrey Canyon</em> catastrophe. If such a disaster
takes place, the destruction of seabirds will be
enormous and immeasurable.</p>

<p><span class="pagenum"><a name="Page_274" id="Page_274">[Pg 274]</a></span></p>

<div class="center">
<table border="0" cellpadding="4" cellspacing="0" summary="Table 4.
Species composition of 8,304 birds killed by oil and examined in
connection with five pollution disasters in the Cattegat, 1969-71.">
    <caption>Table 4. <em>Species composition of 8,304 birds killed by oil
    and examined in connection with five pollution disasters in the
    Cattegat, 1969-71.</em> (After Joensen 1972:12.)</caption>
  <tr>
    <th rowspan="2">Species</th>
    <th colspan="0">Oil incident no.</th>
  </tr>
  <tr>
    <th>1</th>
    <th>2</th>
    <th>3</th>
    <th>4</th>
    <th>5</th>
    <th>Totals</th>
  </tr>
  <tr>
    <td><em>Gavia stellata</em></td>
    <td class="tdn">1</td>
    <td></td>
    <td class="tdn">9</td>
    <td class="tdn">1</td>
    <td class="tdn">4</td>
    <td class="tdn">15</td>
  </tr>
  <tr>
    <td><em>G. arctica</em></td>
    <td class="tdn">2</td>
    <td></td>
    <td class="tdn">2</td>
    <td class="tdn">4</td>
    <td class="tdn">8</td>
    <td class="tdn">16</td>
  </tr>
  <tr>
    <td><em>Gavia sp.</em></td>
    <td class="tdn">4</td>
    <td></td>
    <td class="tdn">1</td>
    <td></td>
    <td></td>
    <td class="tdn">5</td>
  </tr>
  <tr>
    <td><em>Podiceps grisegena</em></td>
    <td class="tdn">4</td>
    <td></td>
    <td class="tdn">1</td>
    <td class="tdn">8</td>
    <td class="tdn">8</td>
    <td class="tdn">21</td>
  </tr>
  <tr>
    <td><em>P. cristatus</em></td>
    <td></td>
    <td></td>
    <td class="tdn">1</td>
    <td></td>
    <td></td>
    <td class="tdn">1</td>
  </tr>
  <tr>
    <td><em>Phalacrocorax carbo</em></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdn">20</td>
    <td></td>
    <td class="tdn">20</td>
  </tr>
  <tr>
    <td><em>Anas platyrhynchos</em></td>
    <td class="tdn">2</td>
    <td></td>
    <td class="tdn">2</td>
    <td></td>
    <td></td>
    <td class="tdn">4</td>
  </tr>
  <tr>
    <td><em>A. clypeata</em></td>
    <td class="tdn">2</td>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdn">2</td>
  </tr>
  <tr>
    <td><em>Aythya marila</em></td>
    <td></td>
    <td></td>
    <td class="tdn">6</td>
    <td class="tdn">2</td>
    <td></td>
    <td class="tdn">8</td>
  </tr>
  <tr>
    <td><em>Clangula hyemalis</em></td>
    <td class="tdn">35</td>
    <td class="tdn">2</td>
    <td class="tdn">26</td>
    <td class="tdn">6</td>
    <td class="tdn">4</td>
    <td class="tdn">73</td>
  </tr>
  <tr>
    <td><em>Melanitta nigra</em></td>
    <td class="tdn">387</td>
    <td class="tdn">241</td>
    <td class="tdn">521</td>
    <td class="tdn">262</td>
    <td class="tdn">77</td>
    <td class="tdn">1,488</td>
  </tr>
  <tr>
    <td><em>M. fusca</em></td>
    <td class="tdn">197</td>
    <td class="tdn">33</td>
    <td class="tdn">417</td>
    <td class="tdn">223</td>
    <td class="tdn">119</td>
    <td class="tdn">989</td>
  </tr>
  <tr>
    <td><em>Somateria mollissima</em></td>
    <td class="tdn">1,683</td>
    <td class="tdn">1,081</td>
    <td class="tdn">947</td>
    <td class="tdn">1,713</td>
    <td class="tdn">19</td>
    <td class="tdn">5,443</td>
  </tr>
  <tr>
    <td><em>Bucephala clangula</em></td>
    <td class="tdn">3</td>
    <td class="tdn">3</td>
    <td class="tdn">13</td>
    <td class="tdn">9</td>
    <td></td>
    <td class="tdn">28</td>
  </tr>
  <tr>
    <td><em>Mergus serrator</em></td>
    <td class="tdn">48</td>
    <td></td>
    <td class="tdn">28</td>
    <td class="tdn">28</td>
    <td class="tdn">2</td>
    <td class="tdn">106</td>
  </tr>
  <tr>
    <td><em>Cygnus olor</em></td>
    <td class="tdn">10</td>
    <td></td>
    <td class="tdn">17</td>
    <td class="tdn">1</td>
    <td></td>
    <td class="tdn">28</td>
  </tr>
  <tr>
    <td><em>C. Cygnus</em></td>
    <td></td>
    <td></td>
    <td class="tdn">1</td>
    <td></td>
    <td></td>
    <td class="tdn">1</td>
  </tr>
  <tr>
    <td><em>Fulica atra</em></td>
    <td class="tdn">1</td>
    <td class="tdn">1</td>
    <td class="tdn">2</td>
    <td class="tdn">5</td>
    <td></td>
    <td class="tdn">9</td>
  </tr>
  <tr>
    <td><em>Larus sp.</em></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdn">13</td>
    <td></td>
    <td class="tdn">13</td>
  </tr>
  <tr>
    <td><em>Alca torda</em></td>
    <td></td>
    <td class="tdn">1</td>
    <td></td>
    <td class="tdn">12</td>
    <td class="tdn">1</td>
    <td class="tdn">14</td>
  </tr>
  <tr>
    <td><em>Uria aalge</em></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdn">1</td>
    <td></td>
    <td class="tdn">1</td>
  </tr>
  <tr>
    <td><em>Cepphus grylle</em></td>
    <td class="tdn">1</td>
    <td></td>
    <td class="tdn">2</td>
    <td class="tdn">16</td>
    <td></td>
    <td class="tdn">19</td>
  </tr>
  <tr>
    <td class="tdp">Total birds examined</td>
    <td class="tdn">2,380</td>
    <td class="tdn">1,362</td>
    <td class="tdn">1,996</td>
    <td class="tdn">2,324</td>
    <td class="tdn">242</td>
    <td class="tdn">8,304</td>
  </tr>
  <tr>
    <td class="tdp">Estimated minimum number of birds killed</td>
    <td class="tdn">10,000</td>
    <td class="tdn">5,000</td>
    <td class="tdn">12,000</td>
    <td class="tdn">15,000</td>
    <td class="tdn">1,500</td>
    <td class="tdn">43,500</td>
  </tr>
  <tr>
    <td class="tdp">Percent of total birds contributed by three species<a name="FNanchor_71_71" id="FNanchor_71_71"></a><a href="#Footnote_71_71" class="fnanchor">[71]</a></td>
    <td class="tdn">95.3</td>
    <td class="tdn">99.5</td>
    <td class="tdn">94.4</td>
    <td class="tdn">94.6</td>
    <td class="tdn">88.8</td>
    <td class="tdn">95.4</td>
  </tr>
</table></div>

<p>As a result of five of the major oil pollution
incidents in the Cattegat from 1969-71, a total
of 43,500 birds were killed, of which 8,304
were examined and enumerated (Table 4). Altogether,
21 or 22 species were involved, but
95% of all birds examined were diving ducks:
common eider and black and velvet scoters.
At present, it has not been possible to identify
any decrease in the number of these ducks in
Danish waters due to oil pollution. However,
if these disasters continue, it can be expected
that duck populations of northern Europe and
the Baltic area will be severely reduced, and
that an overall decline will take place from
which the birds may not be able to recover.</p>

<p>A particularly disastrous year was 1972,
when large numbers of ducks were killed as a
result of rather small oil spills. A tanker disaster
in March 1972 off the eastern coast of Jutland,
in the northern Cattegat, and another in
December 1972 in the Danish Waddensea,
both took place in areas critical to major concentrations
of sea ducks. A total of more than
60,000 birds were killed, of which about 95%
consisted of the same three species of diving
ducks mentioned above. These tragic events
represent a further increase in the annual mortality
of birds caused by oil, and there is reason
to believe that a critical upper limit is
rapidly being approached.</p>

<p>It appears, however, that the measures
taken by pollution control and naval authorities
have greatly improved in recent years. In
January 1973, when a Polish merchant vessel
collided with a Swedish tanker in the Sound,
about 300 tons of heavy fuel oil were released
into the sea. Several Danish and Swedish
ships working in cooperation succeeded in dis<span class="pagenum"><a name="Page_275" id="Page_275">[Pg 275]</a></span>persing
the oil, and no serious effect on seabird
populations took place (Joensen
1973<em>b</em>:118). It seems that the best way of
cleaning up such oil disasters is through a mechanical
removal of the oil, but this is a very
expensive and difficult procedure.</p>


<h6>Pollution by Toxic Chemicals</h6>

<p>Chemical pollution is probably the most
ominous threat to seabirds at present. Since
all toxic chemicals used in agriculture ultimately
end up in the sea, and many large factories
release their industrial wastes directly
into the sea, the effects of this pollution on
marine organisms is attracting a growing interest.
Many students have worked on these
problems, and the results that concern birds
were summarized by Bourne (1972:205). It is
known that organochlorine residues have
been found in seabirds in all the oceans of the
world, including Antarctic waters and Arctic
seas (Bogan and Bourne 1972:358). The
chemicals most often found in birds are DDE
(a metabolite of DDT) and PCB's (polychlorinated
biphenyls), a mixture of related chemical
compounds often originating from industrial
wastes. In addition, some mercury will
always be found, sometimes in increased concentrations.
The present restrictions on the
use of DDT and PCB in Denmark have not yet
resulted in a corresponding decrease in the
amount of these pesticides in birds.</p>

<p>It is well known that marine pollution
reaches a peak in the Baltic. This high level of
pollution is reflected in seabirds. For example,
analyses have shown that eggs from the
colony of common murres on Christians&oslash; in
the Baltic contain about 100 times as much
DDE and 50 times as much PCB as eggs of
murres from the Faroe Islands in the Atlantic
Ocean (Dyck 1975).</p>

<p>A similar difference exists in the mercury
content in birds examined in the two areas.
Feathers of a large sample of black guillemots
and murres from the Cattegat and the Baltic
had higher mercury levels than those from the
Faroe Islands and Greenland. It is interesting
that this difference existed over a hundred
years ago, as evidenced by the analysis of
feathers in museum specimens. The Baltic
populations of both species show very significant
increases in the mercury content in 1965-70,
as compared with the values earlier in this
century. Since 1970 there has been a sharp decrease
in mercury content, and in 1973 the
level was almost as low as it was early in the
century. These results indicate that the strict
control of mercury discharges enforced in
Sweden has resulted in a quick recovery of
nearly normal conditions in the Baltic (Somer
and Appelquist 1974). However, recent
studies by Koeman et al. (1975:286) appear to
show that mercury does not accumulate to the
same extent in seabirds as it does in seals.</p>

<p>High concentrations of chlorinated hydrocarbon
residues accumulate in carnivorous
birds and upset the normal breeding behavior
by making the eggshells too thin and fragile
to survive (Peakall 1970:73; Mueller and
Leach 1974:289). In Denmark, shells of
herring gull eggs from the Baltic population
were thinner, lighter, and more heavily contaminated
with DDE and PCB than were
shells of eggs from other colonies (J&oslash;rgensen
and Kraul 1974:173). This further emphasizes
the pollution of the Baltic Sea.</p>

<p>Massive mortalities of common murres,
such as the one reported in the Irish Sea in the
fall of 1969 which was apparently caused
partly by malnutrition and PCB poisoning
(Parslow and Jefferies 1973:87), are unknown
in Danish waters.</p>

<p>It should be added that the pollution of seawater
with cadmium, so very dangerous for
man, has been high in recent years owing to
the increased use of this element in industry,
but no analysis of its importance for seabirds
in Danish waters has yet been made.</p>

<p>It should also be mentioned that pollution
of fresh water in lagoons or lakes near the sea
can often cause serious declines in numbers of
certain seabirds. This is well illustrated by recent
events in the sanctuary Nakskov Indrefjord
on the island of Lolland. This landlocked
fjord once supported numerous breeding
populations of ducks, grebes, and terns, but in
recent years a number of species (e.g., eared
grebe; common teal; garganey, <em>Anas querquedula</em>;
pintail; and black tern, <em>Chlidonias nigra</em>)
have failed to breed and practically all other
species have declined in numbers. The main
reason for these changes is a severe pollution
from the admission of raw sewage from tributaries
(Bloch et al. 1972). After several out<span class="pagenum"><a name="Page_276" id="Page_276">[Pg 276]</a></span>breaks
of botulism in recent years, procedures
to improve conditions are now being
developed.</p>


<h6>Other Threats to Seabirds</h6>

<p>The most dangerous threats to seabirds are
those discussed above. Authorities are aware
of these dangers and attempts are being made
to improve conditions. Some results have
been achieved in the combat against oil pollution,
and the control of shooting is reaching
an acceptable level. Game management agencies
in Denmark and other Scandinavian
countries (Norway, Sweden, and Finland) are
cooperating on the request of the parliamentary
body of the Nordic Council. If game
biologists in these countries could agree on
proposed changes in the game acts, owing to
the marked decline of a number of bird
species, the parliamentary basis for such a
legal step would be absolutely certain.</p>

<p>However, it must be admitted that the impact
of man on the environment is enormous,
especially in a country like Denmark, which
possesses no raw materials, and where agriculture
has transformed the whole country. In
such a country, the birds have to "face the
music," and by this sharing of resources with
man, they will inevitably decrease in number.
It is the responsibility of biologists and politicians,
without emotional biases, to find the
balance between the requirements of the two
spheres of interest.</p>

<p>Many other dangers that threaten seabirds,
some of which are unrelated to human activities,
are listed here.</p>

<p>&#8226; Land reclamation.&mdash;Reclamation of land
has reduced extensive areas of shallow water,
lagoons, marsh land, etc., from seabirds for
foraging or breeding places. Draining and diking
of coastlands, estuaries, and saltings have
had the same effect. This activity is now almost
stopped, as these projects are no longer
subsidized by the government.</p>

<p>&#8226; Egg-collecting.&mdash;According to the present
game act, collecting gull eggs is permitted
until 24 May. This creates much disturbance
on the breeding grounds, and eggs of terns
and shorebirds are also taken. This practice
should be halted. The "Bird Island Group" of
the Danish Ornithological Society, in a symposium
in 1972, prepared some rules for the
protection of seabirds, among which is a proposal
to stop egg-collecting.</p>

<p>&#8226; Common property.&mdash;The Nature Conservancy
Act regards all land not fenced in, even
small uninhabited islets, as common property.
People have free access to such areas with the
result that seabirds breeding in colonies, or
separately on islands, are disturbed by visitors
arriving by boat. At the same time, noisy
motorboats, bathing parties, or camping visitors
frighten the birds, making successful
breeding almost impossible. Even ornithologists,
bird-banding teams, and bird photographers
add to the destruction. The "Bird Island
Group" of the Danish Ornithological
Society has proposed a general prohibition
against visitors on important bird islands
from 1 March to 15 July to protect the breeding
seabirds.</p>

<p>&#8226; Destruction by predators.&mdash;Fox, ermine,
and stone-marten do not play an essential
role. Rats are more important, even on small
islands, and have caused destruction of tern
and gull colonies. Rat numbers do not decline
until a severe winter with much ice occurs, or
until high tide kills them all. Large gulls also
cause a great deal of destruction, but crows
and magpies are unimportant as predators in
seabird colonies. Numbers of nonbreeding
mute swans or greylag geese may sometimes
be a nuisance, trampling eggs and nestlings in
seabird colonies.</p>

<p>&#8226; Forestry practices.&mdash;The prevailing practice
of the forestry industry in Denmark of
not preserving old trees with holes has considerably
diminished the breeding habitat of
hole-nesting species like the common merganser.
Artificial nest-boxes have now been
established in several areas.</p>

<p>&#8226; Sea conditions.&mdash;During high water, or
rough sea, salt water may flood colonies of
breeding seabirds nesting on low islets, often
reducing the production of young.</p>

<p>&#8226; Aircraft disturbance.&mdash;Disturbances are
also caused by noise from jet aircraft flying
low, especially in military training areas
where air traffic may be heavy.</p>

<p>&#8226; Commercial fisheries.&mdash;Modern commercial
fisheries are depleting so-called industrially
important fish stocks such as sand eels
<em>(Ammodytes)</em>, herrings, and other small fish
over large areas of the sea for the production<span class="pagenum"><a name="Page_277" id="Page_277">[Pg 277]</a></span>
of fish meal. This fishing has undoubtedly
been the main reason for the decline in the
number of terns&mdash;especially sandwich terns
which depend on these small fish species for
food.</p>

<p>&#8226; Unknown factors at sea.&mdash;Large numbers
of pelagic seabirds, particularly fulmars, kittiwakes,
and gannets, are washed up on the
western coast of Jutland in certain years (e.g.,
1959, Joensen 1961:212). These birds died at
sea, for unknown reasons, and apparently as a
result of food shortages or oil pollution.</p>


<h5><em>Conservation</em></h5>

<p>The threats to seabirds mentioned above
are all well known to conservationists, who
are attempting to reduce the impact of these
factors on seabirds where possible. Insofar as
legal protection is concerned, it must be admitted
that there are no marine sanctuaries in
Denmark, although several discussions have
taken place reviewing the possibility of establishing
some in critical areas. There are, however,
a number of sanctuaries on islands
where seabirds breed. In the Sanctuary Act of
1936 these areas were called "Scientific Reserves"
because they were the site of scientific
investigations of bird life. All admission
was forbidden, at least during the breeding
season, and all shooting was prohibited, with
few exceptions. These sanctuaries were administered
by the government's Nature
Conservancy.</p>

<p>The following Scientific Reserves are important
for seabirds: Hirsholmene Islands (in
Cattegat off Frederikshavn), Knotterne Islands
(small islets east of Laes&oslash; Island),
Vejlerne (diked in, landlocked fjords, densely
covered with vegetation, at the Lim Fjord),
Tipperne Peninsula and Klaegbanken Island
(in Ringk&oslash;bing Fjord, western Jutland),
Vars&oslash; Island (Horsens Fjord, eastern Jutland),
and Gr&aelig;holm Island (Christians&oslash;
Archipelago, in the Baltic off Bornholm). A
detailed description of these sites and their
erection, bird life, and ornithological value
was given by Salomonsen (1945). More recently,
two additional Scientific Reserves
have been established: Aegholm Islet (south
of Sealand), and Hessel&oslash; Island in the southern
part of Cattegat.</p>

<p>In addition to these scientific sanctuaries,
there are game reserves and governmental
forest reserves in Denmark. The game reserves
are administered by the Ministry of
Agriculture, which is also responsible for
hunting legislation. The purpose of game reserves
is to support and protect the stock of
game, which includes migrating birds. Shooting
is usually prohibited, but a restricted
shooting season is allowed at some reserves.
More than 50 game reserves are now present
and functioning. Regulations differ widely
from reserve to reserve, but entry to some of
them is not allowed in the breeding season.
Many reserves are important for breeding or
migrating waterfowl and some seabirds. In
fact, a total of 26 game reserves contain seabirds,
the most important of which are the following:
Ulvedybet (landlocked fjord at the
Lim Fjord), Hjarbaek Fjord (landlocked fjord
with brackish water at the Lim Fjord), Felsted
Kog (landlocked fjord at Nissum Fjord),
Jordsand (large stretches, almost 11,000 ha,
of the Danish Waddensea), Stavns Fjord (at
Sams&oslash; Island), Esrum Lake (in northern Sealand),
and Kalvebod Beach (at Amager Island,
near Copenhagen).</p>

<p>In the Nature Conservancy Act of 1969, differences
between scientific and game reserves
were abolished, although regulatory provisions
that were in force for the scientific sanctuaries
were maintained. Unfortunately, the
amalgamation of the two types of reserve has
given more power to the hunters' associations,
which constitute the majority of the administrative
body of the reserves, the so-called
Game Commission ("Vildtn&aelig;vnet").
However, any change in status of the original
scientific reserves will not be tolerated by conservationists
and other environmental groups
in Denmark.</p>


<h4>The Faroe Islands</h4>

<p>The number of seabirds in the Faroe Islands
is greater than in any other region of the
North Atlantic, and is closely related to the
extraordinary richness of the plankton. The
high phytoplankton production is due to a
strong vertical mixing of the water in the
northeast Atlantic, especially at the slopes of
the submarine ridges, where both tidal currents
and oceanic currents are usually strong.
The resulting upwelling enriches the upper
layers of water with large quantities of nutrient
salts for the phytoplankton, and this, in
turn, produces a teeming life of macroplankton
and fish on which the seabirds are dependent
(Salomonsen 1955).</p>

<p><span class="pagenum"><a name="Page_278" id="Page_278">[Pg 278]</a></span></p>

<p>The enormous seabird population of the
Faroes is apparent from the first description
of the islands, "De mensura orbis terrae," a
document written in the year 825 by the Irish
monk Dicuilus, who described the most characteristic
feature of the Faroes as being the
fact that "the islands were full of various
kinds of marine birds." This richness has remained
to the present, and has provided an
important source of food for the resident human
population, particularly in former times.
There are few, if any, countries in the world in
which wild-fowling and other exploitations of
birdlife have played such a major role as in the
Faroes. A number of elaborate and varied
bird-catching methods were invented, and
these have remained essentially the same for
at least the last 500 years. Bird-fowling at
great heights on precipitous sea-cliffs was a
dangerous venture, and each year lives were
lost. The main thing, however, was that food
obtained from fowling meant life and death
for local inhabitants and so was undertaken in
such a well-balanced way that the seabird
populations did not decrease or disappear.
Some fowling still takes place, but on a reduced
scale, since most men are now engaged
in the fishery during the summer. Shooting is
now of much greater importance than in former
times.</p>

<p>The Faroese game acts (from 1897, 1928,
and 1954) are very severe and show a broad
consideration for birdlife. Practically all terrestrial
birds, including shorebirds, are protected,
and existing regulations permit people
to catch or shoot only common murres, razorbills,
puffins, shags <em>(Phalacrocorax aristotelis)</em>,
fulmars, gannets, parasitic jaegers
<em>(Stercorarius parasiticus)</em>, and gulls, as well as
a few "pest" species like crows <em>(Corvus
corone)</em> and ravens <em>(C. corax)</em>. The legal right
of fowling on a "fowling cliff" belongs to the
registered owner of the land on which the cliff
is situated. There are some sound restrictive
laws for these cliffs. For example, shooting
within 3.2 km of any seabird colony is
prohibited.</p>

<div class="center">
<table border="0" cellpadding="4" cellspacing="0" summary="Table 5.
Number of seabirds caught by fowling each year in the Faroe Islands in
the early 1900's.">
    <caption>Table 5. <em>Number of seabirds caught by fowling each
    year in the Faroe Islands in the early 1900's.</em> (From
    Salomonsen 1935.)</caption>
  <tr>
    <th>Species</th>
    <th>Number of birds caught per year</th>
  </tr>
  <tr>
    <td><em>Uria aalge</em></td>
    <td class="tdr">60,000</td>
  </tr>
  <tr>
    <td><em>Fratercula arctica</em></td>
    <td class="tdr">270,000</td>
  </tr>
  <tr>
    <td><em>Puffinus puffinus</em></td>
    <td class="tdr">1,500</td>
  </tr>
  <tr>
    <td><em>Fulmarus glacialis</em></td>
    <td class="tdr">80,000</td>
  </tr>
  <tr>
    <td><em>Morus bassanus</em></td>
    <td class="tdr">1,300</td>
  </tr>
  <tr>
    <td class="tdp">Total</td>
    <td class="tdr">412,800</td>
  </tr>
</table></div>

<p>The annual number of seabirds caught by
fowling in the early 1900's (summarized in
Table 5) were reported in Salomonsen (1935).
This large harvest of birds, taken by fowling
year after year for centuries, did not appear to
influence the seabird populations, as bird
numbers remained stable. However, in recent
years, shooting and a special form of snaring
of murres have increased dramatically and
seem to have endangered the murre population.
The annual number of murres killed is estimated
to be about 120,000, of which 70,000
are snared and at least 50,000 shot (estimates
of birds shot range from 50,000 to 100,000).
This total is almost double the number of
birds caught during fowling, and because of
an apparent decline in murre numbers the provincial
government decided to investigate the
matter, and in 1972 the Danish Ornithological
Society agreed to conduct the study. Figures
from the 1972 census of murres (Table 6) show
that almost 600,000 birds were counted, from
which an estimate of more than 393,000
breeding pairs was calculated (Dyck and Meltofte
1975). In spite of this large number,
Dyck and Meltofte (1975) concluded that the
Faroese murre population has declined by
about 20% during the last 10-15 years. Investigations
are under way to monitor further
changes in murre numbers, and to determine
the trend, and whether reductions in shooting
and snaring are necessary to maintain the
population.</p>

<p>Oil pollution is practically unknown in
Faroese waters, but since drilling for oil will
probably take place in the near future, the importance
of oil to birds in this region may
change. Toxic chemicals do not appear to be<span class="pagenum"><a name="Page_279" id="Page_279">[Pg 279]</a></span>
involved in the decline in murres. Investigations
of concentrations of chemical pollutants
in their eggs show that levels of DDE (mean
1.1 ppm), PCB (mean 2.0 ppm), and mercury
(mean 0.2 ppm) (Dyck and Meltofte 1975) are
relatively low and unlikely to affect reproduction
(Dyck and Meltofte 1975). Levels are
much smaller than those found in seabirds in
Britain, the Baltic, or in albatrosses in the
Pacific (Fisher 1973).</p>

<div class="center">
<table border="0" cellpadding="4" cellspacing="0" summary="Table 6.
Colonies of the common murre, Uria aalge, on the Faroe Islands, based
on a census conducted in 1972.">
    <caption>Table 6. <em>Colonies of the common murre,</em> Uria aalge,
    <em>on the Faroe Islands, based on a census conducted in
    1972.</em> (After Dyck and Meltofte 1975.)</caption>
  <tr>
    <th>Colony</th>
    <th>Number of birds observed</th>
    <th> Number of pairs<a name="FNanchor_72_72" id="FNanchor_72_72"></a><a href="#Footnote_72_72" class="fnanchor">[72]</a></th>
  </tr>
  <tr>
    <td>Suderoy</td>
    <td class="tdr"> 73,945</td>
    <td class="tdr"> 49,500</td>
  </tr>
  <tr>
    <td>L&iacute;tla D&iacute;mun</td>
    <td class="tdr"> 13,220</td>
    <td class="tdr">8,800</td>
  </tr>
  <tr>
    <td>St&oacute;ra D&iacute;mun</td>
    <td class="tdr"> 68,050</td>
    <td class="tdr"> 45,600</td>
  </tr>
  <tr>
    <td>Sandoy</td>
    <td class="tdr"> 101,710</td>
    <td class="tdr"> 68,100</td>
  </tr>
  <tr>
    <td>Hestur</td>
    <td class="tdr">17,290</td>
    <td class="tdr"> 11,600</td>
  </tr>
  <tr>
    <td>Mykines</td>
    <td class="tdr"> 14,500</td>
    <td class="tdr">9,700</td>
  </tr>
  <tr>
    <td>V&aacute;gar</td>
    <td class="tdr"> 4,224</td>
    <td class="tdr">2,800</td>
  </tr>
  <tr>
    <td>Streymoy</td>
    <td class="tdr"> 27,214</td>
    <td class="tdr"> 18,200</td>
  </tr>
  <tr>
    <td>Eysturoy</td>
    <td class="tdr"> 10,520</td>
    <td class="tdr">7,000</td>
  </tr>
  <tr>
    <td>Kalsoy</td>
    <td class="tdr">14,150</td>
    <td class="tdr">9,500</td>
  </tr>
  <tr>
    <td>Vidoy</td>
    <td class="tdr"> 5,980</td>
    <td class="tdr">4,000</td>
  </tr>
  <tr>
    <td>Fugloy</td>
    <td class="tdr">22,730</td>
    <td class="tdr"> 15,200</td>
  </tr>
  <tr>
    <td class="tdp">Totals</td>
    <td class="tdr">587,333</td>
    <td class="tdr"> 393,200<a href="#Footnote_72_72" class="fnanchor">[72]</a></td>
  </tr>
</table></div>


<h4>Greenland</h4>

<p>Greenland, which has an area of 2,175,600
km<sup>2</sup> and extends for a distance of 2,670 km
from the northernmost to the southernmost
point of the country, is almost a continent by
itself. The range of the different species of seabirds,
therefore, is greatly varied, and it is
necessary to classify them according to the
relation between their distributions and the
marine zones. A description of the zones of
the marine environment in the North Atlantic
was given by Salomonsen (1965), and the
breeding distributions of seabirds in Greenland
based on this system are given in
Table 7. The terrestrial area of southernmost
West Greenland belongs to the subarctic zone
of the boreal province, and one boreal bird
species, the black-headed gull, has bred there
in recent years. It is, however, as much a
freshwater bird as a marine one.</p>


<ul>
<li class="caption">Table 7. <em>Distributions of seabirds breeding in
Greenland in relation to marine zones.</em></li>

<li class="ifrst">Marine zone and species<a name="FNanchor_73_73" id="FNanchor_73_73"></a><a href="#Footnote_73_73" class="fnanchor">[73]</a></li>

<li class="indx">Boreo-panarctic</li>
<li class="isub1"><em>Fulmarus glacialis</em></li>
<li class="isub1"><em>Somateria mollissima</em></li>
<li class="isub1"><em>Stercorarius parasiticus</em></li>
<li class="isub1"><em>Rissa tridactyla</em></li>
<li class="isub1"><em>Sterna paradisaea</em></li>
<li class="isub1"><em>Cepphus grylle</em></li>
<li class="isub1"><em>Fratercula arctica</em></li>
<li>Panarctic</li>
<li class="isub1"><em>Larus hyperboreus</em></li>
<li class="isub1"><em>Uria lomvia</em></li>
<li class="isub1"><em>Clangula hyemalis</em></li>
<li class="isub1"><em>Gavia stellata</em></li>
<li>High arctic</li>
<li class="isub1"><em>Somateria spectabilis</em></li>
<li class="isub1"><em>Branta bernicla</em> <em>(hrota)</em></li>
<li class="isub1"><em>Stercorarius longicaudus</em></li>
<li class="isub1"><em>Xema sabini</em></li>
<li class="isub1"><em>Larus thayeri</em></li>
<li class="isub1"><em>Pagophila eburnea</em></li>
<li class="isub1"><em>Cepphus grylle</em> (<em>mandti</em> group)</li>
<li class="isub1"><em>Plautus alle</em></li>
<li class="isub1"><em>Fratercula arctica</em> <em>(naumanni)</em></li>
<li class="isub1"><em>Phalaropus fulicarius</em></li>
<li>Low arctic</li>
<li class="isub1"><em>Larus glaucoides</em></li>
<li class="isub1"><em>Phalaropus lobatus</em></li>
<li>Boreo low arctic</li>
<li class="isub1"><em>Mergus serrator</em></li>
<li class="isub1"><em>Phalacrocorax carbo</em> <em>(carbo)</em></li>
<li class="isub1"><em>Larus marinus</em></li>
<li class="isub1"><em>Alca torda</em></li>
<li class="isub1"><em>Uria aalge</em></li>
<li class="isub1"><em>Cepphus grylle</em> (<em>grylle</em> group)</li>
<li class="isub1"><em>Fratercula arctica</em> <em>(arctica)</em></li>
<li>Boreal</li>
<li class="isub1"><em>Larus ridibundus</em></li>
</ul>
<p><span class="pagenum"><a name="Page_280" id="Page_280">[Pg 280]</a></span></p>
<div class="figcenter" style="width: 451px;">
<img src="images/p_280.jpg" width="451" height="700" alt="" />
<div class="caption"><p>Fig. 1. Breeding range in Greenland of four boreo-panarctic seabirds, <em>Fulmarus glacialis</em>,
<em>Somateria mollissima</em>, <em>Rissa tridactyla</em>, and <em>Fratercula arctica</em>.</p></div>
</div>

<p><span class="pagenum"><a name="Page_281" id="Page_281">[Pg 281]</a></span></p>

<div class="figcenter" style="width: 447px;">
<img src="images/p_281.jpg" width="447" height="700" alt="" />
<div class="caption"><p>Fig. 2. Breeding range in Greenland of three boreo-panarctic seabirds, <em>Sterna paradisaea</em>,
<em>Cepphus grylle</em>, and <em>Stercorarius parasiticus</em>, and one low arctic species, <em>Phalaropus
lobatus</em>.</p></div>
</div>

<p><span class="pagenum"><a name="Page_282" id="Page_282">[Pg 282]</a></span></p>

<div class="figcenter" style="width: 446px;">
<img src="images/p_282.jpg" width="446" height="700" alt="" />
<div class="caption"><p>Fig. 3. Breeding range in Greenland of three panarctic seabirds, <em>Uria lomvia</em>, <em>Larus hyperboreus</em>,
and <em>Clangula hyemalis</em>, and one high arctic species, <em>Stercorarius longicaudus</em>.</p></div>
</div>

<p><span class="pagenum"><a name="Page_283" id="Page_283">[Pg 283]</a></span></p>

<div class="figcenter" style="width: 447px;">
<img src="images/p_283.jpg" width="447" height="700" alt="" />
<div class="caption"><p>Fig. 4. Breeding range in Greenland of three boreo-low arctic seabirds, <em>Mergus serrator</em>, <em>Larus
marinus</em>, and <em>Phalacrocorax carbo</em>, and one high arctic species, <em>Plautus alle</em>.</p></div>
</div>

<p><span class="pagenum"><a name="Page_284" id="Page_284">[Pg 284]</a></span></p>

<p>The widely differing ranges of Greenland
seabirds are shown in Figs. 1-4 and are based
on my new and previously unpublished data.
The borderline between the high arctic and
low arctic zones is situated in Melville Bay on
the west coast, and just south of Scoresby
Sound on the east coast; the innermost parts
of Scoresby Sound belong to the low arctic
zone.</p>

<p>In the low arctic Pacific region the number
of seabirds is said to be about 51 million in
summer and 8 million in winter (Sowl and
Bartonek 1974). No similar estimate is available
for low arctic West Greenland, but I suggest
that it is much lower in summer and
slightly higher in winter.</p>

<p>The human population of Greenland, now
numbering about 50,000 individuals, is restricted
to the seashore, where all cities and
minor outposts are situated. Although shooting
seabirds is an ancient tradition in Greenland,
the true land-birds, which are few in number,
are usually left alone. Seabirds collected
by shooting provide an important source of
food that the Greenlanders could not do without.
Since special shooting and hunting regulations
have not been developed in Greenland,
these activities often resemble a sort of
slaughter rather than true hunting. There is
no game act in Greenland, and practically all
birds can be shot. This condition is similar to
that in Canada, where according to Section
5(7) of the Migratory Birds Regulations
(Canadian Wildlife Service, Ottawa 1973) "an
Indian or Inuk may at any time, without a
permit, take auks, auklets, guillemots,
murres, puffins and scoters and their eggs for
human food and clothing." Much the same
sort of hunting privileges exist for native
peoples of Alaska. What is still worse, however,
is the enormous illegal shooting of
ducks, geese, swans, and cranes that is known
to take place in arctic North America, but is
largely ignored by police and game authorities.
Bartonek et al. (1971) described this
situation very well for Alaska. In Greenland,
it is not possible any more to distinguish between
"native Eskimos" and Greenlanders
(including Danes working in the country), but
the attitude toward animals among the inhabitants
is the same as it has always been&mdash;a
food source to hunt and kill.</p>

<p>With a rapidly growing human population,
and a readily available supply of guns and
speedboats for hunting, the whole natural ecosystem
is beginning to break down, and it cannot
be permitted to continue. The provincial
government is aware of this fact, and various
legal enactments have been issued from both
the government and the local magistrates.
However, since the size of the police force
(mostly Greenlanders) is small, it is of little
help for the preservation of wildlife, and sometimes
even the policemen themselves do not
know the local ordinances. The result has been
that seabirds, previously profusely flourishing,
have considerably decreased in number in
West Greenland.</p>

<p>I have previously described the shooting
and hunting of seabirds in Greenland and the
statutory provisions issued to protect them
(see Salomonsen 1970). At present, the following
seabirds and their eggs are totally protected:
whooper swan; common puffin, <em>Fratercula
arctica</em>; and harlequin duck, <em>Histrionicus
histrionicus</em>. Some other species have a closed
season or are protected in certain parts of the
country: snow goose, <em>Anser caerulescens</em>;
common eider; king eider, <em>Somateria spectabilis</em>;
great cormorant; dovekie, <em>Plautus alle</em>;
black guillemot; and thick-billed murre, <em>Uria
lomvia</em>. Furthermore, all catching and hunting
of birds within 2 km of breeding colonies
of murres and kittiwakes is prohibited. Bird
sanctuaries where hunting, catching, and collecting
of eggs and down are prohibited are
Avsigsut, Nunatsiaq, and Satuarssunguit islands,
which are scattered in Disko Bay, and
Tasiussarssuaq Fjord (the inner part of Arfersiorfik
Fjord, south of Egedesminde).</p>

<p>However, the Greenland Provincial Council
has been alarmed by the serious decline in the
numbers of seabirds due to increases in
human persecution, and it has decided to introduce
a game law similar to those in Denmark
and other European countries. The preparation
of this legislation was left to me, and a
draft of this Greenland game act has been
issued (Salomonsen 1974); the new law was
passed in parliament in 1977 and went into
force on 1 January 1978.</p>

<p>It is not possible to review in detail the different
parts of the new law, but certain important
points should be mentioned. In northern
parts of West Greenland (north of Egedesminde)
the sea is ice-covered for 7-8 months a
year, and seabird hunting is therefore not pos<span class="pagenum"><a name="Page_285" id="Page_285">[Pg 285]</a></span>sible
outside the breeding season. Because of
this, it was necessary to allow some hunting
of murres, eiders, and immature gulls during
the breeding period, but away from nesting
locations. Consumption of seabirds is to be
limited to local residents, and sales to canneries
for shipment to other cities is to cease.
Previously, canneries in northwest Greenland
exported large numbers of thick-billed murres
to South Greenland&mdash;e.g., 25,606 birds in
1971; and 30,029 in 1972 (Anonymous
1974:64). This marketing of murres will end.</p>

<p>Other parts of the proposal important for
seabirds include:</p>

<p>&#8226; A general closed season extending from 15
June to 15 August.</p>

<p>&#8226; Prohibition of shooting at breeding colonies
of seabirds, as is in force at present (cf.
above).</p>

<p>&#8226; Eggs of terns and gulls can be collected for
food in southwest Greenland to 1 July, and in
northwest Greenland to 10 July; fulmar and
murre eggs can also be collected in northwest
Greenland.</p>

<p>&#8226; Each hunter is allowed to shoot or catch 50
birds per day, but the entire bag must be used
for human consumption.</p>

<p>&#8226; All shooting from speedboats, aircraft,
and motor vehicles is prohibited.</p>

<p>&#8226; Catching flightless common eiders, king
eiders, and oldsquaws <em>(Clangula hyemalis)</em> is
prohibited.</p>

<p>&#8226; Practically all seabirds and shorebirds can
be shot; all other birds (except rock ptarmigan
and raven) are totally protected.</p>

<p>The principles of this radical new act must
be taught to the population by all possible
means of communication, including radio,
public meetings, schools, etc.</p>

<hr class="tb" />

<p>Another matter of great concern to seabirds
in Greenland is the Atlantic salmon fishery
off the west coast by Danish, Greenlandic and
foreign fishermen. It is well known that many
birds are killed in the fishing gear, and a
serious political controversy has arisen, especially
between the governments of the United
States and Denmark. The fact that a large
number of thick-billed murres were drowned
in salmon gill nets during their southward
swimming migration along the Greenland
coast was significant. In a resolution sent by
the XV World Conference of the International
Council for Bird Preservation in Texel to the
Danish Government, it was stated that the
annual incidental drowning of murres probably
involved about 250,000 individuals&mdash;a
figure exceeding the reproductive capacity of
the species. This estimate was doubted by
Danish fishery biologists, but recent investigations
carried out by the Canadian Wildlife
Service and the Fisheries Research Board of
Canada have shown that the figure is even
greater, and that the total kill amounts to
about half a million murres annually (Tull et
al. 1972).</p>

<p>Because of this mortality of murres, an
agreement was reached between the American
and Danish governments, namely that:</p>

<div class="blockquot">

<p>From 1 January 1976, all salmon fisheries
outside the 12-mile boundary shall totally stop.
In the years 1972-75 the fishery carried out by
Danish and Faroese fishermen shall be reduced
gradually from 800 to 300 tons of fish, and shall
terminate on 31 December 1975. The fish quota
by Greenland fishermen must amount to no
more than 1,100 tons annually, but from 1976
onwards, the fishery shall be restricted to areas
within the 12-mile limit.</p></div>

<p>This agreement, which has drastically reduced
the number of murres caught, was discussed
at a meeting of the International Committee
of North Atlantic Fisheries in May
1972, and was ratified by the countries involved
in July 1972.</p>

<p>Oil pollution has never occurred in Greenland,
but concessions for offshore oil drilling
along the West Greenland coast have just
been granted by the Danish Government, and
this new development gives rise for concern.
However, it is clearly stated in the concession
that the Ministry for Greenland can lay down
rules for protection against oil pollution and
other damage to human or animal life, and can
adopt measures to fight pollution which has
already taken place (section 5(9)). It is up to
the concessionary to oversee industrial developments
in the area and see that marine
pollution is avoided (section 11).</p>

<p>Toxic chemicals have been found in Greenland
seabirds, as everywhere else in the world,
but it must be emphasized that no pesticides
whatsoever are in use in Greenland itself. Investigations
by Somer and Appelquist (1974)
indicated that the mercury content in black<span class="pagenum"><a name="Page_286" id="Page_286">[Pg 286]</a></span>
guillemots in Greenland has doubled over the
last 20 years, and has now reached 2 ppm,
which is, however, a relatively low figure.
Levels of DDE, PCB, and aldrin in Greenland
birds were investigated by Braestrup et al.
(1974). Common eider, king eider, harlequin
duck, and oldsquaw, as well as thick-billed
murre and great cormorant, were examined;
all were found to be contaminated with pesticides,
although to varying degrees. Highest
concentrations occurred in the cormorant,
which contained 6.5-15 ppm of DDE and 14.1-46.7
ppm of PCB. These specific differences
appear to show that the pesticide level in the
different species of seabirds is influenced
more by the position of the bird in the food
chain than by its migratory habits.</p>

<p>And finally, I wish to mention a more happy
event. On 9 May 1974 a new law of nature protection
in Greenland was passed by the
Danish Parliament. According to this law, a
National Park is to be established covering almost
the entire northeast and north regions of
Greenland, from the Thule District in northern
West Greenland around the entire north
coast of Greenland and south along the east
coast to the northern inner parts of Scoresby
Sound. All hunting, fishing, egg-collecting,
and disturbances to the environment are forbidden
in this enormous area. This is by far
the greatest National Park in the world,
covering about 800,000 km<sup>2</sup>. Of this total
area, the greater part is a lifeless icecap, to be
sure, but about 200,000 km<sup>2</sup> is ice-free land
and suitable habitat for numerous high-arctic
birds.</p>


<h4>References</h4>

<div class="hangindent">

<p>Anonymous. 1974. Gr&oslash;nland 1973, Arsberetning.
Vol. 4. Ministeriet for Gr&oslash;nland, Copenhagen.
106 pp.</p>

<p>Atkinson-Willes, G. L. 1972. The international wildfowl
censuses as a basis for wetland evaluation
and hunting rationalization. Pages 87-110 <em>in</em>
Proc. Int. Conf. Conserv. Wetlands Waterfowl,
Iran 1971.</p>

<p>Bartonek, J. C., J. G. King, and H. K. Nelson. 1971.
Problems confronting migratory birds in Alaska.
Trans. N. Am. Wildl. Nat. Resour. Conf. 36:345-361.</p>

<p>Bloch, D. 1970. The mute swan <em>(Cygnus olor)</em> breeding
in colony in Denmark. [Danish with English
summary.] Dansk Ornith. Foren. Tidsskr. 64:152-162.</p>

<p>Bloch, D. 1971. Ynglebestanden af knopsvane <em>(Cygnus
olor)</em> i Danmark 1966. Danske Vildtunders&oslash;gelser
16. 47 pp.</p>

<p>Bloch, D., C. H. Ovesen, B. H. Fenger, J. Jensen,
and J. Pedersen. 1972. Vildtreservatet Nakskov
Indrefjord. Resume af biologiske, kemiske og
fysiske underse&oslash;gelser i 1970-1971. Vildtbiologisk
Station, Kal&oslash;, R&oslash;nde. 15 pp.</p>

<p>Bogan, J. A., and W. R. P. Bourne. 1972. Organochlorine
levels in Atlantic seabirds. Nature
(Lond.) 240(5380):358.</p>

<p>Bourne, W. R. P. 1972. Threats to seabirds. Int.
Counc. Bird Preserv. Bull. 9:200-218.</p>

<p>Braestrup, L., J. Clausen, and O. Berg. 1974. DDE,
PCB and aldrin levels in arctic birds of Greenland.
Bull. Environ. Contam. Toxicol. 11(4):326-332.</p>

<p>Dyck, J. 1975. Milj&oslash;gifte i danske fugle. Panda-Nyt.
Verdensnaturfonden 1975(1):11-13.</p>

<p>Dyck, J., and H. Meltofte. 1975. The guillemot <em>Uria
aalge</em> population of the Faeroes 1972. Dansk
Ornith. Foren. Tidsskr. 69:55-64.</p>

<p>Fisher, H. I. 1973. Pollutants in North Pacific albatrosses.
Pacific Sci. 27(3):220-225.</p>

<p>Joensen, A. H. 1961. Disaster among fulmars (<em>Fulmarus
glacialis</em> (L.)) and kittiwakes (<em>Rissa tridactyle</em>
(L.)) in Danish waters. [Danish with English
summary.] Dansk Ornith. Foren. Tidsskr.
55:212-218.</p>

<p>Joensen, A. H. 1972<em>a</em>. Oil pollution and seabirds in
Denmark 1935-1968. Dan. Rev. Game Biol. 6(8):1-24.</p>

<p>Joensen, A. H. 1972<em>b</em>. Studies on oil pollution and
seabirds in Denmark 1968-1971. Dan. Rev. Game
Biol. 6(9):1-32.</p>

<p>Joensen, A. H. 1973<em>a</em>. Moult migration and wing-feather
moult of seaducks in Denmark. Dan. Rev.
Game Biol. 8(4):1-42.</p>

<p>Joensen, A. H. 1973<em>b</em>. Danish seabird disasters in
1972. Mar. Pollut. Bull. 4(8):117-118.</p>

<p>Joensen, A. H. 1974. Waterfowl populations in Denmark
1965-1973. A survey of the non-breeding
populations of ducks, swans and coot and their
shooting utilization. Dan. Rev. Game Biol. 9(1):1-206.</p>

<p>J&oslash;rgensen, O. H., and I. Kraul. 1974. Eggshell
parameters, and residues of PCB and DDE in
eggs from Danish herring gulls, <em>Larus a. argentatus</em>.
Ornis Scand. 5(2):173-179.</p>

<p>Koeman, J. H., W. S. M. van de Ven, J. J. M. de
Goejj, P. S. Tjioe, and J. L. van Haaften. 1975.
Mercury and selenium in marine mammals and
birds. Sci. Total Environ. 3:279-287.</p>

<p>Mardal, W. 1974. Ternegruppen. Feltornithologen
16:4-7.</p>

<p>Mueller, W. J., and R. M. Leach, Jr. 1974. Effects of
chemicals on egg shell formation. Annu. Rev.
Pharmacol. 14:289-303.</p>

<p>Nowak, E. 1973. Hunting kill statistics investigation.
Int. Waterfowl Res. Bur. Bull. 35:26-30.</p>

<p>Parslow, J. L. F., and D. J. Jefferies. 1973. Relationship
between organochlorine residues in livers
and whole bodies of guillemots. Environ. Pollut.
5:87-101.</p>

<p><span class="pagenum"><a name="Page_287" id="Page_287">[Pg 287]</a></span></p>

<p>Peakall, D. B. 1970. Pesticides and the reproduction
of birds. Sci. Am. 222(4):72-78.</p>

<p>Salomonsen, F. 1935. Ayes. <em>In</em> R. Sp&auml;rck, ed. Zoology
of the Faroes 3(pt. 2) (lxiv):269 + 6 suppl. A. F.
H&oslash;st, Copenhagen.</p>

<p>Salomonsen, F. 1945. De videnskabelige reservater
i Danmark og deres Fugleliv. Fauna och Flora
40:250-261.</p>

<p>Salomonsen, F. 1954. The Danish game-statistics.
[Danish with English summary.] Dansk Ornith.
Foren. Tidsskr. 48:123-126.</p>

<p>Salomonsen, F. 1955. The food production in the
sea and the annual cycle of Faeroese marine
birds. Oikos 6(1):92-100.</p>

<p>Salomonsen, F. 1963. Oversight over Danmarks
fugle. [With explanatory notes in English.]
Munksgaard, Copenhagen. 156 pp.</p>

<p>Salomonsen, F. 1965. The geographical variation of
the fulmar <em>(Fulmarus glacialis)</em> and the zones of
marine environment in the North Atlantic. Auk
82:327-355.</p>

<p>Salomonsen, F. 1968. The moult migration. Wildfowl
19:5-24.</p>

<p>Salomonsen, F. 1970. Birds useful to man in Greenland.
Pages 169-175 <em>in</em> Productivity and conservation
in northern circumpolar lands. International
Union for Conservation of Nature and
Natural Resources, Morges, Switzerland.</p>

<p>Salomonsen, F. 1974. Forslag til vedt&aelig;gt om jagt
p&aring; fuglene i Gr&oslash;nland. Diskussionsopl&aelig;g til
kommunalbestyrelserne udarbejdet p&aring; opfordring
af Det gr&oslash;nlandske Landsr&aring;d. Tidsskr.
Gr&oslash;nland 1974(5):155-172.</p>

<p>Somer, E., and H. Appelquist. 1974. Changes and
differences in mercury level in the Baltic and
Kattegat compared to the North Atlantic using
<em>Uria</em> sp. (guillemot sp.) and <em>Cepphus grylle</em> (black
guillemot) as indicators. 9th Conf. of the Baltic
Oceanographers, Kiel. 12 pp. Danish Isotope
Centre, Copenhagen.</p>

<p>Sowl, L. W., and J. C. Bartonek. 1974. Seabirds&mdash;Alaska's
most neglected resource. Trans. N. Am.
Wildl. Nat. Resour. Conf. 39:117-126.</p>

<p>Strandgaard, H. 1964. The Danish bag record. I.
Studies in game geography based on the Danish
bag record for the years 1956-57 and 1957-58.
Dan. Rev. Game Biol. 4(2):1-116.</p>

<p>Tull, C. E., P. Germain, and A. W. May. 1972. Mortality
of thick-billed murres in the West Greenland
salmon fishery. Nature (Lond.) 237(5349):42-44.</p></div>

<p><span class="pagenum"><a name="Page_288" id="Page_288">[Pg 288]</a></span></p>


<div class="footnotes"><h4>FOOTNOTES:</h4>

<div class="footnote">

<p><a name="Footnote_67_67" id="Footnote_67_67"></a><a href="#FNanchor_67_67"><span class="label">[67]</span></a> Not counted.</p></div>

<div class="footnote">

<p><a name="Footnote_68_68" id="Footnote_68_68"></a><a href="#FNanchor_68_68"><span class="label">[68]</span></a> No estimate, but number insignificant.</p></div>

<div class="footnote">

<p><a name="Footnote_69_69" id="Footnote_69_69"></a><a href="#FNanchor_69_69"><span class="label">[69]</span></a> Species totally protected.</p></div>

<div class="footnote">

<p><a name="Footnote_70_70" id="Footnote_70_70"></a><a href="#FNanchor_70_70"><span class="label">[70]</span></a> <em>Branta bernicla</em> is fully protected since 1972.</p></div>

<div class="footnote">

<p><a name="Footnote_71_71" id="Footnote_71_71"></a><a href="#FNanchor_71_71"><span class="label">[71]</span></a> <em>Somateria mollissima</em>, <em>Melanitta nigra</em>, and <em>M. fusca</em>.</p></div>

<div class="footnote">

<p><a name="Footnote_72_72" id="Footnote_72_72"></a><a href="#FNanchor_72_72"><span class="label">[72]</span></a> The "number of pairs" is calculated by
multiplying the number of birds observed
by 0.67 (Dyck and Meltofte 1975).</p></div>

<div class="footnote">

<p><a name="Footnote_73_73" id="Footnote_73_73"></a><a href="#FNanchor_73_73"><span class="label">[73]</span></a> A few species breed near freshwater lakes,
but are marine during the nonbreeding season.</p>

<p><span class="pagenum"><a name="Page_289" id="Page_289">[Pg 289]</a></span></p></div></div>
<div class="chapter"></div>
<hr class="chap" />




<h3><a name="Present_Status_and_Trends_in_Population_of" id="Present_Status_and_Trends_in_Population_of">Present Status and Trends in Population of
Seabirds in Norway</a></h3>


<p class="center">by</p>

<p class="center">Einar Brun<a name="FNanchor_74_74" id="FNanchor_74_74"></a><a href="#Footnote_74_74" class="fnanchor">[74]</a></p>

<p class="center">University of Troms&oslash;<br />
Troms&oslash;, Norway
</p>


<h4>Abstract</h4>

<div class="blockquot">

<p>The most numerous seabird in Norway is the puffin <em>(Fratercula arctica)</em>, but its
current breeding population of 1.25 million pairs is slowly declining. The kittiwake
<em>(Rissa tridactyla)</em>, however, is increasing and establishing new colonies; its
population now stands at 510,000 pairs. The population of the common murre
<em>(Uria aalge)</em>, the seabird species most vulnerable to human activity, was about
160,000 breeding pairs in 1964 but is now decreasing at a rate of nearly 5% per
year. Of the other alcids, the razorbill <em>(Alca torda)</em> and thick-billed murre <em>(Uria
lomvia)</em> show similar declines, and the black guillemot <em>(Cepphus grylle)</em> is maintaining
a stable population. The fulmar <em>(Fulmarus glacialis)</em> and the gannet <em>(Sula
bassana)</em> have both spread from the British Isles and have established a number
of breeding colonies in Norway during this century. Evidently immigration of
gannets is still occurring, since the observed rate of increase far exceeds the
population's intrinsic rate of increase. The impact of human activity on bird mortality
varies from species to species. The two most serious factors are coastal oil
pollution and the use of fishing gear; direct hunting pressure accelerates the
decline of murres and razorbills. Persistent toxic chemicals are not yet a serious
problem in Norway.</p></div>

<p>Norway, with a coastline of more than
20,000 km, an abundance of islands, and areas
of offshore upwelling, provides good conditions
for a rich seabird fauna. A regional
study of this seabird fauna has been undertaken
as a sideline of basic marine research.
Although the ultimate aim has been to evaluate
the importance of seabirds in the energy
flow of a marine ecosystem, a more realistic
problem (given priority so far) has been to
study yearly production and the dynamics behind
changes in the breeding populations.</p>

<p>Good population estimates are of fundamental
importance to studies of population
dynamics. Because the available censuses of
seabirds in Norway were few and largely inadequate,
a long-term program was started in
1961. In the beginning, resources and assistance
were very limited, and the work was concentrated
on cliff-breeding seabirds, particularly
the gannet <em>(Sula bassana)</em>, fulmar <em>(Fulmarus
glacialis)</em>, kittiwake <em>(Rissa tridactyla)</em>,
razorbill <em>(Alca torda)</em>, common murre <em>(Uria
aalge)</em>, thick-billed murre <em>(U. lomvia)</em>, and
puffin <em>(Fratercula arctica)</em>. Until 1970, the
study involved making annual censuses in the
approximately 20 major colonies of cliff-breeding
seabirds and mapping the distribution
of the quantitatively less important
colonies.</p>

<p>Since 1970, the Norwegian seabird program
has also involved more detailed studies in
some selected colonies. In these colonies, emphasis
has been on investigation of yearly production
and of the factors limiting this production,
and evaluation of the effects of
human activity on the population growth.</p>
<p><span class="pagenum"><a name="Page_290" id="Page_290">[Pg 290]</a></span></p>


<h4>Material and Methods</h4>

<p>The logistics of census operations have
gradually improved from the use of slow, local
transportation to the use of fast pneumatic
boats and, in more recent years, seaplanes.
Various census methods have been used, depending
on species and circumstances.</p>

<p>For puffins, a method based on measurement
of feeding frequency and on the number
of puffins per time unit that pass a particular
observation post when they return from the
feeding ground was used (Brun 1971<em>a</em>). Kittiwakes
and gannets were readily censused by a
combination of photographic methods and detailed
counts in sample areas (Brun 1971<em>b</em>).
Direct counting is by far the most accurate
method for razorbills, murres, and fulmars;
but in the larger colonies of common murre,
lack of time permitted accurate counts for
only a limited proportion of the cliff. Direct
counts of individuals, the egg/chick ratio, and
estimates of the relative size of the censused
population were used to estimate the total
population of the colony.</p>

<p>In a colony of kittiwakes near Troms&oslash;, environmental
factors that limit breeding success,
such as temperature and wind exposure, were
monitored throughout the breeding season on
a data recorder, and detailed measurements of
temperatures on and inside the eggs have
been recorded. For further information about
the influence of environmental parameters on
incubation rhythm and nest attendance, the
presence of the male and female at a particular
nest was recorded by using radioactive
bands and a Geiger-Muller tube connected to
a pen recorder.</p>

<p>In a study of the effects of human activity,
egg samples of selected species were analyzed
for mercury, PCB, and DDT derivates. An effort
was also made to obtain figures for the
mortality caused by oil pollution and fishing
gear as well as by direct hunting pressure.</p>


<h4>Results</h4>

<h5><em>Status and Trends of
Cliff-breeding Species</em></h5>


<h6>Puffin <em>(Fratercula arctica)</em></h6>

<p>By far the most numerous seabird in Norway
is the puffin (Fig. 1), which is the only
species with a breeding population of more
than 1 million breeding pairs (Tables 1, 2). In
a 1964 census (Brun 1966) the total breeding
population was put at 1.5 million pairs. The
current figure of 1.25 million pairs includes
several newly discovered colonies and some
not censused in 1964; it is more accurate than
the previous census for most of the 15 largest
colonies which make up 99.9% of the total
population. The puffin population is concentrated
in Troms and Nordland (94%), with
only about 3% in Finnmark.</p>

<div class="figcenter" style="width: 504px;">
<img src="images/p_290.jpg" width="504" height="700" alt="" />
<div class="caption"><p>Fig. 1. Distribution of the puffin <em>(Fratercula arctica)</em>
in Norway. Numbers refer to localities listed in
Table 2.</p></div>
</div>


<h6>Kittiwake <em>(Rissa tridactyla)</em></h6>

<p>The second most numerous seabird species
in Norway is the kittiwake, which dominates
in a number of the larger cliff colonies. Its distribution
pattern differs from that of the puffin&mdash;the
main occurrence of the kittiwake
population (about 63%) is in Finnmark
(Table 3).</p>

<p><span class="pagenum"><a name="Page_291" id="Page_291">[Pg 291]</a></span></p>

<div class="center">
<table border="0" cellpadding="4" cellspacing="0" summary="Table 1.
Estimate of the numbers of seabirds breeding on the coast of Norway
1970-1974.">
    <caption>Table 1. <em>Estimate of the numbers of seabirds breeding on
    the coast of Norway 1970-1974. Species are listed in descending
    order of breeding population size.</em></caption>
  <tr>
    <th>Species</th>
    <th>Thousands of pairs<a name="FNanchor_75_75" id="FNanchor_75_75"></a><a href="#Footnote_75_75" class="fnanchor">[75]</a></th>
    <th>Increase (+) or decline (-)</th>
  </tr>
  <tr>
    <td><em>Fratercula arctica</em></td>
    <td class="tdr">1250</td>
    <td class="tdc">-</td>
  </tr>
  <tr>
    <td><em>Rissa tridactyla</em></td>
    <td class="tdr">510</td>
    <td class="tdc">+</td>
  </tr>
  <tr>
    <td><em>Larus argentatus</em></td>
    <td class="tdr">(260)<a name="FNanchor_76_76" id="FNanchor_76_76"></a><a href="#Footnote_76_76" class="fnanchor">[76]</a></td>
    <td class="tdc">+</td>
  </tr>
  <tr>
    <td><em>L. canus</em></td>
    <td class="tdr">(150)<a href="#Footnote_76_76" class="fnanchor">[76]</a></td>
    <td class="tdc">+</td>
  </tr>
  <tr>
    <td><em>Uria aalge</em></td>
    <td class="tdr">100</td>
    <td class="tdc">-</td>
  </tr>
  <tr>
    <td><em>L. marinus</em></td>
    <td class="tdr">(40)<a href="#Footnote_76_76" class="fnanchor">[76]</a></td>
    <td class="tdc">+</td>
  </tr>
  <tr>
    <td><em>Phalacrocorax aristotelis</em></td>
    <td class="tdr">33</td>
    <td class="tdc">+</td>
  </tr>
  <tr>
    <td><em>Alca torda</em></td>
    <td class="tdr">30</td>
    <td class="tdc">-</td>
  </tr>
  <tr>
    <td><em>Cepphus grylle</em></td>
    <td class="tdr">22</td>
    <td class="tdc">0</td>
  </tr>
  <tr>
    <td><em>Sterna paradisaea</em></td>
    <td class="tdr">(21)<a href="#Footnote_76_76" class="fnanchor">[76]</a></td>
    <td class="tdc">-</td>
  </tr>
  <tr>
    <td><em>S. hirundo</em></td>
    <td class="tdr">(13)<a href="#Footnote_76_76" class="fnanchor">[76]</a></td>
    <td class="tdc">-</td>
  </tr>
  <tr>
    <td><em>Phalacrocorax carbo</em></td>
    <td class="tdr">12</td>
    <td class="tdc">+</td>
  </tr>
  <tr>
    <td><em>L. fuscus</em></td>
    <td class="tdr">9<a href="#Footnote_76_76" class="fnanchor">[76]</a></td>
    <td class="tdc">+</td>
  </tr>
  <tr>
    <td><em>Stercorarius parasiticus</em></td>
    <td class="tdr">(8)</td>
    <td class="tdc">0</td>
  </tr>
  <tr>
    <td><em>L. ridibundus</em></td>
    <td class="tdr">4<a href="#Footnote_76_76" class="fnanchor">[76]</a></td>
    <td class="tdc">+</td>
  </tr>
  <tr>
    <td><em>Fulmarus glacialis</em></td>
    <td class="tdr">1.1</td>
    <td class="tdc">+</td>
  </tr>
  <tr>
    <td><em>U. lomvia</em></td>
    <td class="tdr">1.0</td>
    <td class="tdc">-</td>
  </tr>
  <tr>
    <td><em>Hydrobates pelagicus</em></td>
    <td class="tdc">?</td>
    <td class="tdc">?</td>
  </tr>
  <tr>
    <td><em>Sula bassana</em></td>
    <td class="tdr">0.76</td>
    <td class="tdc">+</td>
  </tr>
  <tr>
    <td><em>Oceanodroma leucorrhoa</em></td>
    <td class="tdc">?</td>
    <td class="tdc">?</td>
  </tr>
</table></div>

<p>The annual production of kittiwakes shows
enormous variation, both throughout the
coastline and in different years; however, at
our sample stations in north Norway, the
mean production in 1974 (Table 4) was more
stable and was near the minimum value necessary
to maintain zero population growth.</p>

<p>This minimum production, m<sub>x</sub> (number of
females produced per breeding female), can be
computed from survival rates</p>

<p class="center">m<sub>x</sub> = (1-P)/1<sub>x</sub> = 0.13/0.57 = 0.23
</p>

<p>where P is annual adult survival and 1<sub>x</sub> is survival
of fledged chicks up to first breeding.
Data on survival are taken from Coulson and
White (1959) and from Norwegian banding
recoveries.</p>

<p>The kittiwake has, however, established a
number of new colonies, and although the
local increase in some of these is spectacular,
the long-term change during the last 15 years
is only about 1% increase per year in northern
Norwegian colonies (E. Brun, unpublished
data). In southern Norway, the population
has increased much more rapidly (Brun
1971<em>c</em>).</p>


<h6>Common murre <em>(Uria aalge)</em></h6>

<p>The common murre (Fig. 2) has shown a
considerable decrease. The most spectacular
decrease is at S&oslash;r-Fugl&oslash;y, where a colony of
10,000 pairs in 1940 was reduced to 4,000
pairs in 1961, to 1,100 pairs in 1966, and to
only about 10 breeding pairs in 1974 (Table 5).
Most of the census work was done in 1964 and
1974. The general trend in population change,
as expressed by the yearly decrease or increase,
has been extrapolated forward to 1974
or back to 1964 for those colonies where censuses
were missing for either of these years, to
enable a better comparison (Table 6). The
overall decrease in Norwegian colonies of the
common murre is, thus, near 5% per year; the
few cases with a positive trend are based
either on very small figures or on extrapolation
from old, inadequate censuses.</p>


<h6>Thick-billed Murre <em>(Uria lomvia)</em></h6>

<p>The thick-billed murre (Fig. 3) was first
proved to breed in Norwegian colonies in
1964; it was then found at three localities and
has since been found breeding at eight localities
(Table 7). It is now fairly certain that the
thick-billed murre is not a newcomer but has
remained unnoticed among the common
murre for generations, possibly since the
original immigration of the <em>Uria</em> species after
the last glacial period. Data are not sufficient
to show whether this small population of
thick-billed murres is decreasing at the same
rate as the common murre.</p>

<p><span class="pagenum"><a name="Page_292" id="Page_292">[Pg 292]</a></span></p>

<div class="center">
<table border="0" cellpadding="4" cellspacing="0" summary="Table 2. Status of the puffin (Fratercula arctica) in Norway">
    <caption>Table 2. <em>Status of the puffin</em> (Fratercula arctica) <em>in Norway (cf. Fig. 1)</em>.</caption>
  <tr>
    <th></th>
    <th>Locality</th>
    <th>Year of census</th>
    <th>Number of pairs</th>
    <th>Percent of population</th>
  </tr>
  <tr>
    <td class="tdr">1.</td>
    <td>Kj&oslash;r</td>
    <td class="tdc">1975</td>
    <td class="tdr">80</td>
    <td class="tdr">&lt;0.1</td>
  </tr>
  <tr>
    <td class="tdr">2.</td>
    <td>Heglane</td>
    <td class="tdc">1970</td>
    <td class="tdr">4</td>
    <td class="tdr">&lt;0.1</td>
  </tr>
  <tr>
    <td class="tdr">3.</td>
    <td>Ferkingstad&oslash;yene</td>
    <td class="tdc">1970</td>
    <td class="tdr">5</td>
    <td class="tdr">&lt;0.1</td>
  </tr>
  <tr>
    <td class="tdr">4.</td>
    <td>Utsira</td>
    <td class="tdc">1970</td>
    <td class="tdr">2</td>
    <td class="tdr">&lt;0.1</td>
  </tr>
  <tr>
    <td class="tdr">5.</td>
    <td>Utv&aelig;r</td>
    <td class="tdc">1970</td>
    <td class="tdr">200</td>
    <td class="tdr">&lt;0.1</td>
  </tr>
  <tr>
    <td class="tdr">6.</td>
    <td>Ryggsteinen</td>
    <td class="tdc">1970</td>
    <td class="tdr">2</td>
    <td class="tdr">&lt;0.1</td>
  </tr>
  <tr>
    <td class="tdr">7.</td>
    <td>Veststeinen</td>
    <td class="tdc">1970</td>
    <td class="tdr">1,500</td>
    <td class="tdr">0.1</td>
  </tr>
  <tr>
    <td class="tdr">8.</td>
    <td>Einevarden</td>
    <td class="tdc">1970</td>
    <td class="tdr">1,500</td>
    <td class="tdr">0.1</td>
  </tr>
  <tr>
    <td class="tdr">9.</td>
    <td>Svin&oslash;y</td>
    <td class="tdc">1970</td>
    <td class="tdr">100</td>
    <td class="tdr">&lt;0.1</td>
  </tr>
  <tr>
    <td class="tdr">10.</td>
    <td>Runde</td>
    <td class="tdc">1974</td>
    <td class="tdr">30,000</td>
    <td class="tdr">2.4</td>
  </tr>
  <tr>
    <td class="tdr">11.</td>
    <td>Sal&oslash;y</td>
    <td class="tdc">1970</td>
    <td class="tdr">2</td>
    <td class="tdr">0.1</td>
  </tr>
  <tr>
    <td class="tdr">12.</td>
    <td>Sklinna</td>
    <td class="tdc">1974</td>
    <td class="tdr">2,000</td>
    <td class="tdr">0.2</td>
  </tr>
  <tr>
    <td class="tdr">13.</td>
    <td>Lovunden</td>
    <td class="tdc">1968</td>
    <td class="tdr">60,000</td>
    <td class="tdr">4.8</td>
  </tr>
  <tr>
    <td class="tdr">14.</td>
    <td>Fugl&oslash;y i Gildesk&aring;l</td>
    <td class="tdc">1968</td>
    <td class="tdr">800</td>
    <td class="tdr">0.1</td>
  </tr>
  <tr>
    <td class="tdr">15.</td>
    <td>R&oslash;st</td>
    <td class="tdc">1964</td>
    <td class="tdr">700,000</td>
    <td class="tdr">55.7</td>
  </tr>
  <tr>
    <td class="tdr">16.</td>
    <td>V&aelig;r&oslash;y</td>
    <td class="tdc">1974</td>
    <td class="tdr">70,000</td>
    <td class="tdr">5.6</td>
  </tr>
  <tr>
    <td class="tdr">17.</td>
    <td>Nykv&aring;g</td>
    <td class="tdc">1967</td>
    <td class="tdr">40,000</td>
    <td class="tdr">3.2</td>
  </tr>
  <tr>
    <td class="tdr">18.</td>
    <td>Frugga</td>
    <td class="tdc">1975</td>
    <td class="tdr">5,000</td>
    <td class="tdr">0.4</td>
  </tr>
  <tr>
    <td class="tdr">19.</td>
    <td>Anda</td>
    <td class="tdc">1970</td>
    <td class="tdr">10,000</td>
    <td class="tdr">0.8</td>
  </tr>
  <tr>
    <td class="tdr">20.</td>
    <td>Bleik</td>
    <td class="tdc">1968</td>
    <td class="tdr">40,000</td>
    <td class="tdr">3.2</td>
  </tr>
  <tr>
    <td class="tdr">21.</td>
    <td>S&oslash;r-Fugl&oslash;y</td>
    <td class="tdc">1968</td>
    <td class="tdr">40,000</td>
    <td class="tdr">3.2</td>
  </tr>
  <tr>
    <td class="tdr">22.</td>
    <td>Nord-Fugl&oslash;y</td>
    <td class="tdc">1967</td>
    <td class="tdr">218,000</td>
    <td class="tdr">17.3</td>
  </tr>
  <tr>
    <td class="tdr">23.</td>
    <td>Loppa</td>
    <td class="tdc">1968</td>
    <td class="tdr">180</td>
    <td class="tdr">&lt;0.1</td>
  </tr>
  <tr>
    <td class="tdr">24.</td>
    <td>Hjelms&oslash;y</td>
    <td class="tdc">1964</td>
    <td class="tdr">20,000</td>
    <td class="tdr">1.6</td>
  </tr>
  <tr>
    <td class="tdr">25.</td>
    <td>Gjesv&aelig;rstappen</td>
    <td class="tdc">1973</td>
    <td class="tdr">18,000</td>
    <td class="tdr">1.4</td>
  </tr>
  <tr>
    <td class="tdr">26.</td>
    <td>Kongs&oslash;y</td>
    <td class="tdc">1966</td>
    <td class="tdr">30</td>
    <td class="tdr">&lt;0.1</td>
  </tr>
  <tr>
    <td class="tdr">27.</td>
    <td>Syltefjord</td>
    <td class="tdc">1966</td>
    <td class="tdr">100</td>
    <td class="tdr">&lt;0.1</td>
  </tr>
  <tr>
    <td class="tdr">28.</td>
    <td>Horn&oslash;y</td>
    <td class="tdc">1967</td>
    <td class="tdr">160</td>
    <td class="tdr">&lt;0.1</td>
  </tr>
  <tr>
    <td class="tdr">29.</td>
    <td>Rein&oslash;y</td>
    <td class="tdc">1967</td>
    <td class="tdr">40</td>
    <td class="tdr">&lt;0.1</td>
  </tr>
  <tr>
    <td></td>
    <td>Total</td>
    <td></td>
    <td class="tdr">1,257,705</td>
    <td class="tdr">100.0</td>
  </tr>
</table></div>


<h6>Razorbill <em>(Alca torda)</em></h6>

<p>Another colonial cliff-breeding alcid, the
razorbill (Fig. 4), has a distribution pattern
very similar to that of the common murre, but
the individual colonies (Table 8) are, with one
exception, smaller. The total breeding population
was estimated at 36,000 pairs in 1966-69
(Brun 1969<em>b</em>); some more recent censuses
show a definite decline, but data are not sufficient
to estimate the overall decline in the
Norwegian population. At most, the current
breeding population is 30,000 pairs.</p>


<h6>Fulmar <em>(Fulmarus glacialis)</em></h6>

<p>The fulmar is one of two species of seabirds
that have spread from colonies in the British
Isles and established themselves as breeding
birds in Norway during this century (the
other is the gannet).</p>

<p><span class="pagenum"><a name="Page_293" id="Page_293">[Pg 293]</a></span></p>

<div class="figcenter" style="width: 501px;">
<img src="images/p_293a.jpg" width="501" height="700" alt="" />
<div class="caption"><p><strong>Fig. 2.</strong> Distribution of the common murre <em>(Uria
aalge)</em> in Norway. Numbers refer to localities
listed in Tables 5 and 6.</p></div>
</div>

<div class="figcenter" style="width: 488px;">
<img src="images/p_293b.jpg" width="488" height="700" alt="" />
<div class="caption"><p><strong>Fig. 3.</strong> Distribution of the thick-billed murre <em>(Uria
lomvia)</em> in Norway. Numbers refer to localities
listed in Table 7.</p></div>
</div>

<div class="center">
<table border="0" cellpadding="4" cellspacing="0" summary="Table 3. Status of the kittiwake (Rissa tridactyla) in Norway">
    <caption>Table 3. <em>Status of the kittiwake</em> (Rissa
    tridactyla) <em>in Norway, and a comparison of distribution with
    that of the puffin</em> (Fratercula arctica).</caption>
  <tr>
    <th rowspan="3">County</th>
    <th colspan="0">Breeding pairs</th>
  </tr>
  <tr>
    <th colspan="2"><em>Rissa</em></th>
    <th colspan="0"><em>Fratercula</em></th>
  </tr>
  <tr>
    <th>Number<br />(thousands)</th>
    <th>Percent</th>
    <th>Number<br />(thousands)</th>
    <th>Percent</th>
  </tr>
  <tr>
    <td>Finnmark</td>
    <td class="tdr">321</td>
    <td class="tdr">62.9</td>
    <td class="tdr">38</td>
    <td class="tdr">3.0</td>
  </tr>
  <tr>
    <td>Troms</td>
    <td class="tdr">9</td>
    <td class="tdr">1.8</td>
    <td class="tdr">258</td>
    <td class="tdr">20.5</td>
  </tr>
  <tr>
    <td>Nordland</td>
    <td class="tdr">72</td>
    <td class="tdr">14.1</td>
    <td class="tdr">928</td>
    <td class="tdr">73.7</td>
  </tr>
  <tr>
    <td>Tr&oslash;ndelag (S, N)</td>
    <td class="tdr">1</td>
    <td class="tdr">0.2</td>
    <td class="tdr">2</td>
    <td class="tdr">0.2</td>
  </tr>
  <tr>
    <td>M&oslash;re and Romsdal</td>
    <td class="tdr">105</td>
    <td class="tdr">20.6</td>
    <td class="tdr">30</td>
    <td class="tdr">2.4</td>
  </tr>
  <tr>
    <td>Sogn and Fjordane</td>
    <td class="tdr">1.9</td>
    <td class="tdr">0.4</td>
    <td class="tdr">3</td>
    <td class="tdr">0.2</td>
  </tr>
  <tr>
    <td>Rogaland</td>
    <td class="tdr">0.1</td>
    <td class="tdc">&mdash;</td>
    <td class="tdr">&lt;0.1</td>
    <td class="tdc">&mdash;</td>
  </tr>
  <tr>
    <td class="tdp">Total</td>
    <td class="tdr">510</td>
    <td class="tdr">100.0</td>
    <td class="tdr">1,259</td>
    <td class="tdr">100.0</td>
  </tr>
</table></div>

<p>The fulmar began nesting in the early
1920's on Runde, the only sizeable seabird
colony in south Norway, off Alesund. Further
immigration of birds from the British Isles
probably occurred in the first 25 years, when
the population increased about 10% annually
to about 350 pairs in 1947 (Valeur 1947). Since
then the population increase has slowed down
to about 3% annually, and the population on
Runde in 1971 was about 700 pairs (Table 9).
From Runde, fulmars have spread not only to
a number of islands in the same region, but
also much farther afield&mdash;south to Utsira<span class="pagenum"><a name="Page_294" id="Page_294">[Pg 294]</a></span>
(59&deg;18'N, 4&deg;55'E) and north to Bleik (69&deg;3'N,
15&deg;42'E). The total Norwegian population of
fulmars in 1971 was estimated at 1,100 pairs.</p>

<div class="figcenter" style="width: 491px;">
<img src="images/p_294a.jpg" width="491" height="700" alt="" />
<div class="caption"><p><strong>Fig. 4.</strong> Distribution of the razorbill <em>(Alta torda)</em> in
Norway. Numbers refer to localities listed in
Table 8.</p></div>
</div>

<div class="figcenter" style="width: 483px;">
<img src="images/p_294b.jpg" width="483" height="700" alt="" />
<div class="caption"><p><strong>Fig. 5.</strong> Distribution of the gannet <em>(Sula bassana)</em> in
Norway. Numbers refer to localities listed in
Table 10.</p></div>
</div>

<div class="center">
<table border="0" cellpadding="4" cellspacing="0" summary="Table 4.
Annual production, mx, of the kittiwake (Rissa tridactyla) at some
North Norwegian coastal localities">
    <caption>Table 4. <em>Annual production, m<sub>x</sub>, of the
    kittiwake</em> (Rissa tridactyla) <em>at some North Norwegian
    coastal localities (m<sub>x</sub> = number of females produced
    per female)</em>.</caption>
  <tr>
    <th>Locality</th>
    <th>Year</th>
    <th>Sample size (number)</th>
    <th>m<sub>x</sub></th>
  </tr>
  <tr>
    <td>Ved&oslash;y, R&oslash;st</td>
    <td class="tdc">1972</td>
    <td class="tdr">852</td>
    <td class="tdr">0.21</td>
  </tr>
  <tr>
    <td>Hekkingen, Troms</td>
    <td class="tdc">1974</td>
    <td class="tdr">264</td>
    <td class="tdr">0.46</td>
  </tr>
  <tr>
    <td>Hjelms&oslash;y</td>
    <td class="tdc">1974</td>
    <td class="tdr">357</td>
    <td class="tdr">0.18</td>
  </tr>
  <tr>
    <td>Jarfjord</td>
    <td class="tdc">1974</td>
    <td class="tdr">146</td>
    <td class="tdr">0.31</td>
  </tr>
  <tr>
    <td class="tdp">Total</td>
    <td></td>
    <td class="tdr">1,619</td>
    <td class="tdr">0.25</td>
  </tr>
</table></div>


<h6>Gannet <em>(Sula bassana)</em></h6>

<p>The gannet (Fig. 5), the most recently established
and least numerous of the cliff-breeding
seabirds, has the best-known population
change. Like the fulmar, it was established in
1946 on Runde, and the first individuals were
undoubtedly of British origin. During its entire
breeding history on Runde, and also in
two of the three new colonies in northern Norway
established in the 1960's, the yearly increase
has far exceeded the intrinsic rate of increase
(Table 10); for gannets with a 50%
breeding success, adult mortality of 6%, and
35% survival up to first breeding, the intrinsic
rate of increase is about 2% per year. The
Runde and Syltefjord colonies are naturally
protected by their inaccessibility, but the
colonies at Mosken and Nordmjele, which are
on small islets, are both easily accessible. The
Nordmjele colony, however, has been effectively
protected from its start, whereas the
Mosken colony has been open to visitors; this
difference is probably reflected in their different
breeding success and annual growth
rate (Table 11). The breeding success necessary
to maintain a stable population with the<span class="pagenum"><a name="Page_295" id="Page_295">[Pg 295]</a></span>
mortality figures given above is 34%:</p>

<p class="center">m<sub>x</sub> = (1-P)/1<sub>x</sub> = 0.66/0.35 = 0.17
</p>

<p>For equal sex ratio, breeding success is 2
times m<sub>x</sub> = 0.34.</p>

<p>A British ringed gannet from Ailsa Craig
(55&deg;12'N, 5&deg;07'W) was found nesting when 4
years old in the Nordmjele colony in 1970
(Brun 1972), giving direct evidence that immigration
from colonies in Great Britain (Scotland)
still takes place.</p>

<div class="center">
<table border="0" cellpadding="4" cellspacing="0" summary="Table 5. Status of the common murre (Uria aalge) in Norway">
    <caption>Table 5. <em>Status of the common murre</em> (Uria aalge) <em>in Norway (cf. Fig. 2)</em>.</caption>
  <tr>
    <th rowspan="2" colspan="2">Locality</th>
    <th colspan="2" class="br">Last census</th>
    <th colspan="2">Previous census</th>
    <th rowspan="2">Reference</th>
  </tr>
  <tr>
    <th>Year</th>
    <th>No. of breeding pairs</th>
    <th>Year</th>
    <th>No. of breeding pairs</th>
  </tr>
  <tr>
    <td class="tdn">1.</td>
    <td>Utsira</td>
    <td class="tdc">1970</td>
    <td class="tdn">1</td>
    <td class="tdc">1950</td>
    <td class="tdn">10</td>
    <td>Holgersen 1951</td>
  </tr>
  <tr>
    <td class="tdn">2.</td>
    <td>Utv&aelig;r</td>
    <td class="tdc">1970</td>
    <td class="tdn">17</td>
    <td class="tdc">1948</td>
    <td class="tdn">55</td>
    <td>Willgohs 1952</td>
  </tr>
  <tr>
    <td class="tdn">3.</td>
    <td>Veststeinen</td>
    <td class="tdc">1970</td>
    <td class="tdn">29</td>
    <td class="tdc">1950</td>
    <td class="tdn">40</td>
    <td>Willgohs 1952</td>
  </tr>
  <tr>
    <td class="tdn">4.</td>
    <td>Klovningen</td>
    <td class="tdc">1970</td>
    <td class="tdn">35</td>
    <td class="tdc">1950</td>
    <td class="tdn">20</td>
    <td>Willgohs 1952</td>
  </tr>
  <tr>
    <td class="tdn">5.</td>
    <td>Einevarden</td>
    <td class="tdc">1970</td>
    <td class="tdn">30</td>
    <td class="tdc">1952</td>
    <td class="tdn">25</td>
    <td>Willgohs 1955</td>
  </tr>
  <tr>
    <td class="tdn">6.</td>
    <td>Runde</td>
    <td class="tdc">1974</td>
    <td class="tdn">6,000</td>
    <td class="tdc">1963</td>
    <td class="tdn">7,600</td>
    <td>Brun 1969<em>a</em></td>
  </tr>
  <tr>
    <td class="tdn">7.</td>
    <td>Storholmen</td>
    <td class="tdc">1970</td>
    <td class="tdn">8</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
  </tr>
  <tr>
    <td class="tdn">8.</td>
    <td>R&oslash;st</td>
    <td class="tdc">1974</td>
    <td class="tdn">6,800</td>
    <td class="tdc">1964</td>
    <td class="tdn">9,700</td>
    <td>Brun 1969<em>a</em></td>
  </tr>
  <tr>
    <td class="tdn">9.</td>
    <td>V&aelig;r&oslash;y</td>
    <td class="tdc">1974</td>
    <td class="tdn">1,750</td>
    <td class="tdc">1964</td>
    <td class="tdn">2,400</td>
    <td>Brun 1969<em>a</em></td>
  </tr>
  <tr>
    <td class="tdn">10.</td>
    <td>Nykv&aring;g</td>
    <td class="tdc">1974</td>
    <td class="tdn">350</td>
    <td class="tdc">1966</td>
    <td class="tdn">430</td>
    <td>Brun 1969<em>a</em></td>
  </tr>
  <tr>
    <td class="tdn">11.</td>
    <td>Bleik</td>
    <td class="tdc">1974</td>
    <td class="tdn">60</td>
    <td class="tdc">1952</td>
    <td class="tdn">90</td>
    <td>Regnell 1957</td>
  </tr>
  <tr>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">1964</td>
    <td class="tdn">75</td>
    <td>Brun 1969<em>a</em></td>
  </tr>
  <tr>
    <td class="tdn">12.</td>
    <td>S&oslash;r-Fugl&oslash;y</td>
    <td class="tdc">1974</td>
    <td class="tdn">10</td>
    <td class="tdc">1940</td>
    <td class="tdn">10,000</td>
    <td>Soot-Ryen 1941</td>
  </tr>
  <tr>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">1961</td>
    <td class="tdn">4,000</td>
    <td>Brun 1963</td>
  </tr>
  <tr>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">1966</td>
    <td class="tdn">1,100</td>
    <td>Brun 1969<em>a</em></td>
  </tr>
  <tr>
    <td class="tdn">13.</td>
    <td>Nord-Fugl&oslash;y</td>
    <td class="tdc">1967</td>
    <td class="tdn">9,000</td>
    <td class="tdc">1963</td>
    <td class="tdn">15,000</td>
    <td>L&uuml;tken 1965</td>
  </tr>
  <tr>
    <td class="tdn">14.</td>
    <td>Loppa</td>
    <td class="tdc">1974</td>
    <td class="tdn">500</td>
    <td class="tdc">1966</td>
    <td class="tdn">800</td>
    <td>Brun 1969<em>a</em></td>
  </tr>
  <tr>
    <td class="tdn">15.</td>
    <td>Hjelms&oslash;y</td>
    <td class="tdc">1974</td>
    <td class="tdn">70,000</td>
    <td class="tdc">1964</td>
    <td class="tdn">110,000</td>
    <td>Brun 1965</td>
  </tr>
  <tr>
    <td></td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdc">1967</td>
    <td class="tdn">95,000</td>
    <td>Brun 1969<em>a</em></td>
  </tr>
  <tr>
    <td class="tdn">16.</td>
    <td>Gjesv&aelig;rstappene</td>
    <td class="tdc">1973</td>
    <td class="tdn">580</td>
    <td class="tdc">1967</td>
    <td class="tdn">750</td>
    <td>Brun 1969<em>a</em></td>
  </tr>
  <tr>
    <td class="tdn">17.</td>
    <td>Sv&aelig;rholtklubben</td>
    <td class="tdc">1973</td>
    <td class="tdn">20</td>
    <td class="tdc">1966</td>
    <td class="tdn">25</td>
    <td>Brun 1969<em>a</em></td>
  </tr>
  <tr>
    <td class="tdn">18.</td>
    <td>Omgangsstauran</td>
    <td class="tdc">1973</td>
    <td class="tdn">70</td>
    <td class="tdc">1967</td>
    <td class="tdn">85</td>
    <td>Brun 1969<em>a</em></td>
  </tr>
  <tr>
    <td class="tdn">19.</td>
    <td>Syltefjorden</td>
    <td class="tdc">1974</td>
    <td class="tdn">9,000</td>
    <td class="tdc">1966</td>
    <td class="tdn">12,300</td>
    <td>Brun 1969<em>a</em></td>
  </tr>
  <tr>
    <td class="tdn">20.</td>
    <td>Horn&oslash;y</td>
    <td class="tdc">1974</td>
    <td class="tdn">500</td>
    <td class="tdc">1964</td>
    <td class="tdn">730</td>
    <td>Brun 1969<em>a</em></td>
  </tr>
  <tr>
    <td class="tdn">21.</td>
    <td>Rein&oslash;y</td>
    <td class="tdc">1974</td>
    <td class="tdn">110</td>
    <td class="tdc">1964</td>
    <td class="tdn">160</td>
    <td>Brun 1969<em>a</em></td>
  </tr>
  <tr>
    <td class="tdn">22.</td>
    <td>Kj&oslash;fjord</td>
    <td class="tdc">1970</td>
    <td class="tdn">21</td>
    <td class="tdc">&mdash;</td>
    <td class="tdc">&mdash;</td>
  </tr>
  <tr>
    <td class="tdn">23.</td>
    <td>Skoger&oslash;y</td>
    <td class="tdc">1970</td>
    <td class="tdn">8</td>
    <td class="tdc">1967</td>
    <td class="tdn">6</td>
    <td>Brun 1969<em>a</em></td>
  </tr>
  <tr>
    <td class="tdn">24.</td>
    <td>Sagfjord</td>
    <td class="tdc">1970</td>
    <td class="tdn">9</td>
    <td class="tdc">1967</td>
    <td class="tdn">12</td>
    <td>Brun 1969<em>a</em></td>
  </tr>
  <tr>
    <td class="tdn">25.</td>
    <td>Kobbholmfjorden</td>
    <td class="tdc">1970</td>
    <td class="tdn">2</td>
    <td class="tdc">1967</td>
    <td class="tdn">1</td>
    <td>Brun 1969<em>a</em></td>
  </tr>
</table></div>


<h4><em>Estimates of Total Seabird
Population in Norway</em></h4>

<p>In addition to the more detailed censuses of
the cliff-breeding species dealt with so far,
notes have been made on all seabirds observed
during numerous flights along the Norwegian
coast. Although a first attempt at putting a
figure to all seabird species in Norway may be
somewhat premature, it is believed that even
an extrapolation combined with an educated
guess is of some value until more accurate
censuses covering the whole coast can be
made. Although the data (Table 1) are
arranged in the same way as the results from
"Operation Seafarer" in the British Isles
(Cramp et al. 1974), it must be stressed that
the accuracy of the Norwegian figures, at
least for the non-cliff-breeding birds, is far inferior
to the very fine British data. The table
includes data for two petrels (<em>Hydrobates
pelagicus</em>, <em>Oceanodroma leucorrhoa</em>), which in
Norway breed on R&oslash;st (well north of the Arctic
Circle), where they have adapted to a delayed
breeding season with egg laying in
August because of the conflict of their nocturnal
habits with the continuous daylight
due to the midnight sun. Of the present population
trends that are given for each species in
Table 1, all auks except the black guillemot
<em>(Cepphus grylle)</em> are decreasing, whereas the
gulls, the gannets, and the fulmars are increasing.</p>

<p><span class="pagenum"><a name="Page_296" id="Page_296">[Pg 296]</a></span></p>


<div class="center">
<table border="0" cellpadding="4" cellspacing="0" summary="Table 6.
Population trends in colonies of the common murre (Uria aalge) in
Norway.">
    <caption>Table 6. <em>Population trends in colonies of the common
    murre</em> (Uria aalge) <em>in Norway. Numbers for 1964 and 1974
    are, when not censused those years, extrapolated from present
    trends, using estimated yearly decrease or increase from all
    available census figures.</em></caption>
  <tr>
    <th colspan="2" rowspan="2">Locality</th>
    <th colspan="2">Number of breeding pairs<a name="FNanchor_77_77" id="FNanchor_77_77"></a><a href="#Footnote_77_77" class="fnanchor">[77]</a></th>
    <th rowspan="2">Percentage yearly decrease (-) or increase (+)</th>
  </tr>
  <tr>
    <th>1964</th>
    <th>1974</th>
  </tr>
  <tr>
    <td class="tdr"> 1.</td>
    <td>Utsira</td>
    <td class="tdr">2</td>
    <td class="tdr">1</td>
    <td class="tdc">-12.2</td>
  </tr>
  <tr>
    <td class="tdr"> 2.</td>
    <td>Utv&aelig;r</td>
    <td class="tdr">23</td>
    <td class="tdr">14</td>
    <td class="tdc">-5.5</td>
  </tr>
  <tr>
    <td class="tdr"> 3.</td>
    <td>Veststeinen</td>
    <td class="tdr">32</td>
    <td class="tdr">27</td>
    <td class="tdc">-1.6</td>
  </tr>
  <tr>
    <td class="tdr"> 4.</td>
    <td>Klovningen</td>
    <td class="tdr">30</td>
    <td class="tdr">39</td>
    <td class="tdc">+2.8</td>
  </tr>
  <tr>
    <td class="tdr"> 5.</td>
    <td>Einevarden</td>
    <td class="tdr">28</td>
    <td class="tdr">31</td>
    <td class="tdc">+1.0</td>
  </tr>
  <tr>
    <td class="tdr"> 6.</td>
    <td>Runde</td>
    <td class="tdr">7,438</td>
    <td class="tdr"><em>6,000</em></td>
    <td class="tdc">-2.2</td>
  </tr>
  <tr>
    <td class="tdr"> 7.</td>
    <td>Storholmen</td>
    <td class="tdr">9</td>
    <td class="tdr">7</td>
    <td class="tdc">(-2.2)<a name="FNanchor_78_78" id="FNanchor_78_78"></a><a href="#Footnote_78_78" class="fnanchor">[78]</a>,<a name="FNanchor_79_79" id="FNanchor_79_79"></a><a href="#Footnote_79_79" class="fnanchor">[79]</a></td>
  </tr>
  <tr>
    <td class="tdr"> 8.</td>
    <td>R&oslash;st</td>
    <td class="tdr"><em>9,700</em></td>
    <td class="tdr"><em>6,800</em></td>
    <td class="tdc">-3.6</td>
  </tr>
  <tr>
    <td class="tdr"> 9.</td>
    <td>V&aelig;r&oslash;y</td>
    <td class="tdr"><em>2,400</em></td>
    <td class="tdr"><em>1,750</em></td>
    <td class="tdc">-3.2</td>
  </tr>
  <tr>
    <td class="tdr">10.</td>
    <td>Nykv&aring;g</td>
    <td class="tdr">453</td>
    <td class="tdr"><em>350</em></td>
    <td class="tdc">-2.6</td>
  </tr>
  <tr>
    <td class="tdr">11.</td>
    <td>Bleik</td>
    <td class="tdr"><em>75</em></td>
    <td class="tdr"><em>60</em></td>
    <td class="tdc">-2.3</td>
  </tr>
  <tr>
    <td class="tdr">12.</td>
    <td>S&oslash;r-Fugl&oslash;y</td>
    <td class="tdr">1,844</td>
    <td class="tdr"><em>10</em></td>
    <td class="tdc">-68.5</td>
  </tr>
  <tr>
    <td class="tdr">13.</td>
    <td>Nord-Fugl&oslash;y</td>
    <td class="tdr">13,201</td>
    <td class="tdr">3,681</td>
    <td class="tdc">-13.6</td>
  </tr>
  <tr>
    <td class="tdr">14.</td>
    <td>Loppa</td>
    <td class="tdr">900</td>
    <td class="tdr"><em>500</em></td>
    <td class="tdc">-6.1</td>
  </tr>
  <tr>
    <td class="tdr">15.</td>
    <td>Hjelms&oslash;y</td>
    <td class="tdr"><em>110,000</em></td>
    <td class="tdr"><em>70,000</em></td>
    <td class="tdc">-4.6</td>
  </tr>
  <tr>
    <td class="tdr">16.</td>
    <td>Gjesv&aelig;rstappen</td>
    <td class="tdr">853</td>
    <td class="tdr">556</td>
    <td class="tdc">-4.4</td>
  </tr>
  <tr>
    <td class="tdr">17.</td>
    <td>Sv&aelig;rholtklubben</td>
    <td class="tdr">27</td>
    <td class="tdr">19</td>
    <td class="tdc">-3.2</td>
  </tr>
  <tr>
    <td class="tdr">18.</td>
    <td>Omgangsstauran</td>
    <td class="tdr">94</td>
    <td class="tdr">68</td>
    <td class="tdc">-3.3</td>
  </tr>
  <tr>
    <td class="tdr">19.</td>
    <td>Syltefjorden</td>
    <td class="tdr">13,299</td>
    <td class="tdr"><em>9,000</em></td>
    <td class="tdc">-4.0</td>
  </tr>
  <tr>
    <td class="tdr">20.</td>
    <td>Horn&oslash;y</td>
    <td class="tdr"><em>730</em></td>
    <td class="tdr"><em>500</em></td>
    <td class="tdc">-3.9</td>
  </tr>
  <tr>
    <td class="tdr">21.</td>
    <td>Rein&oslash;y</td>
    <td class="tdr"><em>160</em></td>
    <td class="tdr"><em>110</em></td>
    <td class="tdc">-3.8</td>
  </tr>
  <tr>
    <td class="tdr">22.</td>
    <td>Kj&oslash;fjord</td>
    <td class="tdr">21</td>
    <td class="tdr">20</td>
    <td class="tdc">(-0.4)<a href="#Footnote_78_78" class="fnanchor">[78]</a>,<a href="#Footnote_79_79" class="fnanchor">[79]</a></td>
  </tr>
  <tr>
    <td class="tdr">23.</td>
    <td>Skoger&oslash;y</td>
    <td class="tdr">5</td>
    <td class="tdr">12</td>
    <td class="tdc">+10.1</td>
  </tr>
  <tr>
    <td class="tdr">24.</td>
    <td>Sagfjord</td>
    <td class="tdr">16</td>
    <td class="tdr">6</td>
    <td class="tdc">-10.1</td>
  </tr>
  <tr>
    <td class="tdr">25.</td>
    <td>Kobbholmfjord</td>
    <td class="tdr">1</td>
    <td class="tdr">5</td>
    <td class="tdc">+26.0</td>
  </tr>
  <tr>
    <td></td>
    <td class="tdp">Total</td>
    <td class="tdr">161,341</td>
    <td class="tdr">99,566</td>
    <td class="tdc">-4.9</td>
  </tr>
</table></div>

<div class="center">
<table border="0" cellpadding="4" cellspacing="0" summary="Table 7. Status of the thick-billed murre (Uria lomvia) in Norway">
    <caption>Table 7. <em>Status of the thick-billed murre</em> (Uria lomvia) <em>in Norway (cf. Fig. 3)</em>.</caption>
  <tr>
    <th colspan="2">Locality</th>
    <th>Year</th>
    <th>No. of breeding pairs</th>
    <th>Percentage of total <em>Uria</em> population</th>
  </tr>
  <tr>
    <td class="tdr">1.</td>
    <td>Ved&oslash;y, R&oslash;st</td>
    <td class="tdc">1974</td>
    <td class="tdr">15</td>
    <td class="tdr">0.3</td>
  </tr>
  <tr>
    <td class="tdr">2.</td>
    <td>V&aelig;r&oslash;y</td>
    <td class="tdc">1966</td>
    <td class="tdr">20</td>
    <td class="tdr">0.9</td>
  </tr>
  <tr>
    <td class="tdr">3.</td>
    <td>Hjelms&oslash;y</td>
    <td class="tdc">1974</td>
    <td class="tdr">850</td>
    <td class="tdr">1.2</td>
  </tr>
  <tr>
    <td class="tdr">4.</td>
    <td>Gjesv&aelig;rstappene</td>
    <td class="tdc">1973</td>
    <td class="tdr">25</td>
    <td class="tdr">4.3</td>
  </tr>
  <tr>
    <td class="tdr">5.</td>
    <td>Syltefjord</td>
    <td class="tdc">1970</td>
    <td class="tdr">90</td>
    <td class="tdr">0.9</td>
  </tr>
  <tr>
    <td class="tdr">6.</td>
    <td>Horn&oslash;y</td>
    <td class="tdc">1966</td>
    <td class="tdr">55</td>
    <td class="tdr">8.1</td>
  </tr>
  <tr>
    <td class="tdr">7.</td>
    <td>Rein&oslash;y</td>
    <td class="tdc">1964</td>
    <td class="tdr">1</td>
    <td class="tdr">0.6</td>
  </tr>
  <tr>
    <td class="tdr">8.</td>
    <td>Kj&oslash;fjord</td>
    <td class="tdc">1970</td>
    <td class="tdr">1</td>
    <td class="tdr">4.8</td>
  </tr>
  <tr>
    <td></td>
    <td>Estimated number, Norway, 1974</td>
    <td></td>
    <td class="tdr">&gt;1,000</td>
    <td class="tdr">ca. 1.0</td>
  </tr>
</table><span class="pagenum"><a name="Page_297" id="Page_297">[Pg 297]</a></span></div>

<div class="center">
<table border="0" cellpadding="4" cellspacing="0" summary="Table 8. Status of the razorbill (Alca torda) in Norway">
    <caption>Table 8. <em>Status of the razorbill</em> (Alca torda) <em>in Norway (cf. Fig. 4)</em>.</caption>
  <tr>
    <th colspan="2">Locality</th>
    <th>Year</th>
    <th>No. of breeding pairs</th>
    <th>Percent</th>
  </tr>
  <tr>
    <td class="tdr">1.</td>
    <td>Kj&oslash;r</td>
    <td class="tdc">1970</td>
    <td class="tdr">1</td>
    <td class="tdr">&lt;0.1</td>
  </tr>
  <tr>
    <td class="tdr">2.</td>
    <td>Utsira</td>
    <td class="tdc">1970</td>
    <td class="tdr">25</td>
    <td class="tdr">0.1</td>
  </tr>
  <tr>
    <td class="tdr">3.</td>
    <td>Utv&aelig;r</td>
    <td class="tdc">1970</td>
    <td class="tdr">16</td>
    <td class="tdr">0.1</td>
  </tr>
  <tr>
    <td class="tdr">4.</td>
    <td>Veststeinen</td>
    <td class="tdc">1970</td>
    <td class="tdr">22</td>
    <td class="tdr">0.1</td>
  </tr>
  <tr>
    <td class="tdr">5.</td>
    <td>Klovningen</td>
    <td class="tdc">1970</td>
    <td class="tdr">12</td>
    <td class="tdr">&lt;0.1</td>
  </tr>
  <tr>
    <td class="tdr">6.</td>
    <td>Einevarden</td>
    <td class="tdc">1970</td>
    <td class="tdr">45</td>
    <td class="tdr">0.2</td>
  </tr>
  <tr>
    <td class="tdr">7.</td>
    <td>Runde</td>
    <td class="tdc">1974</td>
    <td class="tdr">2,800</td>
    <td class="tdr">9.5</td>
  </tr>
  <tr>
    <td class="tdr">8.</td>
    <td>Sklinna</td>
    <td class="tdc">1974</td>
    <td class="tdr">15</td>
    <td class="tdr">0.1</td>
  </tr>
  <tr>
    <td class="tdr">9.</td>
    <td>Lovunden</td>
    <td class="tdc">1968</td>
    <td class="tdr">8</td>
    <td class="tdr">&lt;0.1</td>
  </tr>
  <tr>
    <td class="tdr">10.</td>
    <td>R&oslash;st</td>
    <td class="tdc">1974</td>
    <td class="tdr">3,900</td>
    <td class="tdr">13.2</td>
  </tr>
  <tr>
    <td class="tdr">11.</td>
    <td>V&aelig;r&oslash;y</td>
    <td class="tdc">1974</td>
    <td class="tdr">800</td>
    <td class="tdr">2.7</td>
  </tr>
  <tr>
    <td class="tdr">12.</td>
    <td>Nykv&aring;g</td>
    <td class="tdc">1966</td>
    <td class="tdr">250</td>
    <td class="tdr">0.8</td>
  </tr>
  <tr>
    <td class="tdr">13.</td>
    <td>Bleik</td>
    <td class="tdc">1968</td>
    <td class="tdr">28</td>
    <td class="tdr">0.1</td>
  </tr>
  <tr>
    <td class="tdr">14.</td>
    <td>S&oslash;r-Fugl&oslash;y</td>
    <td class="tdc">1974</td>
    <td class="tdr">15</td>
    <td class="tdr">0.1</td>
  </tr>
  <tr>
    <td class="tdr">15.</td>
    <td>Nord-Fugl&oslash;y</td>
    <td class="tdc">1967</td>
    <td class="tdr">10,000</td>
    <td class="tdr">33.8</td>
  </tr>
  <tr>
    <td class="tdr">16.</td>
    <td>Loppa</td>
    <td class="tdc">1969</td>
    <td class="tdr">750</td>
    <td class="tdr">2.5</td>
  </tr>
  <tr>
    <td class="tdr">17.</td>
    <td>Hjelms&oslash;y</td>
    <td class="tdc">1974</td>
    <td class="tdr">7,000</td>
    <td class="tdr">23.7</td>
  </tr>
  <tr>
    <td class="tdr">18.</td>
    <td>Gjesv&aelig;r</td>
    <td class="tdc">1973</td>
    <td class="tdr">2,500</td>
    <td class="tdr">8.5</td>
  </tr>
  <tr>
    <td class="tdr">19.</td>
    <td>Sv&aelig;rholtklubben</td>
    <td class="tdc">1973</td>
    <td class="tdr">18</td>
    <td class="tdr">0.1</td>
  </tr>
  <tr>
    <td class="tdr">20.</td>
    <td>Omgangsstauran</td>
    <td class="tdc">1973</td>
    <td class="tdr">6</td>
    <td class="tdr">&lt;0.1</td>
  </tr>
  <tr>
    <td class="tdr">21.</td>
    <td>Kongs&oslash;y</td>
    <td class="tdc">1966</td>
    <td class="tdr">8</td>
    <td class="tdr">&lt;0.1</td>
  </tr>
  <tr>
    <td class="tdr">22.</td>
    <td>Syltefjorden</td>
    <td class="tdc">1966</td>
    <td class="tdr">1,200</td>
    <td class="tdr">4.1</td>
  </tr>
  <tr>
    <td class="tdr">23.</td>
    <td>Horn&oslash;y</td>
    <td class="tdc">1967</td>
    <td class="tdr">65</td>
    <td class="tdr">0.2</td>
  </tr>
  <tr>
    <td class="tdr">24.</td>
    <td>Rein&oslash;y</td>
    <td class="tdc">1967</td>
    <td class="tdr">55</td>
    <td class="tdr">0.2</td>
  </tr>
  <tr>
    <td class="tdr">25.</td>
    <td>Kj&oslash;fjord</td>
    <td class="tdc">1970</td>
    <td class="tdr">9</td>
    <td class="tdr">&lt;0.1</td>
  </tr>
  <tr>
    <td class="tdr">26.</td>
    <td>Skoger&oslash;y</td>
    <td class="tdc">1970</td>
    <td class="tdr">4</td>
    <td class="tdr">&lt;0.1</td>
  </tr>
  <tr>
    <td class="tdr">27.</td>
    <td>Jarfjordnes</td>
    <td class="tdc">1970</td>
    <td class="tdr">3</td>
    <td class="tdr">&lt;0.1</td>
  </tr>
  <tr>
    <td></td>
    <td class="tdp">Total</td>
    <td></td>
    <td class="tdr">ca. 30,000</td>
  </tr>
</table></div>

<p>Since the coastline of Norway is about the
same length as the coastline of Great Britain
and Ireland, it is interesting to compare the
population figures (Table 12), although the
accuracy is very different. Populations of
auks and gulls are similar in both areas, but
the species composition is different. There are
more terns in the British Isles, but skuas
<em>(Catharacta skua)</em>, shags <em>(Phalacrocorax aristotelis)</em>,
and great cormorants <em>(P. carbo)</em> are
present in similar numbers. The most striking
difference is the very small number of procelli-forms
and gannets in Norway compared to
Britain and Ireland, where they are almost as
numerous as the gulls and the auks.</p>

<div class="center">
<table border="0" cellpadding="4" cellspacing="0" summary="Table 9. Status of the fulmar (Fulmarus glacialis) in Norway.">
    <caption>Table 9. <em>Status of the fulmar</em> (Fulmarus glacialis) <em>in Norway</em>.</caption>
  <tr>
    <th>County</th>
    <th>Number of localities</th>
    <th>Number of breeding pairs</th>
  </tr>
  <tr>
    <td>Nordland</td>
    <td class="tdr">6</td>
    <td class="tdr">140</td>
  </tr>
  <tr>
    <td>M&oslash;re and Romsdal</td>
    <td class="tdr">7</td>
    <td class="tdr">945</td>
  </tr>
  <tr>
    <td>Sogn and Fjordane</td>
    <td class="tdr">2</td>
    <td class="tdr">11</td>
  </tr>
  <tr>
    <td>Rogaland</td>
    <td class="tdr">2</td>
    <td class="tdr">2</td>
  </tr>
  <tr>
    <td class="tdp">Total</td>
    <td class="tdr">17</td>
    <td class="tdr">1,098</td>
  </tr>
</table><span class="pagenum"><a name="Page_298" id="Page_298">[Pg 298]</a></span></div>

<div class="center">
<table border="0" cellpadding="4" cellspacing="0" summary="Table 10. Population increase of the gannet (Sula bassana) in Norway">
    <caption>Table 10. <em>Population increase of the gannet</em> (Sula bassana) <em>in Norway (cf. Fig. 5)</em>.</caption>
  <tr>
    <th rowspan="2" colspan="2">Colony</th>
    <th rowspan="2">Year established</th>
    <th rowspan="2">Mean yearly growth rate 1969-1974<br />(%)</th>
    <th colspan="0">No. of breeding pairs</th>
  </tr>
  <tr>
    <th>1969</th>
    <th>1970</th>
    <th>1971</th>
    <th>1972</th>
    <th>1973</th>
    <th>1974</th>
  </tr>
  <tr>
    <td class="tdn">1.</td>
    <td>Runde</td>
    <td class="tdc">1946</td>
    <td class="tdn">8.4</td>
    <td class="tdn">330</td>
    <td class="tdn">331</td>
    <td class="tdn">383</td>
    <td class="tdn">422</td>
    <td class="tdn">450</td>
    <td class="tdn">494</td>
  </tr>
  <tr>
    <td class="tdn">2.</td>
    <td>Mosken</td>
    <td class="tdc">ca. 1960</td>
    <td class="tdn">5.4</td>
    <td class="tdn">50</td>
    <td class="tdn">83</td>
    <td class="tdn">77</td>
    <td class="tdn">60</td>
    <td class="tdn">62</td>
    <td class="tdn">65</td>
  </tr>
  <tr>
    <td class="tdn">3.</td>
    <td>Nordmjele</td>
    <td class="tdc">1967</td>
    <td class="tdn">83.3</td>
    <td class="tdn">7</td>
    <td class="tdn">36</td>
    <td class="tdn">65</td>
    <td class="tdn">103</td>
    <td class="tdn">127</td>
    <td class="tdn">145</td>
  </tr>
  <tr>
    <td class="tdn">4.</td>
    <td>Syltefjord</td>
    <td class="tdc">1961</td>
    <td class="tdn">14.5</td>
    <td class="tdn">28</td>
    <td class="tdn">29</td>
    <td class="tdn">44</td>
    <td class="tdn">48</td>
    <td class="tdn">51</td>
    <td class="tdn">55</td>
  </tr>
  <tr>
    <td></td>
    <td class="tdp">Total</td>
    <td></td>
    <td class="tdn">12.8</td>
    <td class="tdn">415</td>
    <td class="tdn">479</td>
    <td class="tdn">569</td>
    <td class="tdn">633</td>
    <td class="tdn">690</td>
    <td class="tdn">759</td>
  </tr>
  <tr>
    <td></td>
    <td class="tdp">Yearly growth rate (%)</td>
    <td></td>
    <td></td>
    <td></td>
    <td class="tdl">15.4</td>
    <td class="tdl">18.8</td>
    <td class="tdl">11.2</td>
    <td class="tdl">9.0</td>
    <td class="tdl">10.0</td>
  </tr>
</table></div>


<h4>Discussion</h4>


<h5><em>Impact of Human Activity</em></h5>


<h6>Direct Exploitation</h6>

<p>According to Norwegian laws, all seabirds,
with the exception (for some odd reason) of
the gannet and fulmar, can be hunted from 21
August to 1 March. However, only the two
species of murre and the razorbill are still
regularly hunted and, although no statistics
support it, an estimate based on interviews
with some of the hunters reveals that murres
and razorbills are shot in the ratio of about
50:1. One man can shoot as many as 380
murres and razorbills during a winter season
as a sideline to fishing. Although not many
hunt on this scale, an absolute minimum of
5,000 murres and razorbills are killed this way
each season.</p>

<p>A new law based on modern principles of
conservation has been under consideration for
several years, and this will mean an improvement.
However, the speed of the decline of the
auks, particularly the murres, makes it imperative
to stop this hunting immediately,
and it is of very little economic importance to
the few who take part. Some illegal "fishing"
for auks still takes place at R&oslash;st and Vaer&oslash;y,
where fishnets are anchored over wooden
frames outside the auk colonies at the beginning
of the nesting season. At Ved&oslash;y on R&oslash;st
in 1972, up to 80 murres were taken daily.
Thus an estimated total of 500-700 murres
were taken that year&mdash;about 5% of the breeding
population on this island.</p>

<p>Egg collecting was important during World
War II, but in these more affluent times and
because of the relative inaccessibility of the
auks' nests, egg collecting is now both less attractive
and less important. Human disturbance
of the breeding colonies, however, is
gradually becoming a more serious factor.</p>

<div class="center">
<table border="0" cellpadding="4" cellspacing="0" summary="Table 11.
Comparison of annual growth rate and breeding success in two colonies
of gannets (Sula bassana).">
    <caption>Table 11. <em>Comparison of annual growth rate and breeding
    success in two colonies of gannets</em> (Sula bassana).</caption>
  <tr>
    <th rowspan="2"></th>
    <th colspan="2">Mosken</th>
    <th colspan="0">Nordmjele</th>
  </tr>
  <tr>
    <th>Annual growth rate</th>
    <th>Breeding success (%)</th>
    <th>Annual growth rate</th>
    <th>Breeding success (%)</th>
  </tr>
  <tr>
    <td class="tdc">1969</td>
    <td></td>
    <td class="tdc">62</td>
    <td></td>
    <td class="tdc">&mdash;</td>
  </tr>
  <tr>
    <td></td>
    <td class="tdc">1.66</td>
    <td></td>
    <td class="tdc">5.14</td>
  </tr>
  <tr>
    <td class="tdc">1970</td>
    <td></td>
    <td class="tdc">51</td>
    <td></td>
    <td class="tdc">61</td>
  </tr>
  <tr>
    <td></td>
    <td class="tdc">0.93</td>
    <td></td>
    <td class="tdc">1.81</td>
  </tr>
  <tr>
    <td class="tdc">1971</td>
    <td></td>
    <td class="tdc">36</td>
    <td></td>
    <td class="tdc">46</td>
  </tr>
  <tr>
    <td></td>
    <td class="tdc">0.78</td>
    <td></td>
    <td class="tdc">1.58</td>
  </tr>
  <tr>
    <td class="tdc">1972</td>
    <td></td>
    <td class="tdc">33</td>
    <td></td>
    <td class="tdc">62</td>
  </tr>
  <tr>
    <td></td>
    <td class="tdc">1.03</td>
    <td></td>
    <td class="tdc">1.23</td>
  </tr>
  <tr>
    <td class="tdc">1973</td>
    <td></td>
    <td class="tdc">50</td>
    <td></td>
    <td class="tdc">35</td>
  </tr>
  <tr>
    <td></td>
    <td class="tdc">1.05</td>
    <td></td>
    <td class="tdc">1.14</td>
  </tr>
  <tr>
    <td class="tdc">1974</td>
    <td></td>
    <td class="tdc">12</td>
    <td></td>
    <td class="tdc">39</td>
  </tr>
  <tr>
    <td class="tdc">1969-1974</td>
    <td class="tdc">1.05</td>
    <td class="tdc">40</td>
    <td class="tdc">1.83</td>
    <td class="tdc">46</td>
  </tr>
</table></div>


<h6>Fishing Gear</h6>

<p>Although on a scale different from that in
western Greenland, drift-net and longline fishing
for Atlantic salmon <em>(Salmo salar)</em> outside
the 19-km (12-mile) limit off the northern Norwegian
coast present a serious mortality
hazard to some seabirds. Reliable data exist
only for the longline fisheries. In the 1969 season
(with 75 effective days from mid-March to
mid-June), one boat using 1,040 hooks per day
caught 294 birds: 52 fulmars, 3 gannets, 43
kittiwakes, 107 murres, and 89 puffins. No
razorbills were identified, but they may have
been included in the murre figure. If this
sample is representative, the 100 or so Norwegian
boats using longlines plus about 20
Danish boats (which used 4,000-6,000 hooks
per day and consequently caught more birds)
would have caught roughly 10,000 fulmars,
600 gannets, 9,000 kittiwakes, 21,000 murres,
and 18,000 puffins in the 1969 season. The
drift-nets in Norwegian waters are reported to
be less damaging to seabirds than are the
longlines, but even without adding the figures
from the drift-nets, the numbers are substantial
in view of the size of the Norwegian breeding
populations.</p>

<p><span class="pagenum"><a name="Page_299" id="Page_299">[Pg 299]</a></span></p>

<div class="center">
<table border="0" cellpadding="4" cellspacing="0" summary="Table 12.
Comparison of the number of seabirds breeding on the coasts of Great
Britain and Ireland">
    <caption>Table 12. <em>Comparison of the number of seabirds breeding
    on the coasts of Great Britain and Ireland (Cramp et al. 1974)
    and on the coast of Norway.</em></caption>
  <tr>
    <th rowspan="2">Species</th>
    <th colspan="0">Number of breeding pairs<a name="FNanchor_80_80" id="FNanchor_80_80"></a><a href="#Footnote_80_80" class="fnanchor">[80]</a></th>
  </tr>
  <tr>
    <th>Great Britain and Ireland</th>
    <th>Norway</th>
  </tr>
  <tr>
    <td><em>Fulmarus glacialis</em></td>
    <td class="tdr">306,000</td>
    <td class="tdr">1,100</td>
  </tr>
  <tr>
    <td><em>Puffinus puffinus</em></td>
    <td class="tdr">&gt; 175,000</td>
    <td class="tdc">&mdash;</td>
  </tr>
  <tr>
    <td><em>Hydrobates pelagicus</em></td>
    <td class="tdr">10<sup>5</sup> or 10<sup>6</sup></td>
    <td class="tdr">10<sup>3</sup> or 10<sup>4</sup></td>
  </tr>
  <tr>
    <td><em>Oceanodroma leucorrhoa</em></td>
    <td class="tdr">10<sup>4</sup></td>
    <td class="tdr">10<sup>2</sup></td>
  </tr>
  <tr>
    <td><em>Sula bassana</em></td>
    <td class="tdr">138,000</td>
    <td class="tdr">760</td>
  </tr>
  <tr>
    <td><em>Phalacrocorax carbo</em></td>
    <td class="tdr">8,100</td>
    <td class="tdr">12,000</td>
  </tr>
  <tr>
    <td><em>P. aristotelis</em></td>
    <td class="tdr">31,000</td>
    <td class="tdr">33,000</td>
  </tr>
  <tr>
    <td><em>Stercorarius skua</em></td>
    <td class="tdr">3,100</td>
    <td class="tdr">1<a name="FNanchor_81_81" id="FNanchor_81_81"></a><a href="#Footnote_81_81" class="fnanchor">[81]</a></td>
  </tr>
  <tr>
    <td><em>S. parasiticus</em></td>
    <td class="tdr">1,100</td>
    <td class="tdr">8,000</td>
  </tr>
  <tr>
    <td><em>Larus ridibundus</em></td>
    <td class="tdr">74,000</td>
    <td class="tdr">4,000<a name="FNanchor_82_82" id="FNanchor_82_82"></a><a href="#Footnote_82_82" class="fnanchor">[82]</a></td>
  </tr>
  <tr>
    <td><em>L. canus</em></td>
    <td class="tdr">12,000</td>
    <td class="tdr">(150,000)<a href="#Footnote_82_82" class="fnanchor">[82]</a></td>
  </tr>
  <tr>
    <td><em>L. fuscus</em></td>
    <td class="tdr">47,000</td>
    <td class="tdr">9,000<a href="#Footnote_82_82" class="fnanchor">[82]</a></td>
  </tr>
  <tr>
    <td><em>L. argentatus</em></td>
    <td class="tdr">333,000</td>
    <td class="tdr">(260,000)<a href="#Footnote_82_82" class="fnanchor">[82]</a></td>
  </tr>
  <tr>
    <td><em>L. marinus</em></td>
    <td class="tdr">22,000</td>
    <td class="tdr">(40,000)<a href="#Footnote_82_82" class="fnanchor">[82]</a></td>
  </tr>
  <tr>
    <td><em>Rissa tridactyla</em></td>
    <td class="tdr">470,000</td>
    <td class="tdr">510,000</td>
  </tr>
  <tr>
    <td><em>Sterna sandvicensis</em></td>
    <td class="tdr">12,000</td>
    <td class="tdc">&mdash;</td>
  </tr>
  <tr>
    <td><em>S. dougalli</em></td>
    <td class="tdr">2,300</td>
    <td class="tdc">&mdash;</td>
  </tr>
  <tr>
    <td><em>S. hirundo</em></td>
    <td class="tdr">14,000</td>
    <td class="tdr">(13,000)<a href="#Footnote_82_82" class="fnanchor">[82]</a></td>
  </tr>
  <tr>
    <td><em>S. paradisaea</em></td>
    <td class="tdr">(31,000)</td>
    <td class="tdr">(21,000)<a href="#Footnote_82_82" class="fnanchor">[82]</a></td>
  </tr>
  <tr>
    <td><em>S. albifrons</em></td>
    <td class="tdr">1,800</td>
    <td class="tdc">&mdash;</td>
  </tr>
  <tr>
    <td><em>Alca torda</em></td>
    <td class="tdr">(144,000)</td>
    <td class="tdr">30,000</td>
  </tr>
  <tr>
    <td><em>Uria aalge</em></td>
    <td class="tdr">(577,000)</td>
    <td class="tdr">100,000</td>
  </tr>
  <tr>
    <td><em>U. lomvia</em></td>
    <td class="tdc">&mdash;</td>
    <td class="tdr">1,000</td>
  </tr>
  <tr>
    <td><em>Cepphus grylle</em></td>
    <td class="tdr">8,300</td>
    <td class="tdr">22,000</td>
  </tr>
  <tr>
    <td><em>Fratercula arctica</em></td>
    <td class="tdr">(490,000)</td>
    <td class="tdr">1,250,000</td>
  </tr>
  <tr>
    <td class="tdp">Total</td>
    <td class="tdr">ca. 3,000,000</td>
    <td class="tdr">ca. 2,500,000</td>
  </tr>
</table></div>

<p>Use of fishing gear close inshore, especially
pound nets set near colonies of diving seabirds,
can take a heavy toll under special
weather conditions. In 1969 at Runde, 85
birds, mainly auks, shags, and some diving
ducks, were caught in one net in 24 hours; this
is an exceptionally high figure. The total loss
of diving seabirds in pound nets per year in
Norway (about 6,000 nets fishing for 40 days)
was estimated to be at least 40,000 birds in
1969. The data are too unreliable to give
species composition, however, since fishermen
rarely make note of this.</p>

<p>Amounts of fish offal from offshore
trawlers, drift-netters, and longline fishing
boats have increased in recent years, and
some seabirds, particularly kittiwakes, fulmars,
and gannets make use of this new and
readily available food source. Thus, although
the use of fishing gear is a serious threat to
seabird survival, fish waste from the same
boats provides an abundant food supply for
the more pelagic species.</p>

<p><span class="pagenum"><a name="Page_300" id="Page_300">[Pg 300]</a></span></p>


<h6>Pollution</h6>

<p>No quantitative investigation similar to
those made in Great Britain, Netherlands,
and Belgium (Tanis and Bruyns 1968) has
been carried out on the impact of oil pollution
on seabirds in Norway. The northern Norwegian
population of the most threatened
species, murres and razorbills, winter in
North Sea coastal areas where oil pollution
and oiled birds have most frequently been
found. It is possible that whole populations
winter every year in the same area, and if they
happen to be in a heavily polluted area, a particular
population may be seriously affected.
Such an occurrence is believed to have caused
the dramatic decline in the S&oslash;r Fugl&oslash;y population
(cf. Table 5).</p>

<p>Although not yet serious, pollution by persistent
toxic chemicals such as organochlorines
and mercury is a problem even in
northern Norway, because the northbound
coastal current brings water masses, plankton,
and nekton from areas with industrial
wastes. Analysis of the eggs of herring gull
<em>(Larus argentatus)</em>, murre, razorbill, and kittiwake
in 1972 showed relatively low levels of
mercury; the only species with a relatively
high level of mercury (mean 0.58 ppm) was the
gannet (Fimreite et al. 1974). This elevated
toxic burden may have caused a reduced
breeding success for the gannet. Analysis of
concentrations of PCB's and DDT/DDE
showed that the levels of these organochlorines
were generally also lower in Norwegian
seabirds than in those of Britain (Fimreite
et al. 1977).</p>


<h6>Protection and Necessary
Conservation Measures</h6>

<p>Total protection of some of the important
seabird colonies (including the surrounding
nearshore waters) has proven very effective,
especially when the protection is so strict that
landing is prohibited for a specified period
during incubation and fledging. However, to
reduce the rapid decrease of some species, a
total hunting prohibition of those species
must be instigated, oil pollution must be reduced,
and the fisheries must be regulated to
reduce the mortality caused by fishing gear.</p>


<h5><em>Natural Factors Influencing
Breeding Success</em></h5>

<p>The factors discussed so far are all results
of human activities which directly or indirectly
influence seabird mortality. Yearly production
or breeding success is, however, also
influenced by a number of natural factors
such as food supply, availability of suitable
nest sites, predation, climate (weather), and
population-dependent factors (age, breeding
experience, population density). For the
gannet, whose breeding success has been
studied in some detail (Brun 1974), it was concluded
that the differences in exposure (to
severe weather) and in breeding experience
were the most important factors responsible
for annual fluctuation in breeding success.
For such species as murres, razorbills, and
puffins, food supply is an important limiting
factor. If the spawning of the fish species that
constitute their main food items fails 1 year
for some reason, it may be very difficult for
the seabirds to find an adequate alternative
food supply and most of the chicks starve to
death. To a lesser degree, food supply is limiting
for the kittiwake, which seems to be more
influenced by bad weather (Norderhaug et al.
1977).</p>


<h4>Conclusion</h4>

<p>Two opposite population trends have been
observed&mdash;the decline of the coastal-bound
murres and razorbills and the increase and
spread of the more pelagic gannets, fulmars,
and kittiwakes. These changes are attributed
to a number of factors, which include the
following:</p>

<p>&#8226; The diving murres and razorbills spend a
major part of their time swimming on the surface
and are thus more susceptible to surface
oil pollution than are the pelagic species.</p>

<p>&#8226; The coastal-bound murres and razorbills
are quite heavily hunted, whereas there is no
regular hunting of the pelagic species.</p>

<p>&#8226; The pelagic species are mainly surface
feeders and do not swim under water, and are
thus less affected by the drift-nets than are
diving birds.</p>

<p><span class="pagenum"><a name="Page_301" id="Page_301">[Pg 301]</a></span></p>

<p>&#8226; The pelagic species are the principal beneficiaries
of recently increased supply of fish
offal from trawlers.</p>


<h4>Acknowledgments</h4>

<p>The program was originally sponsored by
Troms&oslash; Museum, later University of Troms&oslash;,
and has been financially supported by the
Norwegian Research Council for Science and
Humanities and the Norwegian Game Fund.
The research grants are gratefully acknowledged.
I also thank my field assistants and
the many local people at the breeding sites
who have been most helpful.</p>


<h4>References</h4>

<div class="hangindent">

<p>Brun, E. 1963. Ornithological features of Nord-Fugl&oslash;y
and S&oslash;r-Fugl&oslash;y. Astarte 1(22):1-13.</p>

<p>Brun, E. 1965. Brunnich's Guillemot, <em>Uria lomvia</em>
(L.), as a breeding bird in Norway. (In Norwegian,
English summary.) Sterna 6:229-250.</p>

<p>Brun, E. 1966. The breeding population of puffins
<em>(Fratercula arctica)</em> in Norway. (In Norwegian,
English summary.) Sterna 7:1-17.</p>

<p>Brun, E. 1969<em>a</em>. The breeding distribution and
population of guillemots <em>(Uria aalge)</em> in Norway.
(In Norwegian, English summary.) Sterna 8:209-224.</p>

<p>Brun, E. 1969<em>b</em>. The breeding distribution and
population of razorbills <em>(Alca torda)</em> in Norway.
(In Norwegian, English summary.) Sterna 8:345-359.</p>

<p>Brun, E. 1971<em>a</em>. Census of Puffins <em>(Fratercula arctica)</em>
on Nord-Fugl&oslash;y, Troms. Astarte 4:41-45.</p>

<p>Brun, E. 1971<em>b</em>. Breeding distribution and population
of cliff-breeding seabirds in S&oslash;r-Varanger,
north Norway. Astarte 4:53-60.</p>

<p>Brun, E. 1971<em>c</em>. Population changes of some seabirds
in south Norway. (In Norwegian, English
summary.) Sterna 10:35-56.</p>

<p>Brun, E. 1972. Establishment and population increase
of the gannet <em>Sula bassana</em> in Norway.
Ornis Scand. 3:27-38.</p>

<p>Brun, E. 1974. Breeding success of gannets <em>Sula
bassana</em> at Nordmjele, And&oslash;ya, north Norway.
Astarte 7:77-89.</p>

<p>Coulson, J. C., and E. White. 1959. The post-fledging
mortality of the kittiwake. Bird Study 6:97-102.</p>

<p>Cramp, S., W. R. P. Bourne, and D. Saunders. 1974.
The seabirds of Britain and Ireland. Collins,
London. 287 pp.</p>

<p>Fimreite, N., J. E. Bjerk, N. Kveseth, and E. Brun.
1977. DDE and PCBs in eggs of Norwegian seabirds.
Astarte 10:15-20.</p>

<p>Fimreite, N., E. Brun, A. Fr&oslash;slie, P. Frederichsen,
and N. Gundersen. 1974. Mercury in eggs of Norwegian
seabirds. Astarte 7:71-75.</p>

<p>Holgersen, H. 1951. Investigations of seabirds in
Rogaland 1949-1950. (In Norwegian, English
summary.) Stavanger Mus. Arb. 1950:61-76.</p>

<p>L&uuml;tken, E. 1965. The breeding birds on Nord-Fugl&oslash;y,
north Norway, their distribution and
numbers. (In Danish, English summary.) Dansk
Orn. For. Tidsskr. 58:166-193.</p>

<p>Norderhaug, M., E. Brun, and G. U. M&oslash;llen. 1977.
Barentshavets sj&oslash;fuglressurser. Forhold i tilknytning
til status, milj&oslash;problemer of forshningsoppgave.
Medd. Norsk Polar Inst. 104:1-119.</p>

<p>Regnell, S. 1957. Fran det nordnorska f&aring;ggelberget
Bleiks&oslash;ya. Fauna Flora, Upps. 52:199-202.</p>

<p>Soot-Ryen, T. 1941. The seabird rookeries of Troms
county. (In Norwegian, English summary.)
Troms&oslash; Mus. Aarsh. 62(1):1-112.</p>

<p>Tanis, J. J. C., and M. F. M. Bruyns. 1968. The impact
of oil pollution on seabirds in Europe. Proc.
Int. Conf. Oil Pollution of the Sea 1968:67-74.</p>

<p>Valeur, P. 1947. Havhesten og havsula p&aring; Rund&oslash;y.
Naturen 70:370-379.</p>

<p>Willgohs, J. F. 1952. On the distribution of some
seabirds in western Norway. Univ. Bergen &Aring;rb.
1951 Naturvit. Rekke (9):1-20.</p>

<p>Willgohs, J. F. 1955. Om forekomsten av endel
kyst-og sj&oslash;fugl p&aring; Vestlandet. Fauna, Oslo 8:16-27.</p></div>

<p><span class="pagenum"><a name="Page_302" id="Page_302">[Pg 302]</a></span></p>


<div class="footnotes"><h4>FOOTNOTES:</h4>

<div class="footnote">

<p><a name="Footnote_74_74" id="Footnote_74_74"></a><a href="#FNanchor_74_74"><span class="label">[74]</span></a> Deceased.</p></div>

<div class="footnote">

<p><a name="Footnote_75_75" id="Footnote_75_75"></a><a href="#FNanchor_75_75"><span class="label">[75]</span></a> Numbers in parentheses are not based on a complete census of the coast.</p></div>

<div class="footnote">

<p><a name="Footnote_76_76" id="Footnote_76_76"></a><a href="#FNanchor_76_76"><span class="label">[76]</span></a> In addition, an unknown number of pairs breeding inland.</p></div>

<div class="footnote">

<p><a name="Footnote_77_77" id="Footnote_77_77"></a><a href="#FNanchor_77_77"><span class="label">[77]</span></a> Numbers in italics were censused from 1964 and 1974.</p></div>

<div class="footnote">

<p><a name="Footnote_78_78" id="Footnote_78_78"></a><a href="#FNanchor_78_78"><span class="label">[78]</span></a> Estimated values from trends in neighboring colonies.</p></div>

<div class="footnote">

<p><a name="Footnote_79_79" id="Footnote_79_79"></a><a href="#FNanchor_79_79"><span class="label">[79]</span></a> Numbers in parentheses are not based on a complete census of the whole coast.</p></div>

<div class="footnote">

<p><a name="Footnote_80_80" id="Footnote_80_80"></a><a href="#FNanchor_80_80"><span class="label">[80]</span></a> Numbers in parentheses are not based on a complete census of the whole coast.</p></div>

<div class="footnote">

<p><a name="Footnote_81_81" id="Footnote_81_81"></a><a href="#FNanchor_81_81"><span class="label">[81]</span></a> New in 1975 (Wim Vader, personal communication).</p></div>

<div class="footnote">

<p><a name="Footnote_82_82" id="Footnote_82_82"></a><a href="#FNanchor_82_82"><span class="label">[82]</span></a> In addition, an unknown number of pairs breeding inland.</p>

<p><span class="pagenum"><a name="Page_303" id="Page_303">[Pg 303]</a></span></p></div></div>



<p><span class="pagenum"><a name="Page_304" id="Page_304">[Pg 304]</a><br /><a name="Page_305" id="Page_305">[Pg 305]</a></span></p>
<div class="chapter"></div>
<hr class="chap" />




<h2>SYMPOSIUM SUMMARY</h2>
<div class="chapter"></div>
<hr class="chap" />




<h3><a name="Conservation_of_Marine_Birds_of_Northern" id="Conservation_of_Marine_Birds_of_Northern">Conservation of Marine Birds of Northern
North America&mdash;A Summary</a></h3>


<p class="center">by</p>

<p class="center">Ian C. T. Nisbet</p>

<p class="center">Massachusetts Audubon Society<br />
Lincoln, Massachusetts 01773
</p>


<p>This is not going to be a straightforward
summary of the conference because it is my
view that a number of important topics have
not been addressed. In particular, what was
supposed to be the main theme of the conference&mdash;the
need for <em>conservation</em> of marine
birds of northern North America&mdash;has been
taken for granted by many speakers and has
been treated by others in what may be a misleadingly
brief way. So instead of simply summarizing
the information that has been presented
in the papers, I want to give my own
views about how we should use this information
to make a case for the conservation of
marine birds. I feel strongly that we can make
a good case for conserving them, and that we
know enough to start doing so. The task of
making a case for conservation and of proposing
priorities for action has been left to me as
the conference summarizer.</p>

<p>Particularly in the first half of this conference,
we heard a long series of accounts of the
birds of the area which stressed our ignorance&mdash;large
amounts of information that was
not known and large amounts of research that
needed to be done. Now, I have an unexpected
advantage over most of these speakers in that
I have very little direct experience in the area.
What I learned from their papers, not having
any very clear picture of the islands, the birds,
their habits, or the food that they eat, is that
we already know quite a lot about the marine
birds of northern North America. We certainly
know enough to decide what we ought
to do next and how to take the basic steps in
conserving them.</p>

<p>After listening to the presentations, reading
the abstracts, and studying the maps
posted in the conference hall, I drew up a list
of 10 points that I will first list and then
elaborate on.</p>

<p>&#8226; We know that we are discussing a very important
biological resource which has been
neglected for a long time.</p>

<p>&#8226; We know roughly what this resource consists
of and which aspects of it are biologically
important.</p>

<p>&#8226; We know why this resource is in its
present condition, and we know something
about the ways in which it is related to other
resources.</p>

<p>&#8226; We know a certain number of things that
the birds do which make them vulnerable to
changes in the environment.</p>

<p>&#8226; We know that the resource has already
been disturbed in the past, both by human-induced
and by natural changes, and we know
that it has already been damaged.</p>

<p>&#8226; We can identify at least some of the major
threats that the resource will face in the next
few years.</p>

<p>&#8226; We know that the resource can be conserved,
at least to a modest and partial extent.</p>

<p>&#8226; We have a fairly good idea of what we
ought to do now to start conserving the
resource.</p>

<p>&#8226; We have some ideas&mdash;so far rather rough
and ill-formulated&mdash;about why we should conserve
the resource.</p>

<p>&#8226; We know&mdash;or so I believe&mdash;that it is practicable
and economically feasible to conserve
the resource.</p>

<p>I am sure that there will be some disagreements
with some of these assertions, especially
with the last two, so I will give reasons
why I believe that we should conserve these
birds and that we can afford to do so.</p>

<p><span class="pagenum"><a name="Page_306" id="Page_306">[Pg 306]</a></span></p>


<h4>Magnitude and Importance
of the Resource</h4>

<p>The papers in the first half of the conference
which reviewed the abundance and distribution
of the birds in the northern North Pacific
Ocean, the Bering Sea, and adjacent seas suggested
that we are dealing with numbers of
birds of the order of 100 million. That is 100
million birds at sea plus some unknown number
of millions of birds along the shore. We do
not have to take these numbers literally&mdash;I
am sure that the persons who produced them
did not mean them to be taken literally&mdash;but
certainly we are talking about something on
the order of tens of millions and not much
more than some hundreds of millions. At
least, it is on the order of a hundred million
rather than ten million or a billion. I do not
think it an exaggeration to say that this is one
of the great neglected biological resources of
the world.</p>


<h4>Characteristics of the
Resource</h4>

<p>Three important aspects of this resource
have not been identified clearly in the papers
delivered at the conference, in part because
the papers summarizing the biological surveys
did not include much of the detail that
was available in the maps posted in the conference
hall. [Maps in this volume do not show
the detail of those posted.] These are the
numerical abundance of the birds, their diversity,
and their unique characteristics.</p>

<p>As to abundance, figures have been mentioned
on the order of 50 million for shearwaters
(<em>Puffinus</em> spp.) and 25 million for
murres (<em>Uria</em> spp.). For other species the
quoted numbers have been less specific, but I
would estimate from what I have read and
heard that the total population must run into
millions for eiders (<em>Somateria</em> spp.), kittiwakes
<em>(Rissea brevirostris)</em>, and fulmars <em>(Fulmarus
glacialis)</em>, and doubtless for other
species. The numbers of the smaller alcids, in
particular, must be very great.</p>

<p>As to diversity, there is an impressive number
of species and a wide variety of habitats.
We have been shown in the photographs some
spectacular island colonies, particularly in the
Bering Sea and the Aleutian Islands, some of
which have a remarkable variety of species.
Several different definitions of "seabird"
have been used at this conference, but certainly
there are dozens, and probably scores,
of genuine marine species that either breed in
the area or use it as a major nonbreeding area.
The collection of birds in the area of the North
Pacific and the Bering seas seems more impressive
in terms of both abundance and diversity
than anything in the north Atlantic
Ocean, which has been so much more fully
studied.</p>

<p>As to the uniqueness, there has been almost
no mention of the endemic species at the conference.
It is therefore important to emphasize
in this summary that a significant group
of marine or coastal birds is endemic to this
area. These birds include the red-legged kittiwake
(<em>Rissa</em> spp.), the Aleutian tern <em>(Sterna
aleutica)</em>, the spectacled eider <em>(Somateria
fischeri)</em>, the emperor goose <em>(Philacte canagica)</em>,
and the red-faced cormorant <em>(Phalacrocorax
urile)</em>; a number of alcids, including the
whiskered <em>(Aethia pygmaea)</em>, parakeet <em>(Cyclorrhynchus
psittacula)</em>, crested <em>(A. cristatella)</em>,
and least auklets <em>(A. pusilla)</em>; the
horned puffin <em>(Fratercula corniculata)</em>; and
Kittlitz's murrelet <em>(Brachyramphus brevirostris)</em>.
In addition, we should not forget some
migrants that make exclusive use of this area
in their nonbreeding season. These include the
short-tailed albatross <em>(Diomedea albatrus)</em>,
the scaled petrel <em>(Pterodroma inexpectata)</em>,
and I believe also Cook's petrel <em>(P. cookii)</em>,
which has not previously been mentioned.
From the little we know about its off-season
distribution, the short-tailed albatross appears
to use these waters exclusively; hence it
has as much claim to be regarded as an endangered
species of the United States as the
whooping crane <em>(Grus americana)</em>.</p>

<p>Perusal of the lists of species presented at
the conference brings out one important
point. Although we are meeting in the United
States and have been looking at the birds
from a United States-Canadian viewpoint,
this is truly an international resource in almost
every respect that I have mentioned.
The most abundant species, in terms of both
numbers and biomass, is probably the short<span class="pagenum"><a name="Page_307" id="Page_307">[Pg 307]</a></span>-tailed
shearwater, a migrant from the southern
hemisphere. The rarest species, and the
most endangered, is the short-tailed albatross,
which breeds only on one island in
Japan. There are migrants in large numbers
from Chile, Australia, New Zealand, and especially
the Soviet Union. All of these use the
area of ocean and shallow sea that we have
been considering as a major area for a substantial
part of their annual cycle.</p>

<p>What more do we need to know about the
extent of this resource? In my opinion we
should not place high priority on determining
the exact numbers of the birds&mdash;whether
there are 25 million or 26 million murres, for
example. It would be difficult, if not impossible,
to determine such numbers in the kind
of geographical and climatic area we are considering.
Moreover, even if we were to measure
the populations with great accuracy and
to determine in a few years that they had
changed by 10%, we would not be able to
draw any conclusions about the reasons for
the change or what should be done about it.</p>

<p>To set priorities for further exploration, I
think it is more important to survey in greater
detail the general distribution of the breeding
colonies. So far, we know the location of only
the largest colonies; we know almost nothing
about the colonies of a mere 10,000 pairs or
less. So I think future surveys should concentrate
on locating the medium-sized colonies
and getting some impression of roughly how
many smaller colonies there are. It is important
to locate and be sure that we know of all
the major colonies that have a considerable
number of species; these large, diverse colonies
should be given priority for conservation.
Most important of all, we need to locate and
survey the endemic species with some precision.
This need is especially great for the
species that we suspect are limited to small
areas or that may otherwise be particularly
vulnerable.</p>

<p>If we are to measure population changes
over the next few decades, it is of course essential
to have a good base-line survey. However,
I do not think it is either practicable or
desirable to try to inventory the entire population
of breeding seabirds with great accuracy.
A more realistic and worthwhile program
would be to select some sample colonies
and to catalogue these sample areas in some
detail, preferably with a photographic record,
so that they can be resurveyed in later years
to determine whether substantial population
changes have taken place. Criteria for selection
of sample colonies for inclusion in this
base-line survey should include not only
numerical size and species diversity but also
ease of access, ease of observation, and the
practicability of obtaining good photographic
records.</p>


<h4>Ecology and Functioning
of the Resource</h4>

<p>In the opening session of this conference,
several speakers reviewed our general knowledge
of the ecology of seabirds; others summarized
our specific knowledge of the birds of
the North Pacific, Bering, and adjacent seas,
and their relation to physical and biological
factors in the environment. There is no need
to summarize these reviews again here except
to point out that information on the relation
between the birds and the marine environment
is being generated very rapidly. We are
beginning to understand the factors that control
the breeding distribution of the individual
species, their foraging strategies, and their
dispersion at sea, at least in summer. However,
it is clear from what has been said at this
conference that we know much less about
their ecology and distribution in winter. This
lack of information is important because conflicting
opinions have been expressed as to
whether factors operating in the winter range
or at the breeding colonies are more critical in
limiting population size.</p>

<p>It is evident from what was said in the opening
session that the distribution of the birds is
very closely related to the distribution of
marine resources. It is clearly no accident that
the distribution of large numbers of many
species of birds coincides with that of the
major fisheries. Similarly, it is no accident
that there is a relation between the distribution
of birds and the extent of the continental
shelf. These coincidences, which reflect the
fundamental dependence of both birds and
fish upon marine productivity, set the stage
for existing and further conflicts between conservation
of the birds and human exploitation
of other resources of the area.</p>

<p><span class="pagenum"><a name="Page_308" id="Page_308">[Pg 308]</a></span></p>

<p>Perhaps the most significant gap in our
knowledge of North Pacific seabirds is in the
area of productivity and demography. As far
as I can judge, we know almost nothing about
the breeding success of these birds, their post-fledging
survival, their longevity, their age at
first breeding, the age structure of their populations,
the fluctuations in their breeding performance,
or their survival from year to year.
For most species, we lack even the most basic
life history and life table information.</p>

<p>If we can argue by analogy from studies
made in other parts of the world, including the
North Atlantic, we can make some basic
generalizations that we would expect to apply
to the birds of our area. We know that as a
class seabirds have some peculiar characteristics
which make them difficult to manage and
cause some of the problems we have in conserving
them. In general, they are long-lived
and breed slowly, most lay small clutches, and
the historical experience is that they take a
very long time to recover from depletion of
population. Many have an irregular breeding
performance; some have long series of bad
years interspersed with occasional years of
good breeding success. Many seabird populations
have traditionally fluctuated, as exemplified
by those of the North Atlantic, whose
fluctuations were described by W. H. Drury
and W. R. P. Bourne.</p>

<p>Some species of seabirds are conservative,
staying in the same colonies for many years or
generations. Others are volatile, dispersing
freely from one site to another and forming
new colonies in an unpredictable way. Seabirds
exhibit a wide range of ecological adaptations;
some are highly specialized, others
are highly generalized and adaptable. These
differences can be very important when their
environment changes, as D. N. Nettleship's
film "Puffins, predators, and pirates" graphically
illustrated.</p>

<p>As M. T. Myres pointed out on the 1st day
of the conference, seabird populations exhibit
both short-and long-term fluctuations. Long-term
fluctuations are those that take place
over times comparable to the generation time
of the species, which may be many years or
even decades for some seabirds. By surveying
populations and measuring changes in them,
we usually obtain information only about
long-term population trends, reflecting long-term
changes in the environment. Short-term
perturbations in the environment are usually
not reflected quickly by changes in total population&mdash;certainly
not by changes that we can
measure with the accuracy of our present-day
census techniques. Many of the man-made
changes we are concerned about are short-term.
To identify their effects we should look
not for changes in total population but rather
for changes in biological parameters, such as
the first-year survival rate or the number of
young raised. I therefore suggest that some of
the most critical parameters to be measured
are changes in age structure of populations.
We should therefore select as biological monitors
species that can readily be aged&mdash;for
example, gulls, which have a sequence of distinguishable
immature plumages.</p>

<p>In specifying gaps in our knowledge of the
ecology of birds, we should set clear priorities
rather than compile a long "shopping list" of
research projects. On the basis of the foregoing
survey, I would suggest the following
as priority items for further study. First, we
need to know a lot more about winter distribution,
not only of the marine birds, but also of
inshore and coastal species. Second, we need
to study in greater detail the relation between
the day-to-day distribution of birds and the
local patchiness of the resources on which
they depend. Evidence that seabirds are able
to locate and use fluctuating and shifting food
sources has been given by several speakers at
the conference. We need to understand how
birds locate these resources and what relation
this has to their survival and vulnerability to
human activities. There is a special need to
study the ecology of endemic species because
their conservation is of special importance.
We need to learn more about the relation of
the birds to the commercial fisheries, both to
resolve existing or alleged conflicts and to
avert future problems.</p>

<p>However, I believe that the highest research
priority should be given to obtaining
basic information on reproductive success and
life table data for some representative species.
Clearly, we cannot study many species in detail,
but in selecting key species for such
studies we should pick a variety of ecological
types&mdash;for example, at least one generalist
species and one specialist, one sedentary
species and one migrant, one species at a high<span class="pagenum"><a name="Page_309" id="Page_309">[Pg 309]</a></span>
trophic level and one at a low trophic level.</p>

<p>For the purpose for which we convened this
symposium&mdash;conservation&mdash;I do not think
that we need detailed knowledge of the factors
which regulate populations. Such knowledge
is, of course, of immense biological interest
and will ultimately be needed for effective
long-term management. However, it does not
have immediate or even medium-term relevance
to the urgent problems of conservation
that we now face. What we do need to do is to
set up some long-term studies of a few carefully
selected species&mdash;preferably long-lived
species&mdash;so that we can trace the effect of environmental
fluctuations on their performance
for a long period.</p>


<h4>Vulnerability of the
Resource</h4>

<p>We already understand a number of factors
that make some of these bird populations particularly
vulnerable to the kind of human activities
which we can envisage in the next
decade or two. Most of the breeding birds concentrate
on islands where they are vulnerable
to predators and to human disturbance. Many
of them concentrate in flocks on human fishing
grounds and over other areas of the continental
shelf which are likely to be the focus of
human activity in the near future. In particular,
some of the birds are known to concentrate
in the passes through the Aleutian Islands,
where they will be particularly vulnerable
to future oil spills. In all these ways the
birds are concentrated in areas where they are
likely to receive disproportionate impacts
from human activity and exploitation.</p>

<p>One point that has been barely mentioned in
this symposium is the effect of molting on the
vulnerability of some of these populations.
The eiders, for example, concentrate on molting
grounds in the Arctic. The exact location
of these molting grounds may not be fully
known, but we certainly know that the birds
molt somewhere in an area where they will be
vulnerable to oil spills (and also to human
hunting if the people who move to the Arctic
choose to hunt them). Nor are eiders the only
species that are flightless when they molt.
Some alcids and loons are also flightless for
short periods and, hence, particularly vulnerable
to oil spills during molt.</p>


<h4>Past Damage to
the Resource</h4>

<p>In the speech opening the symposium, Assistant
Secretary Reed referred to this biological
resource as still relatively unspoiled.
While "relatively" may be an appropriate
word, we do have spectacular evidence of
changes and damage to these bird populations.
The use of the Aleutian Islands for fox
farming seems to me a quite horrifying situation.
We know also that the early whalers and
sealers exploited seabird populations. Although
I know of little specific information
about the effects of such exploitation on birds
in the northern North Pacific, D. G. Ainley in
his survey of historical records from the
Farallon Islands has shown very clearly the
massive effects of human exploitation of
birds, starting early in the 19th century. In
our area of discussion alone, one species (the
spectacled cormorant) is extinct and another
(the short-tailed albatross) became virtually
extinct and is still very rare. I believe that one
or two southern hemisphere species, which
must have been substantial elements in the
northern summer bird population, have also
been seriously depleted as a result of human
activity on their breeding grounds.</p>

<p>Several speakers emphasized the importance
of long-term fluctuations in bird populations
resulting from natural causes, including
some examples from the North Pacific. Other
types of human activity must also have had
some indirect effects on the birds. For
example, whaling and sealing in the 19th century
must have provided large amounts of
food for scavenging birds and eliminated important
competitors for the larger fish-eating
birds. A similar experiment is now in progress
as the predatory fish are being overfished.</p>


<h4>Major Threats to the
Bird Populations</h4>

<p>We now know enough about the distribution
and ecology of the seabirds to identify
the major threats to them that are likely to be
posed by the projected increase in human activity
in the coming decades. The relative importance
of these threats clearly varies from
species to species and from area to area. How<span class="pagenum"><a name="Page_310" id="Page_310">[Pg 310]</a></span>ever,
I think that few of us would disagree
that the largest single threat in the area as a
whole is posed by oil, not only by the prospect
of large-scale drilling for oil on the Alaskan
continental shelf but also by prospective
spills during transportation and deliberate
dumping from ships.</p>

<p>My guess is that the second most important
threat to the seabirds of the northern North
Pacific is the presence of introduced predators,
especially foxes and rats, at the breeding
colonies. Much of the damage inflicted by
these predators may already have been done,
but I think their continuing presence is likely
to have as great a negative effect on the bird
populations as anything else discussed at the
conference.</p>

<p>The relative importance of the other identifiable
threats to the birds is even more conjectural.
Drowning of diving birds in fishnets is
obviously of great potential impact, but its
importance depends greatly on the rapidly
changing practices of fishermen. This problem
must be kept under close surveillance, and the
establishment and enforcement of international
agreements will be critical.</p>

<p>Mineral development has not been mentioned
much. It is my understanding that
there are prospects for substantial onshore,
and perhaps offshore, developments of heavy
metal minerals. These are likely to lead to
local disturbance in the coastal zone, and the
tailings in particular may well pose a threat to
coastal and inshore birds.</p>

<p>Ocean dumping has not been mentioned. I
do not expect that there will be much dumping
of toxic chemicals from Alaskan industries,
but we must remember that this area is
downstream from Japan and the Soviet
Union. I do not know the current practices of
these countries, but the unregulated dumping
of toxic substances from some European
countries apparently has led to large-scale
pollution problems in the North Atlantic.</p>

<p>On present evidence, persistent pesticides
and polychlorinated biphenyls (PCB's) do not
seem to pose a significant threat to north
Pacific seabirds, although high levels of
PCB's have been reported in shearwaters off
the California coast. In my judgment, we have
probably turned the corner in regulating these
chemicals, at least in the northern hemisphere,
and their impact will probably not be
allowed to get worse.</p>

<p>Human disturbance is obviously going to
get very much worse, both from the influx of
new human populations who will be involved
in more industrialized activities in Alaska and
from the likely increase in tourism. A matter
of particular concern is the prospective influx
of natural history tours, which can have
major adverse effects if not carefully
regulated.</p>

<p>Finally, we should not forget the impact of
natural phenomena, including climatic
changes and vulcanism. Bearing in mind the
experience of Katmai, we might expect a natural
disaster to strike a major bird colony at
any moment.</p>


<h4>Practicability of Conservation</h4>

<p>Experience from other countries, as related
in various papers at this conference, has
shown that conservation of seabirds is possible
and practicable, even in remote and inaccessible
areas. We have heard today particularly
about conservation programs and
achievements in Europe and New Zealand.
W. H. Drury spoke briefly about experience in
eastern North America and F. Salomonsen
told us how the bird populations of the
Faeroes Islands have been managed for sustained
yield.</p>

<p>At least in the North Atlantic, where the
history of the bird populations is much better
known, the conservation situation has been,
and probably still is, very much worse than
that now prevailing in the North Pacific.
Looking back on 200 years in the North Atlantic,
we find that two major marine species
have been extinguished, at least one and probably
two or three others became endangered,
and almost all the seabirds were drastically
reduced in numbers (at least in temperate latitudes).
Starting in the late 19th century when
many species first received effective protection,
most showed impressive recoveries, but
some have declined again in the last 30 years.</p>

<p>We can learn several lessons from that experience.
One is that we can do great damage
to seabird populations in a very short time if<span class="pagenum"><a name="Page_311" id="Page_311">[Pg 311]</a></span>
we do things that cause substantial adult
mortality. A second is that seabird populations
can recover well with protection and
modest management&mdash;although most of
them, being slow breeders, recover slowly. A
third lesson is that in the last 30 years we
have caused substantial damage through oil
spills, human disturbance at the breeding
colonies, chemical pollution, and indirectly by
promoting the spread of gulls. Much has been
said at the conference about these present-day
human impacts. However, with the sole
exception of the oil spills which have affected
alcids and sea ducks in parts of northwest
Europe, it seems to me that the damage
caused by human activity in the past 30 years
is considerably less than that in the last 30
years of the 19th century.</p>

<p>Another lesson we can learn from the recent
experience in other areas is that it is possible
to ameliorate some of these adverse human
impacts with local, small-scale, and even
rather amateurish management activities&mdash;for
example, protecting seabird colonies from
gulls, regulating human visits, and controlling
the use of the most toxic chemicals. Our
most conspicuous failure is in controlling oil
pollution. Although safety precautions imposed
on offshore drilling rigs and at shipping
terminals have proved reasonably effective in
averting major damage to seabirds, attempts
to control oil pollution during transportation
have been essentially fruitless. Tanker accidents
and deliberate discharges from vessels
remain the major threat to seabird
populations.</p>

<p>Another lesson from other areas is that public
education has been very effective in putting
pressure behind conservation measures,
and is doing so increasingly. At the same
time, however, it is resulting in an increase of
the disturbances that the birds suffer at their
breeding grounds from casual visitors,
photographers, and sometimes, well-meaning
naturalists.</p>

<p>Finally, in very recent years, there have
been encouraging developments in rehabilitating
oiled birds, captive breeding, and reintroduction
into areas from which they have
been depleted. Restoration of seabird populations
no longer seems an impossible goal.</p>


<h4>Conservation Needs for
North Pacific Seabirds</h4>

<p>We now know enough about the seabirds of
the northern North Pacific to specify in principle
what should be done immediately to conserve
them. I will not address the institutional
arrangements needed for conservation;
R. E. LeResche's paper presented a very clear
picture of the institutional problems involved
in protecting and managing seabirds on an interregional
and international basis. I will
simply endorse his principal recommendation:
that we should try to bring the various responsible
agencies together to formulate comprehensive
management plans.</p>

<p>On the level at which we as individuals and
as a group of biologists can work, we can already
make some positive recommendations.
The most important is that since prevention
of damage is a lot better than cure, measures
to avert damage should have the highest
priority. We have heard a great deal from the
oil industry about the "inevitability" of accidents.
One speaker mentioned the "inherent
fallibility of man." Well, we are all fallible, but
the experience of the last 50 years is that
some people are more fallible than others. No
oil company has a perfect record, but some
have 10 times as many accidents as the best,
and some, I believe, have considerably more
than 10 times as many. This means, very
simply, that it is possible to eliminate most&mdash;not
all, but most&mdash;of the major threats to the
seabirds, merely by upgrading the safety performance
of the entire industry to that already
achieved by its best segments. I suggest
that our major challenge in the coming
years is to work for effective regulation of the
industry: to achieve regulations which will decisively
penalize bad performance and as decisively
reward care.</p>

<p>Perhaps the second priority in conservation
is to protect and manage the existing breeding
colonies. In most cases protection is
legally feasible if we have the will. Most of the
major colonies are in remote areas or in public
ownership where development and disturbance
can be controlled. Management of
the breeding populations is less straightforward,
however, because we do not know<span class="pagenum"><a name="Page_312" id="Page_312">[Pg 312]</a></span>
enough about the functioning of this complex
biological resource. Seabird populations fluctuate
and they have a very long response
time, the environment is not constant, we do
not understand the dynamics of multispecies
communities, and we do not know how they
respond either to external changes or to our
attempts to manage them. Management will
have to be improvisatory for a very long time.
We must recognize that effective conservation
of a bird population with a 20-year
generation time will take at least 20 years to
show results.</p>

<p>Another priority task is to control predators.
I have been impressed by the evidence
we have for major effects of predators on the
seabird populations here. I would regard control
of predators and management of habitats
on some of the major seabird islands as an extremely
urgent task.</p>

<p>A longer-term but no less important program
is public education. This program has
several important aspects: one is to increase
public support for political actions and effective
regulations to protect seabirds; another
is to educate the public about the vulnerability
of seabirds and to prevent disturbance
or deliberate human destruction.</p>

<p>Another aspect of public education is to develop
public interest by making some of the
birds more visible. The great problem with
this biological resource we have been talking
about is that no one knows it is there. Probably
half of us did not know how substantial
and important a resource it is even 5 years
ago. In setting up a large-scale conservation
program, we should not make the mistake of
basing it only on the most remote and inaccessible
colonies, even if these are the most important
numerically. Many of the smaller
colonies are locally very important, both biologically
and for human interest and education.
One example given at this conference
was the State of Washington's program for
conserving what are, by northern Pacific
standards, quite small colonies. This program
is important and impressive because it is conserving
bird populations near people who
want to see the birds. We have the same sort
of situation in Massachusetts and Maine,
where effective protection programs have
been established for extremely small seabird
colonies. We have learned from these programs
that a few hundred birds, or even a few
dozen if properly managed, can be of immense
educational importance. If human access is
carefully managed so that people can see the
birds without disturbing them, these programs
can generate support for conservation
of larger bird populations that may be thousands
of miles away where people may never
see them.</p>


<h4>A Rationale for
Conservation</h4>

<p>As I have tried to show, we know something
about the importance of this biological resource,
and we know in outline what we
should do to conserve it. But why should we?
Almost no one knows the birds are there. We
ourselves do not know whether there are 50
million or 250 million birds in the north
Pacific Ocean. Who cares if 10 million disappear?
If we cannot give a good answer to this
question, we might as well go home and study
chickadees instead.</p>

<p>To justify spending money on conserving
marine birds&mdash;or any other natural resources&mdash;we
must establish their value. Some
of the arguments made in this conference for
assigning economic values to seabirds have
been dangerously weak. The annual value of
"muttonbirds" <em>(Puffinus tenuirostris)</em> in the
New Zealand markets is about $70,000. Some
speakers have tried to argue that seabirds
might play some subtle role that we do not yet
understand in regulating marine communities&mdash;perhaps
they weed out the sick fish. The
direct economic values that we have specified
for seabirds are really not very impressive,
even in terms of the costs involved in conserving
and studying them. The biggest number
we have heard for the value of these seabirds
is the amount of money we are spending on
surveys.</p>

<p>However, this is not the real issue. In judging
the costs and benefits of a conservation
program, we should not look just at the value
of the birds as meat, or oil, or indicators of
pollution. The real issue here, as in all economic
problems, is the rational allocation of
resources. H. Boyd posed the rhetorical question:
"Why should we waste public money on
conserving birds when there are so many<span class="pagenum"><a name="Page_313" id="Page_313">[Pg 313]</a></span>
other things to spend it on?" The question is
more properly posed in reverse: "Why should
the government waste so much public money
on unproductive projects when only a small
amount of money can achieve conservation of
these birds which some people think are
important?"</p>

<p>The fact is that we already know why we
should allocate resources to conservation. I
believe that we have just been evading the
answer. We ought to conserve these birds because
many people want them to be conserved.</p>

<p>This is not, as one speaker said, an elite interest.
The public, as we well know, is already
willing to spend money to conserve natural resources
and is increasingly demonstrating
that willingness. The public, in fact, is ahead
of the administrators and bureaucrats. To appreciate
this, we need only look at some of the
laws already on the books. The Congress of
the United States, in the National Environmental
Policy Act of 1969, declared that it
was the national policy to "create and maintain
conditions under which man and nature
can exist in productive harmony, and fulfill
the social, economic, and other requirements
of present and future generations of Americans."
The Marine Mammal Protection Act of
1972 found that "marine mammals have
proven themselves to be resources of great international
significance, esthetic and recreational
as well as economic, and it is the sense
of Congress that they should be protected and
encouraged to the greatest extent feasible
commensurate with sound policies of resource
management and that the primary object of
their management should be to maintain the
health and stability of the marine ecosystem."
As these laws have been enacted, their
language has become progressively stronger.
The Endangered Species Act of 1973 declares
as the policy of Congress "that all federal departments
and agencies shall seek to conserve
endangered species and threatened species
and shall use their authorities in furtherance
of the purposes of this Act" (P.L. 93-205). It
further directs all Federal departments and
agencies to carry out conservation programs
for the conservation of endangered or
threatened species and to insure that their actions
do not jeopardize the continued
existence of these species or destroy or
modify critical habitat.</p>

<p>These references are not only to Federal
laws passed by remote politicians who can
vote with only a modest responsibility to their
constituents. As we have heard, there are
many State laws and local ordinances which
specify the same kind of thing. All these laws
are on the books for a powerful reason: public
opinion was behind them. The fact that they
have not been enforced and implemented fully
means that we have not been doing our job.</p>

<p>In fact, there is no philosophical problem in
justifying conservation. What we face is an institutional
problem. There is both a public determination
that natural resources should be
conserved and a public apathy and bureaucratic
resistance toward doing it. As concerned
biologists, we should be combating
this apathy by pointing out that conservation
represents a rational allocation of public
resources.</p>

<p>Those who do not learn the lessons of history
are destined to repeat it. If we study the
history of conservation, we find that it developed
most rapidly in those countries which
mismanaged their natural resources earliest.
Within the developed countries there has been
a progressive historical trend toward rational
use and conservation of natural resources.
Conservation of natural resources, in fact,
represents the future and, as biologists, it is
our duty to promote it.</p>


<h4>Economic Feasibility of
Conservation</h4>

<p>Conservation is cheap. Most of us are accustomed
to working on what are essentially
shoestring budgets&mdash;on the order of $100,000,
$10,000, or even $1,000 per year. When we
hear of a million dollars as the cost of doing
something, we tend to think of it as a lot of
money. H. Boyd mentioned a situation on
Baffin Island, where it would cost about a million
dollars to dispose of mine tailings on
shore instead of dumping them into the ocean
under a fulmar colony. I do not think a million
dollars is very much&mdash;certainly not in comparison
with the cost of restoring a colony of
half a million fulmars.</p>

<p>We heard this morning about the acquisition
of Protection Island at a cost of several
hundred thousand dollars. It was pointed out<span class="pagenum"><a name="Page_314" id="Page_314">[Pg 314]</a></span>
that it could have been acquired much more
cheaply only a few years ago. Acquisition of
habitat is cheap if we do it now compared with
what it will cost in a few years or a few
decades. Management is cheap. None of us
gets paid very much, but each of us could
manage several colonies with a couple of students
to help us. Wardens are cheap. Surveys
are cheap. The cost of conserving seabirds is
minuscule in comparison with the amount
spent on the exploitation of resources that
threatens them, and it is minuscule in relation
to the cost of restoring a seabird population
after it has been depleted.</p>

<p>It is far cheaper to avoid oiling birds than it
is to rehabilitate them and to reestablish them
in breeding colonies in the wild. It costs nothing
at all to award leases to companies that
have a good safety record and to refuse leases
to companies with bad records. It costs a little
more to maintain good safety practices in
drilling and transportation. It does cost more
to transport oil in small, double-bottomed
tankers with well-trained crews than to transport
it in big ships flying flags of convenience,
but the cost differential is very small compared
to the value of the shipment.</p>

<p>In considering the economics of conservation,
we have to weigh the costs of conservation
against the value of the resource being
exploited. Full-scale development of oil reserves
on the Alaskan continental shelf would
generate economic values on the order of ten
billion dollars per year. Of this total, 0.1%
would support a reasonably sized management
program for the threatened resources.
About 1% of the total, or 12&cent; per barrel of oil,
would not only support an ample management
program but also permit management of
many other coastal zone resources. Yet the experience
of the last few years has shown that
an increase in oil prices of 1% is barely noticed
by consumers.</p>

<p>The point I am trying to make is that extracting
oil carefully does not cost significantly
more than extracting it carelessly. If
we can solve the institutional problems&mdash;and
I do not underestimate the difficulty of doing
so&mdash;we are not talking about an irrational use
of resources. Conservation is feasible; it is
worthwhile; it is not expensive; and there is a
public demand for it.</p>


<h4>Conclusions</h4>

<p>Practical conservation is an adaptive process.
It is not at present a process that is
firmly based in ecological theory. It is one in
which we have to start by doing something,
see whether it works, and then change our
program in accordance with our early experience.
I do not believe that we can wait for detailed
knowledge of population sizes, or ecology,
or demography, or trophic importance,
or any other biological attribute of these birds
before we start conservation and management.
As scientists we do, of course, find it interesting
and important to study these
things. We should do so; we need to do so; but
we should not use our ignorance of detail as an
excuse for delaying action. If this seabird resource
is worth conserving, we should start
now.</p>


<h4>Summary</h4>

<p>The marine birds of the northern North
Pacific Ocean, Bering Sea, and adjacent seas
constitute one of the great neglected biological
resources of the world.</p>

<p>This resource is impressive in terms of both
total numbers (probably of the order of 100
million birds) and species diversity. A number
of species are endemic to the area and hence of
special interest.</p>

<p>The resource is international in that it includes
major populations of migrants from
Chile, New Zealand, Australia, Japan, the
Soviet Union, and other countries. Several migrant
species appear to use this area exclusively
in their nonbreeding season and should
be included in the list of endemics.</p>

<p>The general relation between the distribution
and abundance of seabirds and other
marine resources is beginning to be understood.
However, comparatively little is known
about the distribution of seabirds in winter,
and there is a serious dearth of information
about breeding success, survival, and
demography.</p>

<p>Seabirds in the north Pacific Ocean and
adjacent seas are concentrated over the continental
shelf and in areas of high biological
productivity. Hence they are especially vul<span class="pagenum"><a name="Page_315" id="Page_315">[Pg 315]</a></span>nerable
to human exploitation.</p>

<p>Seabirds of the northern Pacific Ocean have
already been damaged by human activities in
the past and present. Experience in other
areas shows that seabirds are extremely vulnerable
to human activities and their populations
are often very slow to recover.</p>

<p>The most important threats to the seabird
resource are oil drilling and transportation
and introduced predators, especially foxes.
Other identifiable threats include mineral exploitation,
fishing, ocean dumping of toxic
chemicals, and human disturbance, including
both hunting and tourism.</p>

<p>Experience in other parts of the world, especially
in the North Atlantic, has shown that
seabird populations can be protected and restored
through modest programs of management
and public education. The principal exception
has been the failure to regulate discharges
of oil at sea, which continue to cause
major damage to seabird populations in many
areas.</p>

<p>In the North Pacific and Bering Sea areas,
the most urgent conservation needs are effective
regulation of prospective oil exploitation,
control of introduced predators, and public
education. Regional management plans
should be developed. Public access to bird
colonies should be managed carefully to combine
protection with public education.</p>

<p>Conservation programs for seabirds can be
justified as a response to increasing public demand
for rational management of natural resources.
Conservation programs are inexpensive
in relation to the economic values generated
by oil and mineral development. They
represent a rational allocation of economic
resources.</p>

<p>The following priorities for further study
are suggested:</p>

<p>&#8226; Study of productivity and demography in
a few carefully selected species to provide
basic life table data that will permit rapid
identification of future changes.</p>

<p>&#8226; A base-line census of some carefully selected
breeding colonies, including precise
photographic surveys that can be used to
measure future population changes.</p>

<p>&#8226; Surveys of the distribution of seabirds of
the North Pacific and Bering Sea in winter,
with special emphasis on areas close to shore
where birds may be vulnerable to oil pollution.</p>

<p>&#8226; Special studies of endemic species.</p>

<p>&#8226; Studies of the way in which seabirds locate
and use patchily distributed food resources.</p>

<p>The following conservation measures are
suggested:</p>

<p>&#8226; Adoption of regulations governing exploitation
and transportation of oil which would
provide strong incentives for safe performance
and severe penalties for safety
violations.</p>

<p>&#8226; A conservation tax of a few cents per
barrel of oil to cover the costs of managing the
major seabird colonies and to establish a trust
fund for restoring depleted populations.</p>

<p>&#8226; Equivalent measures for mining and other
exploitative industries in the coastal zone
with a prospective impact on marine
resources.</p>

<p>&#8226; Prohibition of dumping of toxic chemicals
in biologically productive waters.</p>

<p>&#8226; A program to eliminate introduced predators
from the Aleutian Islands and from important
bird colonies elsewhere.</p>

<p>&#8226; Promulgation of effective regulations to
protect birds under the Migratory Bird
Treaty with Japan.</p>

<p>&#8226; Negotiation of migratory bird treaties
with other affected countries, including the
Soviet Union, Australia, New Zealand, and
Chile.</p>

<p>&#8226; Acquisition of major unprotected seabird
colonies into the national wildlife refuge or
other federal landholding systems.</p>

<p>&#8226; Formulation of regional and international
management plans for localized species, especially
endemic species of the Bering Sea.</p>

<p>&#8226; Regulation of public access to major seabird
colonies.</p>

<p><span class="pagenum"><a name="Page_316" id="Page_316">[Pg 316]</a><br /><a name="Page_317" id="Page_317">[Pg 317]</a></span></p>



<p><span class="pagenum"><a name="Page_318" id="Page_318">[Pg 318]</a><br /><a name="Page_319" id="Page_319">[Pg 319]</a></span></p>
<div class="chapter"></div>
<hr class="chap" />




<h2>APPENDIX<br />

Papers and Oral Summaries Presented at the Symposium but Which
Do Not Appear in this Publication</h2>


<div class="hangindent">

<p>Conservation of marine birds of northern
North America&mdash;the symposium theme, by
Nathaniel P. Reed</p>

<p>The marine environment of birds: an overview,
by N. Philip Ashmole</p>

<p>Environmental hazards to birds from petroleum
development in the Beaufort Sea, by
Kees Vermeer [in lieu of a paper on the
status of birds in the Beaufort Sea]</p>

<p>Birds of the marine habitat off northwestern
North America, by Gerald A. Sanger and
James G. King</p>

<p>The pelagic and nearshore birds of the Beaufort
and Chukchi Seas, by George J. Divoky
and George E. Watson</p>

<p>Dispersal and migratory movements, by
George E. Watson, George M. Jonkel, and
F. Graham Cooch</p>

<p>Summary of the session "Status of Marine
Bird Populations," by David N. Nettleship</p>

<p>Breeding habitat of marine birds, by Gerald
A. Bertrand</p>

<p>Exposure of marine birds to environmental
pollutants, by Harry M. Ohlendorf, Robert
W. Risebrough, and Kees Vermeer<a name="FNanchor_83_83" id="FNanchor_83_83"></a><a href="#Footnote_83_83" class="fnanchor">[83]</a></p>

<p>Summary of the session "The Biology and
Ecology of Marine Birds in the North," by
Hugh Boyd</p>

<p>Summary of the session "Conflicts Between
the Conservation of Marine Birds and Uses
of Other Resources," by Joseph P. Linduska</p>

<p>Programs and authorities of the United
States Government related to marine bird
conservation, by Omar Halvorson</p>

<p>Programs and authorities related to marine
bird conservation: Canadian Government,
by Hugh Boyd</p>

<p>Programs and authorities of the State of
Alaska related to marine bird conservation,
by Robert E. LeResche</p>

<p>Seabird programs of private organizations, by
C. Eugene Knoder</p>

<p>Summary of the session "Programs and
Authorities Related to Marine Bird Conservation,"
by William H. Drury</p>

<p>Conservation of marine birds in Antarctica,
by William J. L. Sladen</p>

<p>Seabirds and their conservation in Western
Europe, by W. R. P. Bourne</p>

<p>Summary of the session, "Conservation of
Marine Birds in Other Lands," by Joseph J.
Hickey</p></div>

<div class="footnotes"><h3>FOOTNOTES:</h3>

<div class="footnote">

<p><a name="Footnote_83_83" id="Footnote_83_83"></a><a href="#FNanchor_83_83"><span class="label">[83]</span></a> This paper, because of its length, was published
separately as <cite>Wildlife Research Report 9</cite>.</p></div></div>

<p class="center">U.S. GOVERNMENT PRINTING OFFICE: 1979-681-016/29
</p>

<div id="transnote">

<div class="chapter"></div>
<hr class="chap" />




<h2>TRANSCRIBER'S NOTES</h2>


<p>P. <a href="#Page_228">228</a> the sentence "The factors in Table are largely
self-explanatory" was missing the table reference.</p>

<p>Silently corrected simple spelling, grammar, and typographical errors.</p>

<p>Retained anachronistic and non-standard spellings as printed.</p>
</div>

<div>*** END OF THE PROJECT GUTENBERG EBOOK 49335 ***</div>
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