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+Project Gutenberg (https://www.gutenberg.org) public repository for
+eBook #52382 (https://www.gutenberg.org/ebooks/52382)
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-The Project Gutenberg EBook of Studies of Birds Killed in Nocturnal
-Migration, by Harrison B. Tordoff and Robert M. Mengel
-
-This eBook is for the use of anyone anywhere in the United States and most
-other parts of the world at no cost and with almost no restrictions
-whatsoever. You may copy it, give it away or re-use it under the terms of
-the Project Gutenberg License included with this eBook or online at
-www.gutenberg.org. If you are not located in the United States, you'll have
-to check the laws of the country where you are located before using this ebook.
-
-
-
-Title: Studies of Birds Killed in Nocturnal Migration
-
-Author: Harrison B. Tordoff
- Robert M. Mengel
-
-Release Date: June 20, 2016 [EBook #52382]
-
-Language: English
-
-Character set encoding: ISO-8859-1
-
-*** START OF THIS PROJECT GUTENBERG EBOOK STUDIES OF BIRDS ***
-
-
-
-
-Produced by Judith Wirawan, Chris Curnow, Joseph Cooper
-and the Online Distributed Proofreading Team at
-http://www.pgdp.net
-
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-
-
-
-
- UNIVERSITY OF KANSAS PUBLICATIONS
- MUSEUM OF NATURAL HISTORY
-
-
- Volume 10, No. 1, pp. 1-44, 6 figures in text, 2 tables
-
- September 12, 1956
-
-
- Studies of Birds
- Killed in Nocturnal Migration
-
-
- BY
- HARRISON B. TORDOFF AND ROBERT M. MENGEL
-
-
- UNIVERSITY OF KANSAS
- LAWRENCE
- 1956
-
- * * * * *
-
- UNIVERSITY OF KANSAS PUBLICATIONS, MUSEUM OF NATURAL HISTORY
-
- Editors: E. Raymond Hall, Chairman, A. Byron Leonard,
- Robert W. Wilson
-
-
- Volume 10, No. 1, pp. 1-44, 6 figures in text, 2 tables
- Published September 12, 1956
-
-
- UNIVERSITY OF KANSAS
- Lawrence, Kansas
-
-
- PRINTED BY
- FERD VOILAND, JR., STATE PRINTER
- TOPEKA, KANSAS
- 1956
-
- [Illustration: Logo]
-
- 26-3856
-
-
- * * * * *
-
-Studies of Birds Killed in Nocturnal Migration
-
-BY HARRISON B. TORDOFF AND ROBERT M. MENGEL
-
-
-
-
-Contents
-
-
- PAGE
-
- Introduction 4
-
- Accidents to Migrating Birds in early October, 1954 6
- General 6
- Accidents at Topeka, Kansas 6
- Description of WIBW-TV tower 7
- Weather conditions 7
-
- Acknowledgments 7
-
- Notes on the Species Killed at Topeka 8
-
- Randomness of the Sample 17
-
- Number of Migrants 18
-
- Differential Migration of Sex- and Age-classes 20
- History of the subject 20
- Differential migration of sex- and age-classes as
- shown by the Topeka sample 23
-
- Molt in Relation to Migration 29
- General comment 29
- Molt in the Topeka sample 30
-
- Size Differences according to Sex and Age 31
- Linear measurements 31
- Weights 32
-
- Computations of Longevity and Survival 38
-
- Processing of Samples 38
-
- Summary 39
-
- Literature Cited 41
-
-
-
-
-
-Introduction
-
-
-This paper is primarily an analysis of a sample of migrant birds
-killed in the autumn of 1954 by striking a television tower one mile
-west of Topeka, Shawnee County, Kansas. Secondarily, some aspects of
-migration involved in studies of this kind are discussed and
-historical background is presented.
-
-Considerable interest has been occasioned in recent years in the
-eastern United States by large-scale accidents to night-migrating
-birds. Most accidents have occurred in the autumn. The widespread
-adoption by airports of an instrument called the ceilometer, which
-measures the height of cloud ceilings by reflecting from them a
-high-powered beam of light, has proved under certain conditions to be
-catastrophic to night-flying birds. Among the recent reports of such
-accidents are those of Spofford (1949) and Laskey (1951) for
-Nashville, Tennessee, Howell and Tanner (1951) for Knoxville,
-Tennessee, and Lovell (1952) for Louisville, Kentucky. Recently
-Howell, Laskey, and Tanner (1954) reviewed ceilometer "tragedies"
-without being able to determine the exact reason for their lethal
-effectiveness. Less publicized so far have been mass collisions of
-birds with another class of obstacles, tall radio and television
-towers. These slender towers, usually 500 to 1000 feet tall, are
-increasing rapidly in numbers and there is reason to suppose that they
-will take a correspondingly larger toll of bird life.
-
-Notice has long been given by ornithologists to mass destruction of
-birds by more conventional solid obstructions to passage, and
-newspapers occasionally mention birds killed at such well-known points
-as the Washington Monument and the Empire State Building.
-
-Seventy-five years ago, J. A. Allen (1880) published the results of
-questionnaires circulated by William Brewster to lighthouse keepers.
-Brewster himself (1886) described destruction of birds at a lighthouse
-in the Bay of Fundy, paying keen attention to behavior of the birds
-and the exact conditions under which nocturnal flight and accidents
-occurred. The subject also received attention in several countries
-across the Atlantic. Destruction of birds at Irish lighthouses was
-carefully noted over a period of years and the results were published
-periodically, culminating in R. M. Barrington's massive report (1900)
-which remains in some ways the most thorough of its type.
-
-While conservation-minded individuals have been concerned with the
-tremendous mortality involved in these various events, the ill wind
-blows some good in that, properly used, the data provided by such
-accidents can shed light on many obscure aspects of bird migration.
-Each accidental kill of birds affords a cross-section, approaching in
-variable degree a random sample, of the migrants passing a given point
-on a given date. The types of information provided by such kills are
-numerous, for example: (1) information on the presence of various
-species and the dates of their occurrence; (2) information on the
-relative abundance of species; (3) quantitative data on the relative
-sizes of males and females, and immatures and adults (of importance to
-taxonomic ornithology); (4) information on the relative times of
-migration of males, females, adults, and young; (5) information on
-molts and plumages; (6) quantitative information on composition by
-subspecies of migrants of the same species; (7) physiological data
-(fat condition, _etc._) pertinent to the study of migration; and
-probably others.
-
-In spite of the great potential of this kind of material, the majority
-of ornithologists with access to such data have contented themselves
-with listing the species and sometimes the numbers of birds killed. A
-few have gone further. James T. Tanner (unpublished) attempted to
-compute the longevity of the Ovenbird (_Seiurus aurocapillus_) by
-analysis of ceilometer-killed birds at Knoxville, Tennessee (see
-below). Mention should be made of the reports of Rintoul and Baxter
-(1914) supplemented by Ticehurst (1916) who used rather small numbers
-of birds killed at Scottish lighthouses in studies of molt. However,
-the only effort to utilize the results of accidental kills on a large
-scale over a period of years appears to have been that, already
-mentioned, of Barrington (1900) and his co-workers in Ireland. An idea
-of the potentialities of the large recent kills in the United States
-may be obtained when it is recalled that in the 18 years of
-Barrington's work, which embodied some 1000 reports from lighthouse
-keepers, Barrington obtained for study only about 2000 specimens, many
-of these consisting of wings and feet only (Barrington's paper not
-seen in original; see J. A. Allen, 1901:205). More recently Dobben and
-Bruyns (1939) have analyzed the age and sex classes of some birds
-killed at lighthouses in Holland.
-
-As far as we have learned, there is no previous thorough analysis in
-the literature of large, accidentally-killed samples of birds. On the
-following pages we emphasize some of the uses which can be made of
-such material. We think that intensive analyses of such events,
-whenever they occur, should become a regular part of ornithological
-investigation and that integration of numerous studies of such
-incidents will provide an unprecedented mass of information on
-migration.
-
-
-
-
-Accidents to Migrating Birds in early October, 1954
-
-
-GENERAL.--The few days around the end of the first week of October,
-1954, were notable for a series of accidents which occurred to
-migrating birds over much of eastern United States. So far as we know,
-these were all associated with an extensive belt of bad weather (cold
-fronts and stationary fronts) which covered much of the country during
-that period, and the accidents involved ceilometers and solid
-structures alike. Accidents known to us occurred as far south as
-Macon, Georgia (David W. Johnston, letter: Nov. 1, 1954), as far north
-as New York City, where many migrants were killed at the Empire State
-Building (_New York Times_, Thursday, October 7, 1954, p. 1) and
-elsewhere, and as far west as Smoky Hill Air Force Base at Salina,
-Kansas (ceilometer, October 7, some birds received at the University
-of Kansas). Some of the above, and incidents from a number of other
-localities, were mentioned in varying detail in _Audubon Field Notes_
-(vol. 9, no. 1, pp. 6, 10, 15, 17, 18, 32, February, 1955). Still
-other accidents occurred at Columbia, Missouri (Richard P.
-Grossenheider, verbal communication), and Topeka, Kansas (present
-paper). Some probably have escaped our notice; summaries of some of
-these will probably appear in ornithological journals for some time to
-come. At Robins Air Force Base near Macon, Georgia, at least 50,000
-birds were killed, of which about 2500, representing 54 species, were
-picked up (Johnston, _loc. cit._).
-
-
-ACCIDENTS AT TOPEKA, KANSAS.--At Topeka, Shawnee County, Kansas, all
-birds were killed by collision with the newly-erected (1954)
-television transmitting tower of station WIBW-TV. This tower is one
-mile west of the city.
-
-The first casualties (see Table 1 for all others) were a Sora
-(_Porzana carolina_) and a Yellow-bellied Flycatcher (_Empidonax
-flaviventris_) found on September 7. The major accidents, however,
-occurred on the nights of September 24-25, September 30-October 1,
-October 5-6, and October 6-7. Totals of birds picked up (probably over
-95 per cent of birds killed) are given in Table 1, in which each date
-given is that of the day after the kill, _i. e._, the date on which
-the birds were collected.
-
-All major kills occurred on cloudy and foggy nights associated with
-frontal weather. Throughout the period a few birds struck the tower
-even on fairly clear nights, and minor but appreciable "falls"
-occurred on the nights of October 4-5, 7-8, and 22-23. A few birds
-killed probably were overlooked for a time and found their way into
-later samples. This is especially probable in the case of some birds
-entered under date of October 23, as many of these were somewhat
-desiccated. Weights clearly altered by desiccation or mutilation were
-not recorded. Reports of these accidents have been published by Carson
-(1954 a, b, and c).
-
-According to Carson (1954c:27), the majority of birds killed on nights
-of heavy flight fell "between three and four o'clock in the morning
-when skies were overcast and a cool front moved in from the north. Due
-to the cooperation of the watchmen it is thought that most of the
-birds that were killed were recovered. Of course some injured birds in
-hiding were not found and some were lost to predators."
-
-
-DESCRIPTION OF WIBW-TV TOWER.--The tower is 950 feet tall and stands
-on a hill approximately 1000 feet above sea level. The fact that the
-tower is on a hill places the top of the tower at 1010 feet above the
-elevation of the average local terrain. The tower is triangular in
-cross-section, each face seven feet wide, and is constructed of
-six-inch steel L-beams with three-inch cross-members every seven feet
-and smaller diagonal cross-members. It has no taper and bears a
-transmitting antenna on the top. The tower is supported by 12 guy
-wires, 3 wires attaching at each of 4 levels. The cables extend south,
-WNW, and NNE from the tower and are 1-1/2 inches in diameter. The
-tower is lighted by a series of red lights, some flashing and others
-steady. The transmitter was not in operation when the accidents took
-place.
-
-
-WEATHER CONDITIONS.--All major kills at Topeka occurred when migrating
-birds encountered either a cold front or a stationary front lying over
-eastern Kansas. Typically, this frontal weather included rain, fog,
-and cloud ceilings down to as low as 800 to 1000 feet. Weather of this
-type presumably forces the migrating birds to fly below the cloud
-ceiling and thus brings them within the altitudinal range of the
-television towers.
-
-
-
-
-Acknowledgments
-
-
-We gratefully acknowledge our debt to the Topeka Audubon Society for
-making this study possible by carefully collecting birds killed at the
-television tower. L. B. Carson deserves special mention for his
-general supervision of the bird collecting by the members of the
-Topeka Audubon Society. Members of the Society and others who picked
-up birds under the television tower were: Mrs. Lloyd Biggs, Elaine
-Carson, L. B. Carson, Jesse A. Eddy, Elizabeth Fisher, Mrs. Walter
-Huxman, Florence McKinney, Mrs. Charles Martin, Mrs. Fred P. Martin,
-T. W. Nelson, Fred Prebble, Grace Prebble, Orville Rice, Mrs. G.
-Warren Scholl, E. W. Senne, and Beatrice Swenson.
-
-We received equally important assistance from students and staff of
-the University of Kansas in recording of data and preparation of
-specimens. The following helped in these ways: Rollin H. Baker, R. W.
-Dickerman, David L. Hardy, J. W. Hardy, Jane S. Mengel, Larry D.
-Mosby, Richard Van Gelder, South G. Van Hoose, and Glen E. Woolfenden.
-We are indebted to the Interlibrary Loan Service of the University of
-Kansas Library for help in securing certain reference works. Robert
-Sokal of the University of Kansas gave helpful advice concerning
-statistical procedures.
-
-
-
-
-Notes on the Species Killed at Topeka
-
-
-A list of numbers and kinds of birds killed is given in Table 1.
-Discussion of data afforded by certain species for which, large
-samples were available will be found below. There are additionally
-certain data afforded by the sample and certain comments to be made on
-various species which can be handled most conveniently in an annotated
-list. In this list we have included all weight data (still scarce for
-many North American birds), comments on status in Kansas of various
-species, results of comparisons to determine subspecies, and
-miscellaneous observations. Weights of birds are given in grams and
-were taken on a triple-beam balance. Fat condition is given in the
-scale proposed by McCabe (1943:556). Weight data from birds migrating
-at night should be especially useful because these migrants all have
-relatively empty crops and stomachs, thus reducing variability. Not
-all birds were suitable for weighing and measuring, for a variety of
-reasons. This accounts for discrepancies in totals between Table 1 and
-the annotated list.
-
-All passerine species were aged by noting the degree of ossification
-of the skull. In no case, of the more than a thousand passerines aged
-by examination of the skull, did we find difficulty in determining
-whether an individual was a bird of the year or an adult. We found no
-specimens in which ossification of the skull was nearing completion.
-In the several species in our sample with distinctive first-winter
-plumages, we found complete agreement in age as shown by plumage and
-by condition of the skull. We think this is further proof, if such is
-needed, that this method of aging is thoroughly reliable in early
-autumn for the passerine species included in our sample and for others
-with similar breeding seasons.
-
-TABLE 1.--BIRDS KILLED AT A TELEVISION TOWER AT TOPEKA, KANSAS, IN
-1954
-
-See annotated list for division into sex- and age-classes. Where
-discrepancies exist between totals given here and totals given in the
-annotated list, these result from the fact that some specimens could
-not be sexed and aged.
-
- A: Sept. 25
- B: Oct. 1
- C: Oct. 3
- D: Oct. 4
- E: Oct. 5
- F: Oct. 6
- G: Oct. 7
- H: Oct. 8
- I: Oct. 9
- J: Oct. 10
- K: Oct. 23
- L: Totals
-
- =======================+===+===+===+===+===+===+===+===+===+===+===+====
- | A | B | C | D | E | F | G | H | I | J | K | L
- -----------------------+---+---+---+---+---+---+---+---+---+---+---+----
- Pied-billed Grebe | | 1| | | | 1| | | 1| | | 3
- Green Heron | | | | | | 1| | | | | | 1
- Blue-winged Teal | | 1| | | | 1| 6| | | | | 8
- Virginia Rail | | 3| | | | 1| | | | | | 4
- Sora | 1| 6| | | | 1| 1| 1| | | | 10
- American Coot | | | | | | 3| | | | | | 3
- Mourning Dove | | 8| | | 1| | | | | | 1| 10
- Yellow-billed Cuckoo | | | | | | 1| | | | | | 1
- Black-billed Cuckoo | | | | | | | 1| | | | | 1
- Yellow-shafted Flicker | 3| | | | | | | | | | | 3
- Yellow-bellied | | | | | | | | | | | |
- Flycatcher | | | | | | | 1| | | | | 1
- House Wren | 2| 3| | | | 1| 2| 1| | | 1| 10
- Long-billed Marsh Wren | | 1| | | | 1| 1| | | | | 3
- Short-billed Marsh Wren| 1| 2| | | | | 1| | | | | 4
- Catbird | 1| 28| 1| | 1| 6| 6| | | | | 43
- Brown Thrasher | | 1| | | | 1| | 1| | | | 3
- Wood Thrush | | 3| | | | | | | | | | 3
- Hermit Thrush | | | | | | | | | | 1| | 1
- Olive-backed Thrush | | 14| | 1| | | 1| | | | | 16
- Golden-crowned Kinglet | | | | | | | | | | 1| 5| 6
- Ruby-crowned Kinglet | 2| 1| | | | | 8| 1| 1| | 1| 14
- Yellow-throated Vireo | | 1| | | | | | | | | | 1
- Blue-headed Vireo | 1| 19| | 1| 2| 5| 8| 3| 1| | | 40
- Red-eyed Vireo | 18| 36| | | 2| 13| 2| 3| | | 1| 75
- Philadelphia Vireo | 3| 9| | | | | | | | | | 12
- Warbling Vireo | 8| 19| 1| | 4| 1| 1| | | | | 34
- Black and White Warbler| 1| 1| | | | 3| | | | | | 5
- Tennessee Warbler | | 1| | | 1| 2| 1| | | | | 5
- Orange-crowned Warbler | 7| 14| | | 1| 4| 19| 5| 1| 1| | 52
- Nashville Warbler | 7| 94| 4| | 3| 39| 27| 5| | 1| 1| 181
- Parula Warbler | | | | | | | 1| | 1| | | 2
- Yellow Warbler | 3| 3| | | | 1| 1| | | | | 8
- Magnolia Warbler | | 1| | | | 2| | | | | | 3
- Black-throated Blue | | | | | | | | | | | |
- Warbler | | | | | | | 2| | | | 1| 3
- Myrtle Warbler | | | | | | | | | 1| | | 1
- Black-throated Green | | | | | | | | | | | |
- Warbler | | | | | | | | | | 1| | 1
- Chestnut-sided Warbler | | 1| | | | | | | 1| | | 2
- Bay-breasted Warbler | 1| | | | | 2| | | | | | 3
- Palm Warbler | 3| | | | | | | | | | 1| 4
- Oven-bird | 4| 21| | | | 2| 3| 1| | | 1| 32
- Northern Water-thrush | | 5| | | | | | | | 1| | 6
- Mourning Warbler | 15| 64| | | 2| 11| 2| 1| | | | 95
- Yellow-throat | 10|115| 2| | 4| 25| 18| 1| 1| | | 176
- Yellow-breasted Chat | | 1| | | | | | | | | | 1
- Wilson Warbler | 1| 2| | | | | | | | | | 3
- Canada Warbler | | 2| | | | | | | | | | 2
- American Redstart | 1| | | | | | | | | | | 1
- Bobolink | | 4| | | | | | | | | | 4
- Rose-breasted Grosbeak | | 2| | | | | | | | | | 2
- Indigo Bunting | | 1| | | 2| 3| 1| | | | | 7
- Dickcissel | | 31| | | 1| 3| 1| | | | | 36
- Savannah Sparrow | 1| 6| | 1| | 1| 5| 1| | | 1| 16
- Grasshopper Sparrow | | 7| | | 2| 3| 3| 1| 1| | 1| 18
- Leconte Sparrow | | | | | | | | | | | 3| 3
- Sharp-tailed Sparrow | | | | | | 1| 1| | | 1| | 3
- Slate-colored Junco | | | | | | | | | | | 1| 1
- Clay-colored Sparrow | | 11| 1| | | 2| | 1| | | | 15
- Fox Sparrow | | | | | | | | | | | 1| 1
- Lincoln Sparrow | 41| 7| | | 5| 22| 3| 1| | | 3| 82
- Swamp Sparrow | | 1| | | | 1| 2| | | | | 4
- Song Sparrow | | | | | | | | | | | 2| 2
- Total--species | 22| 41| 6| 3| 13| 31| 29| 16| 10| 8| 15| 61
- Total--individuals | 94|585| 16| 3| 26|146|147| 31| 10| 8| 24|1090
- ------------------------------------------------------------------------
-
-The annotated list may be consulted for further data in connection
-with the species listed in Table 1. As is indicated below, we regard
-the figures of this sample as unreliable to an unknown degree in
-comparing the relative abundance of one species with another.
-Accumulation of such data from various localities, however, should
-prove useful in another type of comparison. Samples of the same
-species killed in the same way at about the same time at different
-localities should be directly comparable. Eventually, this should
-provide us with a means of determining relative abundance of a species
-in different parts of its migratory route.
-
-Approximately 200 of the most interesting specimens were preserved as
-study skins and are in the University of Kansas Museum of Natural
-History. An effort was made to preserve at least one of each species,
-and we fell only a few short of this goal. All of the forms rare in
-Kansas are represented by skins. We could see no reason to list the
-preserved specimens in detail here. Species of which no study skins
-were made, however, are so marked.
-
-So far as we can tell, no truly western subspecies (from west of the
-Great Plains) occurred in the Topeka sample. Probably most or all of
-the birds came from areas more or less directly north of eastern
-Kansas.
-
-In critical areas where different subspecies of the same species occur
-together in migration, data from samples of this kind should prove
-enlightening. In future analyses, conducted in such areas, it might be
-possible to preserve all specimens of some of the variable species, or
-at least to measure all individuals of species in which size is the
-most important variable character. Quantitative study could then be
-made of the different geographic variants occurring, their proportions
-in the migrant population determined, and their origins deduced. In
-studying populations of Painted Buntings (_Passerina ciris_) wintering
-in Mexico, Storer (1951) has provided an interesting demonstration of
-methods which can be applied to such samples.
-
-A few bats killed at the tower provided a surprise. They will be
-discussed separately by Richard Van Gelder.
-
- _Podilymbus p. podiceps._ Pied-billed Grebe.--Weights: male,
- 394.8 (all weights in grams); females, 332.5, 289.7; all
- fat.
-
- _Butorides v. virescens._ Green Heron.--Weight: 1 (unsexed),
- 168.6.
-
- _Anas discors._ Blue-winged Teal.--Weights: 4 males, mean
- 421.2 (391.3-458.1); 3 females, 367.7, 371.6, 393.2; all
- fat.
-
- _Rallus limicola._ Virginia Rail.--Weights: 3 males, 73.7,
- 83.2, 90.5; 1 female, 67.3; moderately fat to fat.
-
- _Porzana carolina._ Sora.--Weights: 4 males, mean 76.8
- (68.7-89.9); 3 females, 62.6, 63.2, 63.5; moderately fat to
- very fat.
-
- _Fulica americana._ American Coot.--Weights: 2 females,
- 385.3, 530.0, both fat. None preserved.
-
- _Zenaidura macroura marginella._ Mourning Dove.--Weights: 2
- adult males, 121.8, 140.2; 3 immature males, 113.1, 126.1,
- 130.0; 3 adult females, 122.5, 126.9, 136.0; 2 immature
- females, 129.4, 132.7; moderately fat to very fat. The
- presence of Mourning Doves in the sample is interesting as
- these birds are not generally regarded as night migrants.
- Conceivably the specimens were local birds going to roost.
- None preserved.
-
- _Colaptes auratus luteus._ Yellow-shafted Flicker.--Weights:
- 2 males, 126.0, 139.4, little fat. Flickers have several
- times been recorded as night migrants.
-
- _Empidonax flaviventris._ Yellow-bellied
- Flycatcher.--Weight: 1 immature male, 11.9, moderately fat.
- This is a rare species in Kansas, the present being the
- ninth preserved specimen for the State.
-
- _Troglodytes aëdon parkmanii._ House Wren.--Weights: 4 adult
- males, mean 10.5 (9.8-10.9), 2 immature males, 9.0, 11.3; 1
- adult female, 9.9, 1 immature female, 7.0; no fat (im.
- [**Female symbol]) to fat.
-
- _Telmatodytes palustris dissaëptus._ Long-billed Marsh
- Wren.--Weights: 1 adult male, 10.8; 1 adult female, 9.2;
- both moderately fat. The specimens are moderately bright and
- rufescent above, being typical of the populations of the
- central plains.
-
- _Cistothorus platensis stellaris._ Short-billed Marsh
- Wren.--Weights: 1 immature male, 8.2; 1 adult female, 8.1;
- immature female, 8.2; all fat.
-
- _Dumetella carolinensis._ Catbird.--Weights: 6 adult males,
- mean 37.5 (34.1-42.5), little fat to very fat; 14 immature
- males, mean 37.57 ± .94 (standard error), S. D. (standard
- deviation) 3.37, little fat to fat; 11 adult females, mean
- 39.09 ± .94, S. D. 2.97, little fat to fat; 12 immature
- females, mean 38.42 ± .83, S. D. 2.74, moderately fat to
- fat.
-
- _Toxostoma r. rufum._ Brown Thrasher.--Weight: 1 immature
- male, 60.2, little fat.
-
- _Hylocichla mustelina._ Wood Thrush.--Weights: 1 adult male,
- 54.2, moderately fat; 2 adult females, 44.6, 45.7, little
- fat and fat, respectively.
-
- _Hylocichla ustulata swainsonii._ Olive-backed
- Thrush.--Weights: 6 immature males, mean 31.0 (28.1-33.2),
- little fat to fat; 6 adult females, mean 29.6 (27.1-35.0),
- moderately fat to fat; 3 immature females, 27.1, 33.8, 35.8,
- little fat to fat. The absence of adult males in our sample
- of 15 birds is noteworthy but inexplicable with our few
- data.
-
- _Regulus s. satrapa._ Golden-crowned Kinglet.--Weights: 1
- adult male, 6.7, moderately fat; 2 immature males, 6.5, 7.4,
- moderately fat and fat; 2 adult females, 7.3, 7.4,
- moderately fat and fat; 1 immature female, 7.2, moderately
- fat.
-
- _Regulus c. calendula._ Ruby-crowned Kinglet.--Weights: 3
- adult males, 6.2, 7.6, 8.2, little fat to fat; 1 immature
- male, 6.6, fat; 4 adult females, mean 6.1 (5.6-6.7),
- moderately fat to fat; 3 immature females, 5.8, 6.6, 7.0,
- moderately fat to fat.
-
- _Vireo flavifrons._ Yellow-throated Vireo.--Weight: 1
- immature male, 21.5, very fat.
-
- _Vireo s. solitarius._ Blue-headed Vireo.--Weights: 9 adult
- males, mean 17.7 (16.6-19.5), little fat to very fat; 17
- immature males, mean 17.53 ± .46, S. D. 1.83, no fat (13.8)
- to very fat (21.3); 7 adult females, mean 17.6 (15.0-21.6),
- moderately fat to very fat; 6 immature females, mean 17.0
- (14.5-18.9), moderately fat to fat. Surprisingly numerous in
- the sample.
-
- _Vireo olivaceus._ Red-eyed Vireo.--Weights: 1 adult male,
- 16.1, moderately fat; 38 immature males, mean 21.21 ± .43,
- S. D. 2.60, little fat (1 specimen) to excessively fat,
- mostly moderately fat or fat; 2 adult females, 18.1, 18.1,
- both fat; 23 immature females, mean 19.28 ± .46, S. D. 2.16,
- little (2 specimens) to very fat, mostly fat.
-
- Wing length: 1 adult male, 79.1; 38 immature males, mean
- 78.05 ± .30, S. D. 1.80; 2 adult females, 76.3, 79.0, 23
- immature females, mean 75.83 ± .42, S. D. 1.99.
-
- As mentioned below, the presence of only 3 adults in the
- sample of 64 Red-eyed Vireos is highly significant and their
- occurrence only in the earlier samples is strong evidence of
- early migration by the adults.
-
- _Vireo philadelphicus._ Philadelphia Vireo.--Weights: 2
- adult males, 12.1, 15.9, moderately fat and very fat; 2
- immature males, 11.1, 13.2, fat and very fat; 2 adult
- females, 13.1, 14.2, both fat; 5 immature females, mean 14.1
- (12.0-15.2), moderately fat to very fat.
-
- This species previously has been collected in Kansas only
- twice. Both records are from Doniphan County in September,
- 1922. Field observers occasionally record the Philadelphia
- Vireo in eastern Kansas. Long (1940:450) calls it a "very
- rare migrant in the extreme east." Our sample of 12 birds
- killed on two nights (and probably after the peak of
- migration of this species) leads us to think that this vireo
- is actually a regular, but overlooked, migrant in fair
- numbers.
-
- _Vireo g. gilvus._ Warbling Vireo.--Weights: 12 adult males,
- mean 15.92 ± .43, S. D. 1.44, moderately fat to very fat; 8
- immature males, mean 16.64 (14.2-17.8), fat to very fat; 5
- adult females, mean 16.1 (13.7-18.0), fat to very fat; 5
- immature females, mean 15.4 (14.1-17.8), little fat to fat.
-
- Wing length: 12 adult males, mean 73.08 ± .49, S. D. 1.64; 8
- immature males, mean 71.15 (69.9-72.8); 5 adult females,
- mean 70.0 (69.2-71.0); 5 immature females, mean 68.4
- (67.7-70.3).
-
- Tail length: 12 adult males, mean 53.33 ± .53, S. D. 1.77; 8
- immature males, mean 50.03 (47.1-51.3); 4 adult females,
- mean 48.6 (47.7-49.8); 5 immature females, mean 49.2
- (47.3-53.0).
-
- There is no indication that western birds (_V. g.
- swainsonii_) make up any part of this sample.
-
- The sample of 34 Warbling Vireos is too small to show the
- significance, if any, of the 2:1 ratio of males to females
- in the sample. Adequate samples of this species, taken at
- intervals, would add interesting information on time of
- migration of the four sex- and age-classes.
-
- _Mniotilta varia._ Black and White Warbler.--Weights: 1
- adult male, 12.5, fat; 2 adult females, 10.0, 10.0, little
- fat, fat.
-
- _Vermivora peregrina._ Tennessee Warbler.--Weights: 1 adult
- male, 10.9, very fat; 1 immature male, 12.9, very fat; 2
- adult females, 9.1, 12.5, moderately fat and very fat. The
- relative scarcity of Tennessee Warblers in the sample is
- surprising. They are common in the area in spring.
-
- _Vermivora c. celata._ Orange-crowned Warbler.--Weights: 9
- adult males, mean 8.8 (7.7-10.9), little fat to fat; 13
- immature males, mean 8.92 ± .15, S. D. .53, little fat to
- fat; 5 adult females, mean 8.8 (8.3-10.3), little fat to
- moderately fat; 17 immature females, mean 9.13 ± .08, S. D.
- .72, little fat to fat. Of the 19 Orange-crowned Warblers
- killed on October 7, 11 had little fat, 6 were moderately
- fat, and only 2 were fat. No one-night sample of any other
- warbler killed at Topeka had less fat than this group of
- warblers. Furthermore, our sample (including 11 males) from
- October 7 (all sex- and age-classes) averaged 8.81 grams;
- the sample of 13 (including only 4 males) from October 1
- averaged 9.1 grams. If one can assume, for any one species,
- that individuals undertake nocturnal migration only when
- they are physiologically ready, and this includes a certain
- amount of fat as a fuel source (Wolfson, 1954), then this
- further assumption seems justified: birds killed in
- migration with little fat must have flown longer or farther
- or both than birds killed with more fat. No further
- speculation on this point is permissible with our data, but
- the possibilities for study of future large kills,
- especially where actual time of death of the birds is known,
- are obvious.
-
- _Vermivora r. ruficapilla._ Nashville Warbler.--More
- Nashville Warblers were picked up at Topeka than any other
- species and they are discussed in detail elsewhere in this
- report. The four sex- and age-classes can be identified with
- fair accuracy on plumage characteristics alone. Adult males
- have a large amount of reddish-brown in the crown, not
- completely veiled by the gray tips of the crown feathers.
- Immature males have a smaller but distinct crown patch,
- usually completely veiled. All males, compared with females,
- are grayer on the sides of the head, have a more nearly
- white eye-ring, and show clearer yellow on the throat. Adult
- females differ from immature females in that they more often
- have a trace of rufous in the crown and tend to be brighter
- below than the immatures. Of 177 specimens, 20 were very
- fat, 108 were fat, 46 were moderately fat, and 3 had little
- fat.
-
- _Parula americana._ Parula Warbler.--Weight: 1 adult female,
- 7.9, fat.
-
- _Dendroica petechia aestiva._ Yellow Warbler.--Weights: 1
- immature male, 10.2, fat; 3 adult females, 8.8, 9.5, 10.1,
- moderately fat; 2 immature females, 9.0, 9.4, little fat and
- fat.
-
- _Dendroica magnolia._ Magnolia Warbler.--Weights: 1 adult
- female, 9.0, moderately fat; 2 immature females, 7.9, 10.3,
- moderately fat and fat.
-
- _Dendroica c. caerulescens._ Black-throated Blue
- Warbler.--Weights: 2 immature males, 13.8, 14.1, excessively
- fat; 1 immature female, 11.4, fat. This species is rare in
- Kansas. Although its breeding range is almost entirely east
- and north of Kansas, records in files at the University of
- Kansas show that more specimens have been taken in western
- than in eastern Kansas.
-
- _Dendroica c. coronata._ Myrtle Warbler.--Weight: 1 immature
- female, 11.6, fat.
-
- _Dendroica pensylvanica._ Chestnut-sided Warbler.--Weights:
- 2 immature females, 8.1, 10.0, little fat. Only one specimen
- from Kansas had been preserved previously although the
- species is a regular transient in small numbers throughout
- the state.
-
- _Dendroica castanea._ Bay-breasted Warbler.--Weights: 1
- adult male, 19.2, excessively fat; 1 adult female, 11.7,
- little fat; 1 immature female, 11.2, moderately fat. Only 5
- specimens of this warbler have been taken previously in
- Kansas, 4 in spring (Ruth, 1952:18-19) and 1 in fall.
-
- _Dendroica p. palmarum._ Palm Warbler.--Weights: 2 immature
- males, 9.9, 10.9, moderately fat; 2 unsexed immatures, 9.1,
- 9.4, moderately fat. This species has been taken in fall in
- Kansas only once before (KU 26353, taken by Wetmore, at
- Lawrence, on October 5, 1907), but probably occurs regularly
- in both spring and fall migration.
-
- _Seiurus a. aurocapillus._ Oven-bird.--Weights: 2 adult
- males, 22.5, 23.8, fat and very fat; 14 immature males, mean
- 21.89 ± .66, S. D. 2.46, fat to very fat; 8 adult females,
- mean 21.4 (18.3-25.7), moderately fat to fat; 6 immature
- females, mean 18.2 (15.6-20.0), moderately fat to fat.
-
- _Seiurus noveboracensis notabilis._ Northern
- Water-thrush.--Weights: 3 immature males, 18.1, 18.6, 22.2,
- moderately fat to fat; 1 immature female, 22.2, fat.
- Referring these birds to _notabilis_ is a somewhat arbitrary
- procedure. They display some intermediacy of characters and
- probably stem from a population, intermediate between
- _notabilis_ and _noveboracensis_, occupying much of central
- North America (cf. McCabe and Miller, 1933).
-
- _Oporornis philadelphia._ Mourning Warbler.--Weight data
- presented elsewhere. The birds killed at Topeka provide the
- latest fall dates for this species in Kansas. Fifteen were
- killed on September 25, 64 on October 1, 2 on October 5, 11
- on October 6, 2 on October 7, and 1 on October 8. We find no
- other records later than September 15. Of 93 specimens
- examined, 1 was excessively fat, 22 were very fat, 45 were
- fat, 21 were moderately fat, and 4 had little fat. The
- abundance of this secretive species in the sample was a
- great surprise. It had previously been considered a rather
- rare migrant in this area.
-
- _Geothlypis trichas occidentalis [>brachidactyla?]._
- Yellow-throat.--Weight data presented elsewhere. This
- species was second in numbers only to the Nashville Warbler
- in the total kill at Topeka. Of 167 birds examined, 29 were
- very fat, 114 were fat, 23 were moderately fat, and 1 had
- little fat.
-
- The Yellow-throats are greatly in need of meaningful and
- comprehensive revision, which to date has been restricted to
- the western subspecies (Behle, 1950). Since the appearance
- of the 1931 A. O. U. Check-List a great deal of scattered
- taxonomic work on the species, as yet unsynthesized, has
- made the picture of its geographic variation a blurry one so
- far as the details are concerned. Made in the absence of
- adequate comparative material, the above identification is
- to be regarded as tentative. Also, it is, unfortunately,
- based only on those 6 of our 176 specimens preserved as
- skins. Five of these are adult males, the sixth being an
- immature female. Compared with a series of Kentucky
- specimens regarded as typical _brachidactyla_, these birds
- are paler and brighter above (tending toward gray-green
- rather than brownish olive), brighter and more extensively
- yellow below, with broader, more nearly white superciliary
- stripes above their black masks (in males). In size they are
- close to _occidentalis_ (see Behle, 1950:202). Five males
- have an average wing-length of 56.6 mm. (53-59); one female
- measures 53. Six males from Kentucky: 55.1 (53-56); four
- females, 51.1 (48-56). Our birds may be assumed to have
- stemmed from a population to the north and west which, if
- not _occidentalis_ (or _campicola_ Behle and Aldrich, of
- which no comparative material is at hand), is intermediate
- between _brachidactyla_ and more western birds. Judging from
- Behle's map (1950:fig. 32), these birds may have come from
- an area near the confluence of three subspecies
- (_campicola_, _occidentalis_, _brachidactyla_). Long
- (1940:452) reports three subspecies breeding in Kansas
- (_brachidactyla_, northeast; _occidentalis_, west;
- _trichas_, southeast). The occurrence in Kansas of _G. t.
- trichas_ as currently understood is completely out of the
- question.
-
- _Icteria v. virens._ Yellow-breasted Chat.--Weight: 1
- unsexed immature, 29.7, moderately fat.
-
- _Wilsonia p. pusilla._ Wilson Warbler.--Weights: 2 adult
- females, 7.5, 7.8, fat, moderately fat; 1 unsexed adult,
- 8.3, fat.
-
- _Wilsonia canadensis._ Canada Warbler.--Weight: 1 immature
- female, 10.0, little fat. We know of only five other
- specimens from Kansas, although this warbler seems to be a
- regular migrant in small numbers in the state.
-
- _Setophaga r. ruticilla._ American Redstart.--Weight: 1
- immature female, 9.1, moderately fat.
-
- _Dolichonyx oryzivorus._ Bobolink.--Weights: 2 adult
- females, 39.5, 42.9; 2 immature females, 38.8, 42.0; all
- excessively fat. Specimens of the Bobolink previously have
- been taken in fall in Kansas only on September 20 and 24,
- 1933, near Lawrence, by Long and Preble (Long, 1934).
-
- _Pheucticus ludovicianus._ Rose-breasted Grosbeak.--Weights:
- 1 adult male, 50.4, fat; one immature male, 54.5, very fat.
-
- _Passerina cyanea._ Indigo Bunting.--Weights: 1 adult male,
- 18.4, fat; 2 immature males, 17.2, 17.2, fat and very fat; 2
- adult females, 14.3, 16.9, moderately fat and very fat; 1
- immature female, 13.4, little fat. The sample was carefully
- checked for Lazuli Buntings (_Passerina amoena_); none was
- found.
-
- _Spiza americana._ Dickcissel.--Weight data presented
- elsewhere in this paper. Dickcissels were picked up at the
- television tower on October 1 (31), 5 (1), 6 (3), and 7 (1).
- These birds, together with an adult female taken 3 miles
- east and 3 miles south of Lawrence, on October 11, 1953, by
- Tordoff, are the only specimens of this species taken as
- late as October in Kansas. The Dickcissel becomes
- inconspicuous in late summer and many observers here and
- elsewhere have thought the species disappeared much earlier
- than it really does (see Ganier, 1949). Of 34 specimens, 20
- were very fat and 14 were fat.
-
- _Passerculus sandwichensis nevadensis._ Savannah
- Sparrow.--Weights: 1 adult male, 19.4, fat; 2 immature
- males, 18.3, 19.0, moderately fat; 5 adult females, mean
- 17.2 (14.8-19.5), little fat to fat; 4 immature females,
- mean 18.0 (16.9-19.6), moderately fat to fat. Many of the
- Savannah Sparrows migrating through Kansas have in the past
- been referred to the subspecies _P. s. anthinus_ (=
- _alaudinus_ of the 1931 A. O. U. Check-List) by various
- workers (see Long, 1940:454). As Peters and Griscom
- (1938:464-5) have shown, true _anthinus_, breeding in the
- far northwest, ordinarily occurs in migration only in the
- western part of the country, the breeding Savannah Sparrows
- of a large part of the central continental region (east to
- southern Wisconsin) being _P. s. nevadensis_ as now
- understood. Migrants of this pale, clay-colored subspecies
- should be abundant in Kansas, and all of the specimens in
- the present sample are referable to it.
-
- _Ammodramus savannarum perpallidus._ Grasshopper
- Sparrow.--Weights: 3 adult males, 16.4, 17.6, 20.6,
- moderately fat, fat, fat; 5 immature males, mean 18.1
- (16.0-20.2), little fat to fat; 5 adult females, mean 17.9
- (16.8-18.9), moderately fat to very fat; 5 immature females,
- mean 18.1 (16.8-20.6), fat to very fat.
-
- _Passerherbulus caudacutus._ Leconte Sparrow.--Weights: 1
- immature male, 11.2, moderately fat; 1 immature female,
- 12.2, moderately fat.
-
- _Ammospiza caudacuta nelsoni._ Sharp-tailed
- Sparrow.--Weights: 2 adult males, 15.2, 17.1, moderately fat
- and very fat; 1 adult female, 13.3, little fat. Five
- specimens of this species have been taken previously in
- Kansas, all in October in the eastern part of the state.
- Additionally, several observers have reported birds seen but
- not collected. The three birds from Topeka were picked up on
- October 6, 7, and 10 and are the only specimens taken since
- 1907. Possibly our specimens from Topeka struck the tower on
- the same night. Tordoff noticed, upon preparation, that the
- specimens from October 7 and 10 showed progressive drying of
- the extremities and spoilage as compared with the bird
- picked up on October 6.
-
- _Junco hyemalis cismontanus._ Slate-colored Junco.--Weight:
- 1 immature female, 16.4, little fat. Juncos of hybrid type,
- whether _J. h. hyemalis_ × _J. oreganus_ subsp. or true _J.
- h. cismontanus_, are fairly common in eastern Kansas.
-
- _Spizella pallida._ Clay-colored Sparrow.--Weights: 2 adult
- males, 11.6, 12.2, both fat; 1 immature male, 11.8, fat; 1
- adult female, 12.5, fat; 7 immature females, mean 11.1
- (9.7-12.5), little fat to fat.
-
- _Passerella iliaca iliaca._ Fox Sparrow.--Weight: 1 adult
- female, 29.4, little fat. A trifle grayer above than any of
- several Kentucky specimens, this bird nevertheless seems
- well within the range of variation of _iliaca_.
-
- _Melospiza l. lincolnii._ Lincoln Sparrow.--Weights and
- measurements are discussed elsewhere. Of 81 specimens, 15
- were very fat, 47 were fat, 12 were moderately fat, and 7
- had little fat. Interestingly, there is no evidence that the
- large southern montane subspecies (_M. l. alticola_) has
- contributed to the present sample. No bimodality is evident
- in the curve of wing-length in our birds, the largest of
- which barely approach, the small extreme recorded for
- _alticola_ by Miller and McCabe (1935:156).
-
- _Melospiza georgiana ericrypta._ Swamp Sparrow.--Weights: 3
- immature females, 14.3, little fat, 16.7, 17.0, moderately
- fat. Swamp Sparrows examined were all more or less brightly
- colored and seem to belong to this northern subspecies.
-
- _Melospiza melodia juddi._ Song Sparrow.--Weights: 1 adult
- female, 19.4, little fat; 1 unsexed immature, 16.0, little
- fat. A large proportion of the migrant and wintering Song
- Sparrows in eastern Kansas probably originate from the range
- of this subspecies in the northern plains. _Melospiza
- melodia euphonia._--One immature female (not weighed) was
- picked up below the tower on October 27, 1954, and thus does
- not appear in Table 1. The specimen proved typical of this
- generally more eastern subspecies upon comparison with a
- large series from Kentucky. For what it may be worth we
- refer the single specimen to this subspecies. Long
- (1940:456) reported two eastern subspecies from Kansas
- ("_beata_," _melodia_). All Kansas specimens genuinely of
- eastern origin probably originate from the range of
- _euphonia_, as now understood.
-
-
-
-
-Randomness of the Sample
-
-
-The reliability of certain of the conclusions which might be drawn
-from data of the kind presented herein depends largely on the
-randomness of the sample. To what degree does this sample provide a
-true cross-section of the nocturnal migrants present over the area on
-a given night or succession of nights? As far as the relative
-abundance of species in the sample is concerned, there is little doubt
-that it is not at all random. The absence of such species as the
-Gray-cheeked Thrush (_Hylocichla minima_), among the passerines, and
-many of the shorebirds known to be migrating through the area at the
-time is evidence for this statement. Quite possibly many seminocturnal
-species did not strike the tower at all for the simple reason that
-they could see it, and certain large-eyed diurnal species (such as
-thrushes and shorebirds) may avoid collision to some extent, thus not
-appearing in the sample in proportion to their actual numbers.
-Finally, some or all of the species concerned probably migrate partly
-by day. The sample may to some degree reflect the true relative
-abundance of closely related species. For example, there is little
-doubt that, as shown by the sample, Nashville Warblers are more
-numerous locally at this season than Tennessee Warblers, a fact that
-can readily be corroborated by ordinary field observation. Also, the
-sample is useful in suggesting the actual abundance of species which
-are furtive and/or difficult to identify under normal field
-conditions, for example, the Mourning Warbler and Philadelphia Vireo.
-It is obvious that the sample should reflect the true relative
-abundance at one place and time of any two species with equal tendency
-to migrate by night and equal tendency to strike the tower. Since the
-facts in regard to both tendencies are at present unknown for most
-species, we think that interspecific comparisons should be avoided or
-approached with extreme caution.
-
-In respect to the relative abundance of the various sex- and
-age-classes within a given species, the sample is, we think, as close
-to random as is possible to obtain. Certainly it is greatly superior
-to samples obtained by field collecting, where possible differences in
-habits, wariness, and experience of the birds, and subconscious (if
-not conscious) selection by collectors can all bias the results.
-Dwight (1900:128-9) believed that the greater wariness of adult birds
-was almost entirely responsible for the seemingly disproportionate
-number of immatures in autumn and gave some observational evidence in
-favor of his views. The large percentage of adults in some of the
-samples here treated tends to reinforce Dwight's position. To a
-somewhat lesser extent, this advantage in randomness of accidental
-kills over routine collecting may be supposed to apply also in
-demonstrating the composition by subspecies of a single migrant
-species.
-
-So far as particulars already mentioned are concerned, the present
-sample or other samples of tower-killed birds would seem to be in no
-way superior (that is, more nearly random) to samples obtained in
-connection with lighthouses and other lighted objects, and
-ceilometers. In one important respect, however, it is probably
-somewhat superior to these as the dimly red-lighted structure has not
-been shown to have any important collecting or attracting influence.
-Thus, in computations intended to estimate the over-all abundance of
-migrants, the sample should be more reliable than samples involving
-bright light with its possible attracting effect.
-
-
-
-
-Number of Migrants
-
-
-If it can be assumed that nocturnally migrating birds are
-approximately uniformly spaced across the sky and that the red lights
-did not attract birds which would otherwise have missed the tower, it
-is possible to compute the volume of migration from the sample killed.
-In regard to the first assumption, both Stone (1906:250-251) and
-Lowery (1951:409-413) have presented evidence of fairly uniform
-distribution of nocturnal migrants. We have no information on the
-second assumption beyond the facts that birds do not strike the high
-towers on clear nights or lower towers even on stormy nights.
-
-On nights when large numbers of birds struck the 950 foot Topeka
-tower, only a few struck a 500 foot radio tower, also lighted with red
-lights, at Lawrence, 24 miles east, under similar weather conditions.
-Most of the birds found at Topeka were fairly close to the base of the
-tower, indicating that they struck the tower itself or that they were
-flying high enough to strike guy wires only fairly close to the tower.
-The scarcity of birds under the guy wires some distance from the tower
-at Topeka and at the radio tower at Lawrence causes us to think that
-most of the birds were flying more than 450 feet above the ground. On
-this basis, we have computed numbers of migrants passing through a
-plane one mile long and 500 feet high (2,640,000 square feet),
-intersecting the assumed path of migration at right angles.
-Vertically, the theoretical plane begins at 450 feet above ground and
-has its top edge at 950 feet. The solid (discounting spaces between
-girders, _etc._) cross-sectional area of the tower intersecting this
-plane was computed by actual measurement to be 1685 square feet. On
-the night of September 30-October 1, 585 birds were killed. By
-computation (585/1685 = X/2,640,000), approximately 916,000 birds
-passed through the mile-long plane that night. On each of the nights
-of October 5-6 and October 6-7, approximately 230,000 birds passed
-through this plane. By comparison, Lowery (1951:436) recorded maximum
-station densities in one night in spring of 63,600 birds at Tampico,
-Mexico, and 51,600 at Lawrence, Kansas, as determined by
-moon-watching. Lowery's figures refer to numbers of birds crossing any
-part of a circle one mile in diameter and are roughly comparable to
-ours if, as we think, most of the birds at Topeka were flying at
-altitudes between 450 and 950 feet above the ground.
-
-It must be realized that these figures are only approximations. One
-variable ignored is the frontal extent (or area, viewed from the
-front, subject to damage by striking an obstruction) of the birds
-themselves. Since practically all birds killed showed head or trunk
-injuries, rather than a high proportion with only broken wings, we
-chose to disregard frontal extent of the birds in our calculations. If
-our figures are inaccurate by as much as 50 per cent in either
-direction, which seems unlikely to us, they still give some idea of
-the tremendous volume of nocturnal migration under some conditions.
-
-It may be more meaningful to compute numbers of migrants by species.
-This can be done readily by making appropriate substitutions from
-Table 1 in the equation given above. For example, on the night of
-September 30-October 1, approximately 147,000 Nashville Warblers
-passed through the mile-long plane and on the same night, 100,000
-Mourning Warblers and 14,000 Philadelphia Vireos. Neither of the last
-two species would be judged to be abundant migrants in autumn in
-eastern Kansas by ordinary field observations; the television tower
-sample, however, indicates that these as well as other species must
-often be overlooked when they do stop in Kansas.
-
-
-
-
-Differential Migration of Sex- and Age-classes
-
-
-HISTORY OF THE SUBJECT.--For a long time it has been known in a
-general way that old and young birds and males and females of some
-species do not always migrate at the same times, by the same routes,
-or even to the same places. This is a subject about which much has
-been written. Reading the summaries of some general texts, it is easy
-to acquire the impression that the facts of the matter are well known.
-On the contrary, they are poorly known and much remains to be learned
-before differential migration is understood. This can best be
-indicated by a brief survey of the literature.
-
-The importance of the subject was emphasized by Meinertzhagen
-(1930:52) in one of the later reviews of differential migration: "The
-main problem concerns the Cause of Migration, the Stimulus which
-compels Migration and the Origin of the Migratory Habit.... There is,
-however, a minor problem which affords valuable evidence in helping us
-to solve the major problem, bearing very directly on it, namely, the
-order of sex and age on migration."
-
-The mystery of how birds, especially the young, find their way in
-migration has fascinated students since the earliest times. The quite
-natural though purely anthropomorphic conclusion of early scholars was
-that the old birds led the young on migration. This attractive idea
-persisted long after ornithology began to grow into a science. The
-classic theory was restated by Palmén (1876:267), in one of the first
-thorough reviews of the subject of migration, as follows: "Directe
-Beobachtungen in der Natur ergeben, dass die Schaaren von ziehenden
-Vögeln allgemein ältere und stärkere Individuen als Anführer des Zuges
-haben." Variously modified, this view continued to crop up for some
-time and still found support in the 1890's (see Dixon, 1892:69). Gätke
-(1895:101) correctly questioned the credibility of Palmén's "direct
-observations."
-
-With the gradual abandonment of the unsupportable classic theory,
-diametrically opposed views were adopted by workers on opposite sides
-of the Atlantic. The American stand was ably expressed by Brewster
-(1886), who went to great pains to state his case and give evidence,
-and who was later supported by Allen (1896:144-147; 1909:17). The
-Americans held that adult birds nearly always preceded the young in
-migration, and this was based on much evidence, whether or not
-correctly interpreted. Dwight (1900:127) also gave evidence in favor
-of this theory. Equally definite, if, as has later been shown,
-somewhat vaguely documented, was the famous work of Gätke (1895:see
-pp. 100-113), who after many years' observation of migrant birds in
-Heligoland concluded the exact opposite, that young in general precede
-adults (see critiques of Allen, 1896:144-147; Wiegold, 1926:5).
-Gätke's dissenting opinion was for a time supported enthusiastically
-by British workers (Gurney, 1923:579-580).
-
-As so often happens, neither extreme has withstood the test of time,
-and more recent summaries (Meinertzhagen, 1930:55-56; Thomson, 1926,
-1936:488-489; Wiegold, 1926) have tended to compromise. Many
-exceptions to Gätke's extreme conclusion have been detected.
-Exceptions to the Brewster-Allen stand have also been discovered,
-although work along these lines on the American side has lagged
-somewhat. Rowan (1926) has given further evidence on the migration of
-certain shorebirds, and some evidence has accrued in relation to
-particular species and groups as a result of life-history and banding
-studies (see Pitelka, 1946). Authors of major works on migration,
-however, have either been preoccupied with other phases of migration
-or avoided the issue. In an able study (one of several on related
-subjects) of the composition by sex and age of migrant populations in
-north Germany, Drost (1935:177) did not go into the question of order
-on migration.
-
-One is left with the impression that the whole subject is still
-unsettled. While earlier workers sought to reduce the entire matter to
-law, the latest disclaim the possibility of generalization. After
-summarizing Brewster's and Gätke's opinions, Thomson (1926:79) wrote:
-"It would seem, in any event, that no general rule can be laid down."
-Meinertzhagen's summation (1930:56-57) still represents fairly well
-the status of our knowledge: "Order of sex and age on autumn passage
-is very difficult to arrive at, as evidence is conflicting. But, on
-the whole, it seems that birds flock together, old and young,
-preparatory to moving south, and do in many cases initiate migration
-in company.... But once movement is initiated, among birds which do
-not habitually fraternise in flocks, adults, and especially males,
-will naturally outstrip the less virile females and still less virile
-offspring.... The consequence is that any observer at an intermediate
-station such as Heligoland is, in noting birds of the year as first
-arrivals, has not had an opportunity of noting the flocks of adults
-which have passed without alighting. On the other hand, there is very
-definite evidence to show that among certain species, adults follow
-their offspring on migration. The reason for different behaviour
-among different types of birds remains obscure." We regard much of
-this as still theory.
-
-[Illustration: FIG. 1. Composition by age and sex as found in one or
-more series of each of eight species of birds included in the Topeka
-sample. Each separate series is represented by a single histogram, the
-histograms for a species being grouped with the earliest series on the
-left. Each histogram expresses the numbers of adults (left-hand
-column) and immatures (right-hand column) in terms of percentage of
-the whole series. Thus the two bars of each couplet add up to 100 per
-cent. The hatched portion of each bar represents males, the clear
-portion females.]
-
-It would be difficult to imagine a better way of resolving the
-problems and uncertainties just reviewed than by the detailed
-analysis of large samples of migratory birds killed at random at
-various points and times. An analysis of the sample of birds
-accidentally killed at Topeka is presented here as an initial step in
-this direction. Although the conclusions based on this sample are
-tentative and may in time be altered, the data themselves are
-definite. If this general type of analysis is repeated again and
-again--applied to samples taken on many dates and in many
-localities--a mass of hitherto unparalleled evidence for the study of
-migration will emerge.
-
-
-DIFFERENTIAL MIGRATION OF SEX- AND AGE-CLASSES AS SHOWN BY THE TOPEKA
-SAMPLE.--Smaller samples have not been treated. Species affording
-samples seemingly large enough to justify at least preliminary
-analysis were: Catbird, Red-eyed Vireo, Mourning Warbler, Dickcissel,
-Nashville Warbler, Orange-crowned Warbler, Yellow-throat, and Lincoln
-Sparrow (Fig. 1). For all of these except the Catbird and Dickcissel,
-at least two samples from a week or more apart were available for
-comparison in an effort to detect trends in migration. Fig. 1 shows
-the actual ratios of sex- and age-classes observed in samples of the
-species listed above. Each of the last four species provided two
-separate samples, of sufficient size to warrant an attempt at
-measuring the statistical significance of the observed changes in
-adult-immature ratios (Table 2).
-
-TABLE 2.--STATISTICS OF THE RATIOS OF ADULTS TO IMMATURES IN FOUR
-SPECIES
-
- =======================================================================
- Species | Dates |Total | Number | Differ- |P[3]
- | of samples |number| and | ence |
- | | |percentage |(in %)[2]|
- | | |of adults[1]| |
- --------------|--------------------|------|------------|---------|----
- Nashville |Oct. 1 (93) | 156 | 45 (.484) | .071 |.36
- Warbler |Oct. 5-7 (63) | | 26 (.413) | |
- | | | | |
- Orange-crowned|Sept. 25-Oct. 1 (19)| 44 | 3 (.158) | .282 |.05
- Warbler |Oct. 5-9 (25) | | 11 (.440) | |
- | | | | |
- Yellow-throat |Oct. 1 (115) | 159 | 62 (.540) | .085 |.34
- |Oct. 5-8 (44) | | 20 (.455) | |
- | | | | |
- Lincoln |Oct. 1-3 (44) | 71 | 27 (.614) | .318 |.01
- Sparrow |Oct. 6-10 (27) | | 8 (.296) | |
- ----------------------------------------------------------------------
-
- Footnotes:
-
- [1] Percentage of immatures equals 1.000 minus percentage of
- adults.
-
- [2] Standard error of the difference between ratios was
- computed by the formula
- [Greek: s]^e = sqrt(P_e Q_e (1/N_1 + 1/N_2))
- where P_e equals percentage of adults and Q_e equals
- percentage of immatures in the entire sample.
-
- [3] Probability of error; _i. e._, a P of .01 means there is
- one chance in 100 that the difference observed does not
- represent an actual difference in nature.
-
-Upon the application of statistical methods it soon became evident
-that, unless changes in ratio between two samples are marked, large
-samples would be required in order to reach conclusions of high
-statistical significance in a single study of the present type. In
-this case (see Table 2), the Lincoln Sparrow and Orange-crowned
-Warbler, though represented by only moderate-sized series, show marked
-changes in age composition over the period studied, and the
-statistical treatment indicates a high degree of probability that
-these changes are real. Assurance that the lesser changes observed in
-the Nashville Warbler and Yellow-throat are real, on the other hand,
-is much less, even though the samples are larger. Few if any of the
-samples here discussed are as large as might be desired. Therefore,
-conclusions based upon them (see below) are to be regarded as
-tentative. Many other, future, samples will perhaps also be
-insufficient in size in themselves. There are, however, statistical
-advantages to repetition which will serve to make the repeated
-analysis even of small samples significant and valuable.
-
-Certain of the samples not treated statistically show ratios that can
-be seen by inspection to be probably significant. For example the
-almost complete absence of adults from the three samples of Red-eyed
-Vireos (Fig. 1E) cannot be disregarded in view of the size of the
-whole sample of the species. The same applies to the high percentage
-of adult females and the near absence of adult males in the sample of
-the Dickcissel (Fig. 1F). The continuity in direction of changes
-observed in the three samples of the Mourning Warbler (Fig. 1G) and
-Red-eyed Vireo is likewise probably significant, even though some of
-the samples compared are small. It seems to us that the application of
-statistical methods to these species should await the accumulation of
-more material. For anyone desiring to treat them statistically now,
-the data are inherent in this paper.
-
-We have not computed the standard errors of the ratios of sexes within
-age groups (except experimentally in a few cases). This can easily be
-done, however, and the significance of a given ratio determined, on
-the assumption (perhaps sometimes dubiously justifiable) that the
-sex-ratio in the species concerned is one:one. Obviously there is no
-point in computation of the standard errors of adult-immature ratios
-in single samples (such as that of the Dickcissel) until the actual
-ratio prevailing in the species in nature at the season in question is
-known for comparison with the observed ratio. Our formal statistical
-treatment, therefore, has been limited to an examination of the
-significance of the _changes_ between adult-immature ratios in samples
-of the same species taken a number of days apart.
-
-The samples suggest several patterns of differential migration of sex-
-and age-classes. Indeed, the important consideration brought out--in
-our opinion not hitherto sufficiently emphasized in literature--seems
-to be that in generalizing about adults and immatures, one must be
-careful to take sexes into account, and conversely, in generalizing
-about males and females, one must consider also age. In other words,
-there are really four classes to be considered. This poses additional
-problems in analysis and introduces the need for still larger samples
-in order to reach significant conclusions. To illustrate: an
-adult-immature ratio of 40:20 (N = 60) may be satisfactorily
-significant, while within the 40 adults a ratio of 25 males:15 females
-may not be. Were the original sample 80:40 (N = 120) with male adults
-50 and female adults 30, it is obvious that the significance of the
-latter ratio would be greater. The same applies in reverse if the
-greater emphasis is placed on sex and the lesser on age. Because of
-the moderate size of the samples this problem has been felt in the
-present study in respect to sex ratios within age groups, many of
-which must at present be regarded as of tentative significance.
-
-In short, what the earlier ornithologists regarded as a simple problem
-is in reality a complex one. There are only two patterns in what may
-be called the Brewster-Gätke argument: adults first or immatures first
-(with of course the further possibility of both at the same time).
-Both patterns occur, as is now known, at least to some extent. But
-actual patterns, as suggested by our samples, are more complex when
-all classes are considered. It will readily be seen that, if adult
-males, immature males, adult females, and immature females be regarded
-as units, each with certain migratory characteristics, the
-combinations of these units in various orders of migratory precedence
-are potentially numerous. In fact, of course, they do not behave
-strictly as units (or perhaps very rarely so), but our data strongly
-indicate that the tendency exists in many cases. This may be stated
-another way. The present samples may be reduced to two basic patterns,
-fitting the classic early American (adults first) and early European
-(immatures first) theories. But, either such simple arrangement is
-compounded in some, perhaps in truth in all, instances by differential
-migration of the sexes _within_ each age class. This proposition can
-also be stated backwards: the samples show differential times of
-migration of the sexes, compounded by differential times of migration
-of the age groups within each sex. The order in which these matters
-are approached depends on what one is trying to find out. Influenced
-by the literature, in which most emphasis has been placed on age, we
-have approached the problem from that standpoint. The data and figures
-here given, however, can be juggled if one wishes to place first
-emphasis on the order of sexes in migration.
-
-Bearing in mind what has just been said, particularly in respect to
-sizes of samples necessary for significance, let us consider the
-patterns of migration suggested by the Topeka sample. These are as
-follows:
-
-(1) _An early migration largely composed of adults, giving way later
-on to a preponderance of immatures._ Regardless of variations among
-them, samples showing this basic pattern are in line with the opinions
-of Brewster (1886) and his followers. This pattern is here shown by
-the Lincoln Sparrow, Yellow-throat, Nashville Warbler, Catbird (one
-sample only), and Red-eyed Vireo (Fig. 1, A, B, C, D, E). The evidence
-of these and all other samples would admittedly be more conclusive if
-the samples were further apart in time or, better still, were there
-more of them. There is evidence that differences in migration of the
-sexes, within age classes, influence this pattern, sharply in some
-instances. In the later samples of Lincoln Sparrow, Yellow-throat, and
-Red-eyed Vireo (Fig. 1, A, B, E) there are relatively fewer males,
-both adult and immature, than in the earlier samples and this may be
-true also of the Catbird, judging from the single sample. The Red-eyed
-Vireo (Fig. 1, E) is characterized by small number, or absence of,
-males in each sample but the samples are not significantly different,
-and can be regarded as one. Although the samples of the Dickcissel and
-Mourning Warbler (Fig. 1, F, G) show a somewhat different over-all
-pattern and are discussed further on, they also contain few adult
-males. Since these samples are from a period that is near the end of
-the migration of Red-eyed Vireos, Mourning Warblers, and Dickcissels,
-it may be assumed tentatively that the adult males have already
-migrated. Meinertzhagen (1930:56) postulated that in many species
-there is an earlier or more rapid migration of adults, particularly
-males, and the data for the above species in our sample tend to
-support his assumption. But our data suggest in addition that in some
-species _immature males_ migrate earlier, or more rapidly, than do
-_immature females_, just as adult males precede adult females in some
-instances. Within this general pattern (adults first) another
-variation is shown by the Nashville Warbler (Fig. 1, C) in which the
-later sample of adults is heavily weighted towards males, even though
-an increasing over-all proportion of immatures is evidenced. In this
-case, and contrary to Meinertzhagen's suggestion, it would seem that
-adult females have preceded or outstripped adult males in migration.
-
-(2) _An early preponderance of immatures, followed by a preponderance
-of adults._ The several species of birds at Topeka that display this
-pattern conform with the conclusions of Gätke and other early Old
-World ornithologists that in most species immatures precede adults in
-migration. In the present sample two variations of this pattern occur.
-
-(a) In the Dickcissel (Fig. 1, F) and the Mourning Warbler (Fig. 1,
-G), immatures decrease more markedly than adults (visible in samples
-of Mourning Warbler; inferred in Dickcissel), leaving the adults in
-the majority in the closing phase of migration. The distinctive and
-interesting feature in each of these two species is the ascendancy in
-numbers of adults _despite_ the almost complete disappearance of adult
-males. The relative increase of adults is here caused by a retarded
-migration of adult females, which linger conspicuously behind all
-other classes. Something of this nature was suggested, in theory, by
-Dixon (1892:70) who thought that adult females are delayed by
-"maternal duties." It was hinted at also by Dwight (1900:127) who
-thought that in some species females molted later than males as a
-result of prolongation of parental responsibilities. As mentioned
-already, there is need for caution in interpreting the present samples
-because the Dickcissel is represented only by one sample and two of
-the three samples of Mourning Warblers are small. In the case of the
-Mourning Warbler, the samples may be regarded as one, nearly lacking
-in adult males. The progressive increase of adult females, however,
-may be significant; at least there are enough of these to make
-division of the birds into three samples enlightening. There is, of
-course, some chance that the majority of adult males have not yet
-migrated, or are migrating by a different route. This seems unlikely
-in both cases. October 1 is late in the migration of the Dickcissel
-and it seems that large-scale migration would not occur much later,
-and in the case of the Mourning Warbler adult males are rare in all
-three samples, extending over a considerable period and reaching late
-into the probable migration period of the species. It is interesting
-to conjecture just when and where adult male Mourning Warblers do
-migrate in autumn. Brewster (1886:16) wrote: "This species arrives at
-Cambridge [Massachusetts] about September 12, and during the remainder
-of the month is ... abundant.... The adults, however, are so very
-uncommon that I have never known them [to] represent more than five
-per cent of the total number of individuals. They do not seem to be
-more numerous in the earlier flights than towards the close of the
-month, and I am very sure that they cannot be found in this locality
-before the young begin to appear." While the present samples show an
-abundance of adult _females_ of this species (could Brewster have
-failed to recognize these as adults?) the whereabouts of the adult
-males remains a mystery.
-
-(b) Another variation is displayed by the Orange-crowned Warbler (Fig.
-1, H). Here also there is an increase of adults towards the end of
-migration, but this increase is marked by a growing percentage not of
-females but of males. Locally this species is a late migrant compared
-with most others of the Parulidae. Thus the first sample, composed of
-birds taken September 25-October 1, may be regarded as fairly early in
-the fall migration. Immature birds compose 84.2 per cent of this
-sample, there being no adult males at all. By October 5-9 the picture
-has changed markedly, the sample being composed of 44 per cent adults
-(82 per cent of which, in turn, are males) and 56 per cent immatures.
-In view of this trend one can not help suspecting that a still later
-sample would show a majority of adults, perhaps nearly all males. This
-of course does not necessarily follow; the migration of immatures
-could simply be more protracted, and could have commenced earlier,
-than that of adults.
-
-Little imagination is required to see how enlightening it might be
-could we analyze thoroughly the patterns of all migrating species.
-When the detailed facts are available, it seems likely that general
-trends will emerge which may be of great significance to the study of
-migration in general. A final point which must eventually be clarified
-is determination of the extent of variability in the pattern of each
-species from year to year and locality to locality.
-
-Once patterns of precedence in migration of different classes are
-established, search into the life-histories of the species concerned
-may help to explain the peculiarities discovered. In the present case,
-for instance, we find a possible clue to the reason for the high
-proportion of adult females of the Dickcissel late in migration, as
-shown by our sample. Gross (1921:14-15) presented evidence that adult
-female Dickcissels molt considerably later than their mates, and we
-have independent evidence that individuals of this species are at
-times almost flightless when molting the remiges!
-
-
-
-
-Molt in Relation to Migration
-
-
-GENERAL COMMENT.--The exact relationship between molt and migration
-seems not to have been definitely established. The subject has
-received cursory attention in the literature and conflicting opinions
-have been expressed. Dwight (1900:126-128) believed that molt is
-completed or nearly completed before migration in nearly all passerine
-species that occur in New York save for certain swallows and
-flycatchers. Molt has since been found to precede migration of at
-least one of the flycatchers (_Empidonax virescens_) considered by
-Dwight to be an exception to this rule (Mengel, 1952). In Great
-Britain the subject of molt in migration was considered in some detail
-by Rintoul and Baxter (1914) and Ticehurst (1916), who arrived at
-conclusions similar to Dwight's. These workers also found certain
-swallows to be exceptions to the rule.
-
-The above authors and others have shown that, at least among
-passerines, some body molt is frequently found in migrating
-individuals but that molt of tail feathers is much less often found
-and molt of remiges almost nonexistent. Baxter and Rintoul noted only
-four cases of molting wing feathers among hundreds of migrants. Among
-the diverse non-passerine orders the picture seems to be more
-complicated, as might be expected. We do not, however, comprehend the
-reasoning which led Meinertzhagen (1930:56) to summarize: "... on the
-whole it can be said that though birds seldom migrate when flight
-feathers are in quill, moult in general does not influence migration."
-This seems to us an obvious _non sequitur_. Meinertzhagen (_loc.
-cit._) went on to say: "Males and females of one species are believed
-to moult simultaneously [see, however, Dwight, 1900:127], and there is
-no doubt that in some cases the two sexes migrate at slightly
-different times, and occasionally prefer different winter quarters.
-Birds of the year never moult their quills previous to their first
-autumn migration [Consultation of Dwight, 1900, who gives many
-examples of this, would have spared Meinertzhagen this error.], and
-yet they frequently follow adults on passage and sometimes precede
-them. There are no grounds for believing that adults have moulted
-their quills before birds of the year are prepared to migrate [but
-there are, in many cases; _cf._ Dwight, 1900:127], in the case where
-adults precede the latter. Neither is there any evidence to show that
-adults have not moulted their quills till after their offspring are
-ready for passage, in the case where they follow their offspring. It
-does not, therefore, appear that moult is an important factor."
-
-Comments interpolated above show our feeling that this summary is
-inadequate and misleading. To us it seems that the extreme rarity in
-migration of birds with remiges in molt is strong evidence that molt
-_does_ influence at least the time of migration. It is immaterial
-whether this molt occurs before or after migration, although in the
-majority of cases it seems to take place before. Much more needs to be
-known of the migration pattern and molt of each species before
-generalizations can safely be made.
-
-Analysis of samples of migrants can show only the presence and nature
-or the absence of molt in birds actually migrating. In the present
-instance shortage of time and manpower for preserving some and
-processing all of the sample resulted in incomplete data being kept on
-molt. We include this section to emphasize uncertainties still
-prevalent and to stimulate further work.
-
-
-MOLT IN THE TOPEKA SAMPLE.--Our limited findings coincide with those
-of Rintoul and Baxter (1914). Body molt was noted in a number of
-individuals and species. When present, this molt almost invariably was
-in its final stages. One immature male Rose-breasted Grosbeak (October
-1) was in heavy body molt. It is perhaps worthy of mention here that
-this grosbeak evidently migrates at times in extensive molt. An adult
-male (RMM 1102) taken by Mengel near Henderson, Kentucky, on September
-9, 1949, was molting plumage of body, wings, and tail, no feather of
-the last being longer than one half inch. This remarkable specimen had
-only five primaries on one side and four on the other fully
-functional. The outermost on the left and two outermost on the right
-were from the previous plumage, not yet dropped; the three innermost
-of each wing were new and full-length.
-
-In the present sample molt of remiges was noted in one specimen, an
-adult female Indigo Bunting (October 1) with outer primaries sheathed
-and with molt in progress in the body plumage. The one (immature)
-Yellow-breasted Chat in the sample (October 1) had all of its tail
-feathers nearly full-length but in quill, possibly as a result of
-accident, and two feathers were being replaced also in the tail of an
-immature Clay-colored Sparrow (October 6), which was also in body molt
-and had some, juvenal feathers on the belly and flanks.
-
-Body molt near completion was further noted as follows: immature male
-Yellow-throated Vireo (October 1), adult male Blue-headed Vireo
-(October 1), immature female Leconte Sparrow (October 23), several
-Lincoln Sparrows (various dates).
-
-
-
-
-Size Differences according to Sex and Age
-
-
-LINEAR MEASUREMENTS.--Taxonomists long have recognized in many species
-that males differ in size from females. Less attention, until
-recently, has been paid to the relative sizes of adult and immature
-birds. Many taxonomists, however, seem to have had an uneasy suspicion
-that immature birds are "untrustworthy" in comparison with adults, and
-immatures have often been excluded from samples when recognizable.
-Since, however, there are still relatively few reliably aged specimens
-in collections, for the most part only those immature birds
-immediately recognizable as such by obvious plumage differences (which
-are often present only in juvenal plumage) have been excluded from
-series. The majority of birds in first winter plumage so closely
-resemble adults that the two ages have been included in series for
-measurement. In most passerines these younger birds still bear the
-juvenal feathers in wing and tail and are, in size of these important
-parts, quite as "untrustworthy" as birds still in juvenal body
-plumage. Even if a complete postjuvenal molt occurs we still should
-not assume that first winter feathers are as long as adult winter
-feathers without first determining that this is so. Although aware of
-this problem, systematists until recently seemingly have been more or
-less content to disregard it, or forced to do so for practical
-reasons. Miller (1941:179) had little choice but to hope that size
-differences between adult and immature juncos were unimportant. Behle
-(1942:217) wrote of Horned Larks, _Eremophila alpestris_: "... the
-plumages of first-year birds and adults seem indistinguishable, though
-I have never quite satisfied myself that there are no differences in
-lengths of rectrices and remiges." He added, with logic confusing to
-us: "Since it is a difficult problem to determine the ages of horned
-larks that have passed the postjuvenal molt, this similarity of
-plumages is fortunate for the systematist."
-
-In recent years, some workers have analyzed size differences between
-adults and immatures. Sibley (1950:115) showed that adult Red-eyed
-Towhees (_Pipilo erythrophthalmus_) had notably longer wings and tails
-than immatures, and the same was demonstrated in Red Crossbills
-(_Loxia curvirostra_) by Tordoff (1952). In work with jays
-(_Aphelocoma_), Pitelka (1951:199) found that: "... in comparisons of
-dimensions of sex and age groups within a given sample, although
-magnitude of difference varies from one character to another, most of
-the averages are successively smaller for first-year males and adult
-and first-year females." He listed exceptions and concluded:
-"Segregation [of sex and age classes] has proved to be of extreme
-significance in an interpretation of individual and geographic
-variation."
-
-Much along these lines can be learned by examination of large random
-samples such as that afforded by the Topeka accident. Although only a
-few species in this sample were measured, the results secured seem to
-show further the need for segregation of age classes in taxonomic work
-with some species.
-
-Figure 2 shows the variation in the lengths of wing and tail in the
-Nashville Warbler. It is evident from the figure that in both of these
-characters the four sex- and age-classes differ significantly. An
-accurate idea of geographic variation in this species could not be
-obtained without separating these classes in comparisons. Age classes
-in spring and summer, long after the skull is completely ossified, can
-be segregated only if differences in plumage can be found. In the
-Nashville Warbler, such differences occur in fall (see annotated list)
-but these probably are obliterated by the partial prenuptial molt.
-These facts emphasize the importance, for taxonomic studies, of aged
-specimens collected in late summer or early fall on their breeding
-ground and in fresh winter plumage. Figure 3 shows the variation in
-length of wing in the Lincoln Sparrow in which age seems to be of much
-less importance than in the Nashville Warbler. Males and females of
-the Lincoln Sparrow differ significantly in wing-length, but adults
-and immatures are of nearly the same size. It would seemingly not be
-necessary to separate age classes in studies of geographic variation
-in the Lincoln Sparrow. Size data for some other species are given in
-the annotated list.
-
-
-WEIGHTS.--Little seems to have been done to determine the correlation
-of weights with sex- and age-classes. Weight may be the best single
-index of over-all size and is especially useful to students of the
-physiology of migration. Weight, however, is subject to the
-considerable variable imposed by fat condition. In large and
-comparable series, this variable is probably removed insofar as
-comparability of means is concerned, but the high variability of
-weight in most cases naturally results in more overlap (less
-separability) between populations with means close together than is
-found with most linear measurements.
-
-[Illustration: FIG. 2. Statistics of variation in length of wing and
-tail in the Nashville Warbler. The solid lines represent the observed
-ranges in millimeters. The stippled boxes represent two standard
-errors ([Greek: s]m) to each side of the means (vertical lines). The
-open boxes represent one standard deviation ([Greek: s]) to each side
-of the means.]
-
-[Illustration: FIG. 3. Statistics of variation in length of wing in
-the Lincoln Sparrow.]
-
-Figures 4-6 show variation in weight in the samples of Nashville
-Warbler, Mourning Warbler, Yellow-throat, Dickcissel, and Lincoln
-Sparrow. Each figure is essentially self-explanatory. It will be seen
-that in the Nashville Warbler and Lincoln Sparrow, weight is roughly
-proportional to wing-length (shown in figs. 2 and 3), giving about
-equally good separation of sex- and age-classes in the latter and
-poorer separation in the former. Data for these and other species
-indicate a generally greater weight of males than of females, but less
-difference according to age. In some other species, for example the
-Yellow-throat, immatures seem to be a little heavier on the average
-than adults. It is unfortunate that wing-lengths are not at present
-available for these.
-
-[Illustration: FIG. 4. Statistics of variation in weight in the
-Nashville Warbler and Mourning Warbler.]
-
-[Illustration: FIG. 5. Statistics of variation in weight in the
-Yellow-throat and Dickcissel.]
-
-[Illustration: FIG. 6. Statistics of variation in weight in the
-Lincoln Sparrow.]
-
-These comments on weight suggest an additional factor which may play a
-part in rate of migration and which some day may be profitably
-studied. Suppose that in some species adults and immatures are nearly
-the same in weight but that immatures have shorter wings. In such a
-species the immatures are relatively shorter-winged for their weight
-than adults. In aerodynamic terms, they would have a higher
-"wing-loading." (Wing-loading is the result obtained by dividing area
-of effective wing by total weight; it is here assumed that in a single
-species wing area is directly proportional to wing length.) This being
-the case, immatures with higher "wing-loading" would require more
-energy (derived from fat) to fly the same distance as adults, or with
-the same amount of fat they would fly a shorter distance. Thus they
-might tend to be outstripped in migration by adults starting at the
-same time. The reverse, of course, would also be theoretically true,
-if adults possessed a higher wing-loading than immatures. Physical
-factors such as these rather than the differential "virility"
-postulated by Meinertzhagen (1930:56) might account for the arrival of
-certain classes of some species on the wintering grounds in advance of
-others. There are, of course, many other factors which must be taken
-into account before the effect, if any, of the wing-loading factor can
-be evaluated. Data for illuminating calculations will become
-available, however, with the accumulation of abundant information on
-weights, measurements, and migration patterns.
-
-
-
-
-Computations of Longevity and Survival
-
-
-Tanner (ms., and letter, April 21, 1955) recently devoted considerable
-ingenuity to computing by actuarial methods the longevity of the
-Oven-bird, using the adult-immature ratio in samples killed at a
-ceilometer at Knoxville, Tennessee. Tanner's computations were based
-on the simple assumption that
-
-S (survival rate) = Number of adults in population (or sample) / Total
-size of population (or sample).
-
-Further application of such techniques may prove desirable and
-rewarding. It would seem at present, however, to be a risky procedure,
-as it has been abundantly shown (see above) that adults and immatures
-often do not migrate at the same times and rates, and the ratios of
-adults to immatures in samples of migrants are likely to be far from
-representative of the true proportions in the populations concerned.
-It should be added that Tanner is perfectly aware of this objection.
-
-
-
-
-Processing of Samples
-
-
-Thorough processing of large samples of birds killed accidentally is
-time-consuming. We were fortunate in having considerable assistance;
-even so, all desirable data could not be obtained from the 1090 birds
-of the present sample. As aids to others conducting studies of this
-kind we should mention a few points which may be of assistance.
-
-Birds should be picked up as soon as possible after death, certainly
-by the end of the day after the accident and preferably much sooner.
-They should be weighed as soon as possible after collection (weights
-decrease rapidly, even under refrigeration), and the weights (in
-grams, to one tenth of a gram) written on tags attached to a leg of
-each specimen. The sample should then be sorted by species or groups
-of species of approximately equal size (to avoid crushing of smaller
-birds by larger ones), placed in boxes, paper bags, or better,
-air-tight containers clearly marked with date, locality, and other
-necessary particulars, and relegated to a deep-freeze compartment. For
-all but the smallest samples, such freezing units are indispensable to
-complete study. Once frozen, the birds may be selected for study at
-leisure, but time is still important, as, even when frozen, gonads may
-eventually deteriorate, and birds eventually become desiccated which
-is a disadvantage if skins are to be made.
-
-In the cases of large kills, or limited manpower, or both, it may be
-impossible to process all birds, however desirable this might be. If
-possible, however, all should be collected, identified, the numbers
-and species recorded, and rarities saved. Further, partial analysis,
-or more properly, complete analysis of a partial sample, can be made.
-Analyses which for any reason require randomness of sample pose a
-special problem. We think that in very large kills the best way to
-solve this problem is probably to make one or more transects across
-the area where dead birds are found. These transects should cross both
-the areas of greatest and least density (to allow for fast and slow
-flying species). Their width may be adjusted to give the desired
-number of birds, that is, the number that can be adequately processed.
-Another alternative would be to decide to study certain abundant
-species and pick up all of these. There are other possibilities, but
-in any event the method of sampling should be thoroughly described
-wherever all birds have not been processed.
-
-
-
-
-Summary
-
-
-The foregoing paper discusses accidents in which large numbers of
-night-migrating birds are killed. A brief historical review of
-ornithological interest in such occurrences is given, and the types of
-data provided by the accidents are listed and discussed. In
-particular, recent accidents occurring in early October, 1954, through
-much of eastern United States are cited, and detailed analysis is
-presented of a sample of 1090 birds killed one mile west of Topeka,
-Shawnee County, Kansas, between September 25 and October 23, 1954.
-
-At Topeka during the period mentioned, 1090 birds representing 61
-species were collected and were processed at the University of Kansas.
-For all specimens, weight, sex, age, and fat condition were recorded,
-and certain species were measured as well. Some notes on molt were
-taken. A total of 193 birds was preserved as study skins, and 49 as
-skeletons. Comments on weight, size, sex, age, subspecific identity,
-and status in Kansas are presented in an annotated list.
-
-Randomness of this and other similar samples is discussed. A
-theoretical computation is given for several nights of the numbers of
-migrants passing through a plane one mile in width, from 450 to 950
-feet above ground level, and oriented to face the arriving migrants.
-The computed totals give some idea of the tremendous volume of
-nocturnal migration under some conditions. Potentialities of further
-study of such events are discussed and a comparison is made with lunar
-observations.
-
-Differential migration of sex- and age-groups as shown by the larger
-samples taken at Topeka (Catbird, Red-eyed Vireo, Nashville Warbler,
-Yellow-throat, Mourning Warbler, Dickcissel, Lincoln Sparrow) is
-discussed and the history of theories on this subject reviewed. It is
-shown that age and sex must both be taken into account in studies of
-differential migration. Several patterns of migration are displayed by
-the species analyzed, adults migrating earlier than immatures in some
-instances, young earlier than adults in others, but all seemingly
-being complicated to varying degrees by differential migration of
-sexes within age groups. It is suggested that explanations of these
-patterns may be sought in the life histories of the species involved.
-
-Molt in relation to migration is discussed briefly, and it is held
-that there is an important relationship between molt and time of
-migration. Specimens noted to be in molt are listed.
-
-Size differences, in wing length, tail length, and weight are
-discussed in relation to sex and age, and variation in one or more of
-these characters is analyzed as found in the Topeka samples of
-Nashville Warbler, Mourning Warbler, Yellow-throat, Dickcissel, and
-Lincoln Sparrow. It seems that in some instances significant size
-differences prevail between adults and immatures and that these age
-classes should be separated in taxonomic work with species where
-differences in size are known to exist. When the facts are not known
-they should be determined, and the large samples collected in
-accidents to nocturnal migrants present excellent opportunities for
-ascertaining the facts.
-
-Other uses of material obtained in large migration accidents are
-discussed, such as computations of longevity and the problems of
-processing large, accidentally-killed samples. Care should be taken to
-select samples as nearly random as possible when all birds cannot be
-processed.
-
-Repeated and thorough analysis of accidental kills should provide a
-mass of valuable data bearing on many questions and problems which
-have thus far been little studied.
-
-
-
-
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-_Transmitted June 30, 1955._
-
-
-
-
-Transcriber's Notes:
-
-
-Words surrounded by _ are italicized.
-
-Small capitals are presented as all capitals in this e-text.
-
-Obvious printer's errors have been repaired, other inconsistent
-spellings have been kept, for example inconsistent use of hyphen (e.g.
-"age-classes" and "age classes") and diacritical mark (e.g. "Zool."
-and "Zoöl.").
-
-
-
-
-
-End of the Project Gutenberg EBook of Studies of Birds Killed in Nocturnal
-Migration, by Harrison B. Tordoff and Robert M. Mengel
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-<pre>
-
-The Project Gutenberg EBook of Studies of Birds Killed in Nocturnal
-Migration, by Harrison B. Tordoff and Robert M. Mengel
-
-This eBook is for the use of anyone anywhere in the United States and most
-other parts of the world at no cost and with almost no restrictions
-whatsoever. You may copy it, give it away or re-use it under the terms of
-the Project Gutenberg License included with this eBook or online at
-www.gutenberg.org. If you are not located in the United States, you'll have
-to check the laws of the country where you are located before using this ebook.
-
-
-
-Title: Studies of Birds Killed in Nocturnal Migration
-
-Author: Harrison B. Tordoff
- Robert M. Mengel
-
-Release Date: June 20, 2016 [EBook #52382]
-
-Language: English
-
-Character set encoding: ISO-8859-1
-
-*** START OF THIS PROJECT GUTENBERG EBOOK STUDIES OF BIRDS ***
-
-
-
-
-Produced by Judith Wirawan, Chris Curnow, Joseph Cooper
-and the Online Distributed Proofreading Team at
-http://www.pgdp.net
-
-
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-</pre>
-
-<hr class="chap" />
-
-
-
-<p class="center"><span class="smcap">University of Kansas Publications</span></p>
-
-<p class="center"><span class="smcap">Museum of Natural History</span></p>
-
-<hr />
-
-<p class="center">Volume 10, No. 1, pp. 1-44, 6 figures in text, 2 tables</p>
-
-<p class="center">&mdash;&mdash;&mdash;September 12, 1956&mdash;&mdash;&mdash;</p>
-
-
-<h1 class="space-above">Studies of Birds<br />
-Killed in Nocturnal Migration</h1>
-
-
-<p class="center">BY<br />
-HARRISON B. TORDOFF AND ROBERT M. MENGEL</p>
-
-
-<p class="center space-above"><span class="smcap">University of Kansas<br />
-Lawrence</span><br />
-1956</p>
-
-<hr class="chap" />
-
-<p class="center"><span class="smcap">University of Kansas Publications, Museum of Natural History</span></p>
-
-<p class="center">Editors: E. Raymond Hall, Chairman, A. Byron Leonard,
-Robert W. Wilson</p>
-
-<p class="center space-above">Volume 10, No. 1, pp. 1-44, 6 figures in text, 2 tables<br />
-Published September 12, 1956</p>
-
-
-<p class="center space-above"><span class="smcap">University of Kansas</span><br />
-Lawrence, Kansas</p>
-
-<p class="center space-above">PRINTED BY<br />
-FERD VOILAND, JR., STATE PRINTER<br />
-TOPEKA, KANSAS<br />
-1956</p>
-
-<div class="figcenter" style="width: 111px;">
-<img src="images/logo.png" width="111" height="44" alt="" />
-</div>
-
-<p class="center">26-3856</p>
-
-<hr class="chap" />
-
-<p><span class="pagenum"><a name="Page_3" id="Page_3">[Pg 3]</a></span></p>
-
-<p class="center"><big>Studies of Birds<br />
-Killed in Nocturnal Migration</big></p>
-
-<p class="center">BY<br />
-HARRISON B. TORDOFF AND ROBERT M. MENGEL</p>
-
-
-
-<h2>Contents</h2>
-
-
-<div class="center">
-<table border="0" cellpadding="4" cellspacing="0" summary="Contents">
-<tr><td></td><td align="right"><span class="smcap lowercase">PAGE</span></td></tr>
-<tr><td align="left">Introduction</td><td align="right"><a href="#Page_4">4</a></td></tr>
-<tr><td align="left">Accidents to Migrating Birds in early October, 1954</td><td align="right"><a href="#Page_6">6</a></td></tr>
-<tr><td align="left"><span style="margin-left: 1em;">General</span></td><td align="right"><a href="#Page_6">6</a></td></tr>
-<tr><td align="left"><span style="margin-left: 1em;">Accidents at Topeka, Kansas</span></td><td align="right"><a href="#Page_6">6</a></td></tr>
-<tr><td align="left"><span style="margin-left: 1em;">Description of WIBW-TV tower</span></td><td align="right"><a href="#Page_7">7</a></td></tr>
-<tr><td align="left"><span style="margin-left: 1em;">Weather conditions</span></td><td align="right"><a href="#Page_7">7</a></td></tr>
-<tr><td align="left">Acknowledgments</td><td align="right"><a href="#Page_7">7</a></td></tr>
-<tr><td align="left">Notes on the Species Killed at Topeka</td><td align="right"><a href="#Page_8">8</a></td></tr>
-<tr><td align="left">Randomness of the Sample</td><td align="right"><a href="#Page_17">17</a></td></tr>
-<tr><td align="left">Number of Migrants</td><td align="right"><a href="#Page_18">18</a></td></tr>
-<tr><td align="left">Differential Migration of Sex- and Age-classes</td><td align="right"><a href="#Page_20">20</a></td></tr>
-<tr><td align="left"><span style="margin-left: 1em;">History of the subject</span></td><td align="right"><a href="#Page_20">20</a></td></tr>
-<tr><td align="left"><span style="margin-left: 1em;">Differential migration of sex- and age-classes as shown by the Topeka sample</span></td><td align="right"><a href="#Page_23">23</a></td></tr>
-<tr><td align="left">Molt in Relation to Migration</td><td align="right"><a href="#Page_29">29</a></td></tr>
-<tr><td align="left"><span style="margin-left: 1em;">General comment</span></td><td align="right"><a href="#Page_29">29</a></td></tr>
-<tr><td align="left"><span style="margin-left: 1em;">Molt in the Topeka sample</span></td><td align="right"><a href="#Page_30">30</a></td></tr>
-<tr><td align="left">Size Differences according to Sex and Age</td><td align="right"><a href="#Page_31">31</a></td></tr>
-<tr><td align="left"><span style="margin-left: 1em;">Linear measurements</span></td><td align="right"><a href="#Page_31">31</a></td></tr>
-<tr><td align="left"><span style="margin-left: 1em;">Weights</span></td><td align="right"><a href="#Page_32">32</a></td></tr>
-<tr><td align="left">Computations of Longevity and Survival</td><td align="right"><a href="#Page_38">38</a></td></tr>
-<tr><td align="left">Processing of Samples</td><td align="right"><a href="#Page_38">38</a></td></tr>
-<tr><td align="left">Summary</td><td align="right"><a href="#Page_39">39</a></td></tr>
-<tr><td align="left">Literature Cited</td><td align="right"><a href="#Page_41">41</a></td></tr>
-</table></div>
-
-<hr class="chap" />
-
-<p><span class="pagenum"><a name="Page_4" id="Page_4">[Pg 4]</a></span></p>
-
-
-
-<h2>Introduction</h2>
-
-
-<p>This paper is primarily an analysis of a sample of migrant birds
-killed in the autumn of 1954 by striking a television tower one mile
-west of Topeka, Shawnee County, Kansas. Secondarily, some aspects
-of migration involved in studies of this kind are discussed and
-historical background is presented.</p>
-
-<p>Considerable interest has been occasioned in recent years in the
-eastern United States by large-scale accidents to night-migrating
-birds. Most accidents have occurred in the autumn. The widespread
-adoption by airports of an instrument called the ceilometer,
-which measures the height of cloud ceilings by reflecting from them
-a high-powered beam of light, has proved under certain conditions
-to be catastrophic to night-flying birds. Among the recent reports
-of such accidents are those of Spofford (1949) and Laskey (1951)
-for Nashville, Tennessee, Howell and Tanner (1951) for Knoxville,
-Tennessee, and Lovell (1952) for Louisville, Kentucky. Recently
-Howell, Laskey, and Tanner (1954) reviewed ceilometer "tragedies"
-without being able to determine the exact reason for their lethal
-effectiveness. Less publicized so far have been mass collisions of
-birds with another class of obstacles, tall radio and television towers.
-These slender towers, usually 500 to 1000 feet tall, are increasing
-rapidly in numbers and there is reason to suppose that they will
-take a correspondingly larger toll of bird life.</p>
-
-<p>Notice has long been given by ornithologists to mass destruction
-of birds by more conventional solid obstructions to passage, and
-newspapers occasionally mention birds killed at such well-known
-points as the Washington Monument and the Empire State Building.</p>
-
-<p>Seventy-five years ago, J. A. Allen (1880) published the results
-of questionnaires circulated by William Brewster to lighthouse
-keepers. Brewster himself (1886) described destruction of birds
-at a lighthouse in the Bay of Fundy, paying keen attention to behavior
-of the birds and the exact conditions under which nocturnal
-flight and accidents occurred. The subject also received attention
-in several countries across the Atlantic. Destruction of birds at
-Irish lighthouses was carefully noted over a period of years and the
-results were published periodically, culminating in R. M. Barrington's
-massive report (1900) which remains in some ways the most
-thorough of its type.</p>
-
-<p><span class="pagenum"><a name="Page_5" id="Page_5">[Pg 5]</a></span></p>
-
-<p>While conservation-minded individuals have been concerned with
-the tremendous mortality involved in these various events, the ill
-wind blows some good in that, properly used, the data provided
-by such accidents can shed light on many obscure aspects of bird
-migration. Each accidental kill of birds affords a cross-section,
-approaching in variable degree a random sample, of the migrants
-passing a given point on a given date. The types of information
-provided by such kills are numerous, for example: (1) information
-on the presence of various species and the dates of their occurrence;
-(2) information on the relative abundance of species; (3) quantitative
-data on the relative sizes of males and females, and immatures
-and adults (of importance to taxonomic ornithology); (4)
-information on the relative times of migration of males, females,
-adults, and young; (5) information on molts and plumages; (6)
-quantitative information on composition by subspecies of migrants
-of the same species; (7) physiological data (fat condition, <i>etc.</i>)
-pertinent to the study of migration; and probably others.</p>
-
-<p>In spite of the great potential of this kind of material, the majority
-of ornithologists with access to such data have contented
-themselves with listing the species and sometimes the numbers of
-birds killed. A few have gone further. James T. Tanner (unpublished)
-attempted to compute the longevity of the Ovenbird
-(<i>Seiurus aurocapillus</i>) by analysis of ceilometer-killed birds at
-Knoxville, Tennessee (see below). Mention should be made of the
-reports of Rintoul and Baxter (1914) supplemented by Ticehurst
-(1916) who used rather small numbers of birds killed at Scottish
-lighthouses in studies of molt. However, the only effort to utilize
-the results of accidental kills on a large scale over a period of years
-appears to have been that, already mentioned, of Barrington (1900)
-and his co-workers in Ireland. An idea of the potentialities of the
-large recent kills in the United States may be obtained when it is
-recalled that in the 18 years of Barrington's work, which embodied
-some 1000 reports from lighthouse keepers, Barrington obtained for
-study only about 2000 specimens, many of these consisting of wings
-and feet only (Barrington's paper not seen in original; see J. A.
-Allen, 1901:205). More recently Dobben and Bruyns (1939) have
-analyzed the age and sex classes of some birds killed at lighthouses
-in Holland.</p>
-
-<p>As far as we have learned, there is no previous thorough analysis
-in the literature of large, accidentally-killed samples of birds. On
-the following pages we emphasize some of the uses which can be
-made of such material. We think that intensive analyses of such<span class="pagenum"><a name="Page_6" id="Page_6">[Pg 6]</a></span>
-events, whenever they occur, should become a regular part of
-ornithological investigation and that integration of numerous studies
-of such incidents will provide an unprecedented mass of information
-on migration.</p>
-
-
-
-<hr class="chap" />
-<h2>Accidents to Migrating Birds in early October, 1954</h2>
-
-
-<p><span class="smcap">General.</span>&mdash;The few days around the end of the first week of
-October, 1954, were notable for a series of accidents which occurred
-to migrating birds over much of eastern United States. So
-far as we know, these were all associated with an extensive belt of
-bad weather (cold fronts and stationary fronts) which covered
-much of the country during that period, and the accidents involved
-ceilometers and solid structures alike. Accidents known to us
-occurred as far south as Macon, Georgia (David W. Johnston, letter:
-Nov. 1, 1954), as far north as New York City, where many
-migrants were killed at the Empire State Building (<i>New York
-Times</i>, Thursday, October 7, 1954, p. 1) and elsewhere, and as far
-west as Smoky Hill Air Force Base at Salina, Kansas (ceilometer,
-October 7, some birds received at the University of Kansas). Some
-of the above, and incidents from a number of other localities, were
-mentioned in varying detail in <i>Audubon Field Notes</i> (vol. 9, no.
-1, pp. 6, 10, 15, 17, 18, 32, February, 1955). Still other accidents
-occurred at Columbia, Missouri (Richard P. Grossenheider, verbal
-communication), and Topeka, Kansas (present paper). Some probably
-have escaped our notice; summaries of some of these will probably
-appear in ornithological journals for some time to come. At
-Robins Air Force Base near Macon, Georgia, at least 50,000 birds
-were killed, of which about 2500, representing 54 species, were
-picked up (Johnston, <i>loc. cit.</i>).</p>
-
-
-<p><span class="smcap">Accidents at Topeka, Kansas.</span>&mdash;At Topeka, Shawnee County,
-Kansas, all birds were killed by collision with the newly-erected
-(1954) television transmitting tower of station WIBW-TV. This
-tower is one mile west of the city.</p>
-
-<p>The first casualties (see Table 1 for all others) were a Sora (<i>Porzana
-carolina</i>) and a Yellow-bellied Flycatcher (<i>Empidonax flaviventris</i>)
-found on September 7. The major accidents, however,
-occurred on the nights of September 24-25, September 30-October
-1, October 5-6, and October 6-7. Totals of birds picked up (probably
-over 95 per cent of birds killed) are given in Table 1, in which
-each date given is that of the day after the kill, <i>i. e.</i>, the date on
-which the birds were collected.</p>
-
-<p><span class="pagenum"><a name="Page_7" id="Page_7">[Pg 7]</a></span></p>
-
-<p>All major kills occurred on cloudy and foggy nights associated
-with frontal weather. Throughout the period a few birds struck
-the tower even on fairly clear nights, and minor but appreciable
-"falls" occurred on the nights of October 4-5, 7-8, and 22-23. A few
-birds killed probably were overlooked for a time and found their
-way into later samples. This is especially probable in the case of
-some birds entered under date of October 23, as many of these
-were somewhat desiccated. Weights clearly altered by desiccation
-or mutilation were not recorded. Reports of these accidents have
-been published by Carson (1954 a, b, and c).</p>
-
-<p>According to Carson (1954c:27), the majority of birds killed on
-nights of heavy flight fell "between three and four o'clock in the
-morning when skies were overcast and a cool front moved in from
-the north. Due to the cooperation of the watchmen it is thought
-that most of the birds that were killed were recovered. Of course
-some injured birds in hiding were not found and some were lost to
-predators."</p>
-
-
-<p><span class="smcap">Description of WIBW-TV TOWER</span>.&mdash;The tower is 950 feet tall
-and stands on a hill approximately 1000 feet above sea level. The
-fact that the tower is on a hill places the top of the tower at 1010
-feet above the elevation of the average local terrain. The tower
-is triangular in cross-section, each face seven feet wide, and is
-constructed of six-inch steel L-beams with three-inch cross-members
-every seven feet and smaller diagonal cross-members. It has no
-taper and bears a transmitting antenna on the top. The tower is
-supported by 12 guy wires, 3 wires attaching at each of 4 levels.
-The cables extend south, WNW, and NNE from the tower and are
-1-1/2 inches in diameter. The tower is lighted by a series of red lights,
-some flashing and others steady. The transmitter was not in operation
-when the accidents took place.</p>
-
-
-<p><span class="smcap">Weather conditions.</span>&mdash;All major kills at Topeka occurred when
-migrating birds encountered either a cold front or a stationary front
-lying over eastern Kansas. Typically, this frontal weather included
-rain, fog, and cloud ceilings down to as low as 800 to 1000 feet.
-Weather of this type presumably forces the migrating birds to fly
-below the cloud ceiling and thus brings them within the altitudinal
-range of the television towers.</p>
-
-
-
-<hr class="chap" />
-<h2>Acknowledgments</h2>
-
-
-<p>We gratefully acknowledge our debt to the Topeka Audubon Society for
-making this study possible by carefully collecting birds killed at the television
-tower. L. B. Carson deserves special mention for his general supervision of
-<span class="pagenum"><a name="Page_8" id="Page_8">[Pg 8]</a></span>the bird collecting by the members of the Topeka Audubon Society. Members
-of the Society and others who picked up birds under the television tower were:
-Mrs. Lloyd Biggs, Elaine Carson, L. B. Carson, Jesse A. Eddy, Elizabeth Fisher,
-Mrs. Walter Huxman, Florence McKinney, Mrs. Charles Martin, Mrs. Fred P.
-Martin, T. W. Nelson, Fred Prebble, Grace Prebble, Orville Rice, Mrs. G.
-Warren Scholl, E. W. Senne, and Beatrice Swenson.</p>
-
-<p>We received equally important assistance from students and staff of the
-University of Kansas in recording of data and preparation of specimens. The
-following helped in these ways: Rollin H. Baker, R. W. Dickerman, David
-L. Hardy, J. W. Hardy, Jane S. Mengel, Larry D. Mosby, Richard Van Gelder,
-South G. Van Hoose, and Glen E. Woolfenden. We are indebted to the Interlibrary
-Loan Service of the University of Kansas Library for help in securing
-certain reference works. Robert Sokal of the University of Kansas gave helpful
-advice concerning statistical procedures.</p>
-
-
-
-<hr class="chap" />
-<h2>Notes on the Species Killed at Topeka</h2>
-
-
-<p>A list of numbers and kinds of birds killed is given in Table 1.
-Discussion of data afforded by certain species for which, large
-samples were available will be found below. There are additionally
-certain data afforded by the sample and certain comments to
-be made on various species which can be handled most conveniently
-in an annotated list. In this list we have included all weight
-data (still scarce for many North American birds), comments on
-status in Kansas of various species, results of comparisons to determine
-subspecies, and miscellaneous observations. Weights of
-birds are given in grams and were taken on a triple-beam balance.
-Fat condition is given in the scale proposed by McCabe (1943:556).
-Weight data from birds migrating at night should be especially
-useful because these migrants all have relatively empty
-crops and stomachs, thus reducing variability. Not all birds were
-suitable for weighing and measuring, for a variety of reasons. This
-accounts for discrepancies in totals between Table 1 and the annotated
-list.</p>
-
-<p>All passerine species were aged by noting the degree of ossification
-of the skull. In no case, of the more than a thousand passerines
-aged by examination of the skull, did we find difficulty in determining
-whether an individual was a bird of the year or an adult.
-We found no specimens in which ossification of the skull was nearing
-completion. In the several species in our sample with distinctive
-first-winter plumages, we found complete agreement in
-age as shown by plumage and by condition of the skull. We think
-this is further proof, if such is needed, that this method of aging is
-thoroughly reliable in early autumn for the passerine species included
-in our sample and for others with similar breeding seasons.</p>
-
-<p><span class="pagenum"><a name="Page_9" id="Page_9">[Pg 9]</a></span></p>
-
-<p class="center"><span class="smcap">Table 1.&mdash;Birds Killed at a Television Tower at Topeka, Kansas, in 1954</span></p>
-
-<p>See annotated list for division into sex- and age-classes. Where discrepancies
-exist between totals given here and totals given in the annotated list, these
-result from the fact that some specimens could not be sexed and aged.</p>
-
-<div class="center">
-<table border="1" cellpadding="4" cellspacing="0" summary="Birds Killed">
-<tr><td></td>
- <td>Sept. 25</td><td>Oct. 1</td><td>Oct. 3</td><td>Oct. 4</td><td>Oct. 5</td><td>Oct. 6</td><td>Oct. 7</td><td>Oct. 8</td><td>Oct. 9</td><td>Oct. 10</td><td>Oct. 23</td>
- <td>Totals</td></tr>
-<tr><td align="left">Pied-billed Grebe</td>
- <td></td><td>1</td><td></td><td></td><td></td><td>1</td><td></td><td></td><td>1</td><td></td><td></td>
- <td>3</td></tr>
-<tr><td align="left">Green Heron</td>
- <td></td><td></td><td></td><td></td><td></td><td>1</td><td></td><td></td><td></td><td></td><td></td>
- <td>1</td></tr>
-<tr><td align="left">Blue-winged Teal</td>
- <td></td><td>1</td><td></td><td></td><td></td><td>1</td><td>6</td><td></td><td></td><td></td><td></td>
- <td>8</td></tr>
-<tr><td align="left">Virginia Rail</td>
- <td></td><td>3</td><td></td><td></td><td></td><td>1</td><td></td><td></td><td></td><td></td><td></td>
- <td>4</td></tr>
-<tr><td align="left">Sora</td>
- <td>1</td><td>6</td><td></td><td></td><td></td><td>1</td><td>1</td><td>1</td><td></td><td></td><td></td>
- <td>10</td></tr>
-<tr><td align="left">American Coot</td>
- <td></td><td></td><td></td><td></td><td></td><td>3</td><td></td><td></td><td></td><td></td><td></td>
- <td>3</td></tr>
-<tr><td align="left">Mourning Dove</td>
- <td></td><td>8</td><td></td><td></td><td>1</td><td></td><td></td><td></td><td></td><td></td><td>1</td>
- <td>10</td></tr>
-<tr><td align="left">Yellow-billed Cuckoo</td>
- <td></td><td></td><td></td><td></td><td></td><td>1</td><td></td><td></td><td></td><td></td><td></td>
- <td>1</td></tr>
-<tr><td align="left">Black-billed Cuckoo</td>
- <td></td><td></td><td></td><td></td><td></td><td></td><td>1</td><td></td><td></td><td></td><td></td>
- <td>1</td></tr>
-<tr><td align="left">Yellow-shafted Flicker</td>
- <td>3</td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td>
- <td>3</td></tr>
-<tr><td align="left">Yellow-bellied Flycatcher</td>
- <td></td><td></td><td></td><td></td><td></td><td></td><td>1</td><td></td><td></td><td></td><td></td>
- <td>1</td></tr>
-<tr><td align="left">House Wren</td>
- <td>2</td><td>3</td><td></td><td></td><td></td><td>1</td><td>2</td><td>1</td><td></td><td>1</td><td></td>
- <td>10</td></tr>
-<tr><td align="left">Long-billed Marsh Wren</td>
- <td></td><td>1</td><td></td><td></td><td></td><td>1</td><td>1</td><td></td><td></td><td></td><td></td>
- <td>3</td></tr>
-<tr><td align="left">Short-billed Marsh Wren</td>
- <td>1</td><td>2</td><td></td><td></td><td></td><td></td><td>1</td><td></td><td></td><td></td><td></td>
- <td>4</td></tr>
-<tr><td align="left">Catbird</td>
- <td>1</td><td>28</td><td>1</td><td></td><td>1</td><td>6</td><td>6</td><td></td><td></td><td></td><td></td>
- <td>43</td></tr>
-<tr><td align="left">Brown Thrasher</td>
- <td></td><td>1</td><td></td><td></td><td></td><td>1</td><td></td><td>1</td><td></td><td></td><td></td>
- <td>3</td></tr>
-<tr><td align="left">Wood Thrush</td>
- <td></td><td>3</td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td>
- <td>3</td></tr>
-<tr><td align="left">Hermit Thrush</td>
- <td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td>1</td><td></td>
- <td>1</td></tr>
-<tr><td align="left">Olive-backed Thrush</td>
- <td></td><td>14</td><td></td><td>1</td><td></td><td></td><td>1</td><td></td><td></td><td></td><td></td>
- <td>16</td></tr>
-<tr><td align="left">Golden-crowned Kinglet</td>
- <td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td>1</td><td>5</td>
- <td>6</td></tr>
-<tr><td align="left">Ruby-crowned Kinglet</td>
- <td>2</td><td>1</td><td></td><td></td><td></td><td></td><td>8</td><td>1</td><td>1</td><td></td><td>1</td>
- <td>14</td></tr>
-<tr><td align="left">Yellow-throated Vireo</td><td></td><td>1</td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td>
- <td>1</td></tr>
-<tr><td align="left">Blue-headed Vireo</td>
- <td>1</td><td>19</td><td></td><td>1</td><td>2</td><td>5</td><td>8</td><td>3</td><td>1</td><td></td><td></td>
- <td>40</td></tr>
-<tr><td align="left">Red-eyed Vireo</td>
- <td>18</td><td>36</td><td></td><td></td><td>2</td><td>13</td><td>2</td><td>3</td><td></td><td></td><td>1</td>
- <td>75</td></tr>
-<tr><td align="left">Philadelphia Vireo</td>
- <td>3</td><td>9</td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td>
- <td>12</td></tr>
-<tr><td align="left">Warbling Vireo</td>
- <td>8</td><td>19</td><td>1</td><td></td><td>4</td><td>1</td><td>1</td><td></td><td></td><td></td><td></td>
- <td>34</td></tr>
-<tr><td align="left">Black and White Warbler</td>
- <td>1</td><td>1</td><td></td><td></td><td></td><td>3</td><td></td><td></td><td></td><td></td><td></td>
- <td>5</td></tr>
-<tr><td align="left">Tennessee Warbler</td>
- <td></td><td>1</td><td></td><td></td><td>1</td><td>2</td><td>1</td><td></td><td></td><td></td><td></td>
- <td>5</td></tr>
-<tr><td align="left">Orange-crowned Warbler</td>
- <td>7</td><td>14</td><td></td><td></td><td>1</td><td>4</td><td>19</td><td>5</td><td>1</td><td>1</td><td></td>
- <td>52</td></tr>
-<tr><td align="left">Nashville Warbler</td>
- <td>7</td><td>94</td><td>4</td><td></td><td>3</td><td>39</td><td>27</td><td>5</td><td></td><td>1</td><td>1</td>
- <td>181</td></tr>
-<tr><td align="left">Parula Warbler</td>
- <td></td><td></td><td></td><td></td><td></td><td></td><td>1</td><td></td><td>1</td><td></td><td></td>
- <td>2</td></tr>
-<tr><td align="left">Yellow Warbler</td>
- <td>3</td><td>3</td><td></td><td></td><td></td><td>1</td><td>1</td><td></td><td></td><td></td><td></td>
- <td>8</td></tr>
-<tr><td align="left">Magnolia Warbler</td>
- <td></td><td>1</td><td></td><td></td><td></td><td>2</td><td></td><td></td><td></td><td></td><td></td>
- <td>3</td></tr>
-<tr><td align="left">Black-throated Blue Warbler</td>
- <td></td><td></td><td></td><td></td><td></td><td></td><td>2</td><td></td><td></td><td></td><td>1</td>
- <td>3</td></tr>
-<tr><td align="left">Myrtle Warbler</td>
- <td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td>1</td><td></td><td></td>
- <td>1</td></tr>
-<tr><td align="left">Black-throated Green Warbler</td>
- <td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td>1</td><td></td>
- <td>1</td></tr>
-<tr><td align="left">Chestnut-sided Warbler</td>
- <td></td><td>1</td><td></td><td></td><td></td><td></td><td></td><td></td><td>1</td><td></td><td></td>
- <td>2</td></tr>
-<tr><td align="left">Bay-breasted Warbler</td>
- <td>1</td><td></td><td></td><td></td><td></td><td>2</td><td></td><td></td><td></td><td></td><td></td>
- <td>3</td></tr>
-<tr><td align="left">Palm Warbler</td>
- <td>3</td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td>1</td>
- <td>4</td></tr>
-<tr><td align="left">Oven-bird</td>
- <td>4</td><td>21</td><td></td><td></td><td></td><td>2</td><td>3</td><td>1</td><td></td><td></td><td>1</td>
- <td>32</td></tr>
-<tr><td align="left">Northern Water-thrush</td>
- <td></td><td>5</td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td>1</td><td></td>
- <td>6</td></tr>
-<tr><td align="left">Mourning Warbler</td>
- <td>15</td><td>64</td><td></td><td></td><td>2</td><td>11</td><td>2</td><td>1</td><td></td><td></td><td></td>
- <td>95</td></tr>
-<tr><td align="left">Yellow-throat</td>
- <td>10</td><td>115</td><td>2</td><td></td><td>4</td><td>25</td><td>18</td><td>1</td><td>1</td><td></td><td></td>
- <td>176</td></tr>
-<tr><td align="left">Yellow-breasted Chat</td>
- <td></td><td>1</td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td>
- <td>1</td></tr>
-<tr><td align="left">Wilson Warbler</td>
- <td>1</td><td>2</td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td>
- <td>3</td></tr>
-<tr><td align="left">Canada Warbler</td>
- <td></td><td>2</td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td>
- <td>2</td></tr>
-<tr><td align="left">American Redstart</td>
- <td>1</td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td>
- <td>1</td></tr>
-<tr><td align="left">Bobolink</td>
- <td></td><td>4</td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td>
- <td>4</td></tr>
-<tr><td align="left">Rose-breasted Grosbeak</td>
- <td></td><td>2</td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td>
- <td>2</td></tr>
-<tr><td align="left">Indigo Bunting</td>
- <td></td><td>1</td><td></td><td></td><td>2</td><td>3</td><td>1</td><td></td><td></td><td></td><td></td>
- <td>7</td></tr>
-<tr><td align="left">Dickcissel</td>
- <td></td><td>31</td><td></td><td></td><td>1</td><td>3</td><td>1</td><td></td><td></td><td></td><td></td>
- <td>36</td></tr>
-<tr><td align="left">Savannah Sparrow</td>
- <td>1</td><td>6</td><td></td><td>1</td><td></td><td>1</td><td>5</td><td>1</td><td></td><td></td><td>1</td>
- <td>16</td></tr>
-<tr><td align="left">Grasshopper Sparrow</td>
- <td></td><td>7</td><td></td><td></td><td>2</td><td>3</td><td>3</td><td>1</td><td>1</td><td></td><td>1</td>
- <td>18</td></tr>
-<tr><td align="left">Leconte Sparrow</td>
- <td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td>3</td>
- <td>3</td></tr>
-<tr><td align="left">Sharp-tailed Sparrow</td>
- <td></td><td></td><td></td><td></td><td></td><td>1</td><td>1</td><td></td><td></td><td>1</td><td></td>
- <td>3</td></tr>
-<tr><td align="left">Slate-colored Junco</td>
- <td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td>1</td>
- <td>1</td></tr>
-<tr><td align="left">Clay-colored Sparrow</td>
- <td></td><td>11</td><td>1</td><td></td><td></td><td>2</td><td></td><td>1</td><td></td><td></td><td></td>
- <td>15</td></tr>
-<tr><td align="left">Fox Sparrow</td>
- <td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td>1</td>
- <td>1</td></tr>
-<tr><td align="left">Lincoln Sparrow</td>
- <td></td><td>41</td><td>7</td><td></td><td></td><td>5</td><td>22</td><td>3</td><td>1</td><td></td><td>3</td>
- <td>82</td></tr>
-<tr><td align="left">Swamp Sparrow</td>
- <td></td><td>1</td><td></td><td></td><td></td><td></td><td>1</td><td>2</td><td></td><td></td><td></td>
- <td>4</td></tr>
-<tr><td align="left">Song Sparrow</td>
- <td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td></td><td>2</td>
- <td>2</td></tr>
-<tr><td align="left">Total&mdash;species</td>
- <td>22</td><td>41</td><td>6</td><td>3</td><td>13</td><td>31</td><td>29</td><td>16</td><td>10</td><td>8</td><td>15</td>
- <td>61</td></tr>
-<tr><td align="left">Total&mdash;individuals</td>
- <td>94</td><td>585</td><td>16</td><td>3</td><td>26</td><td>146</td><td>147</td><td>31</td><td>10</td><td>8</td><td>24</td>
- <td>1090</td></tr>
-</table></div>
-
-<p><span class="pagenum"><a name="Page_10" id="Page_10">[Pg 10]</a></span>The annotated list may be consulted for further data in connection
-with the species listed in Table 1. As is indicated below, we
-regard the figures of this sample as unreliable to an unknown degree
-in comparing the relative abundance of one species with another.
-Accumulation of such data from various localities, however,
-should prove useful in another type of comparison. Samples of the
-same species killed in the same way at about the same time at different
-localities should be directly comparable. Eventually, this
-should provide us with a means of determining relative abundance
-of a species in different parts of its migratory route.</p>
-
-<p>Approximately 200 of the most interesting specimens were preserved
-as study skins and are in the University of Kansas Museum
-of Natural History. An effort was made to preserve at least one of
-each species, and we fell only a few short of this goal. All of the
-forms rare in Kansas are represented by skins. We could see no
-reason to list the preserved specimens in detail here. Species of
-which no study skins were made, however, are so marked.</p>
-
-<p>So far as we can tell, no truly western subspecies (from west of
-the Great Plains) occurred in the Topeka sample. Probably most
-or all of the birds came from areas more or less directly north of
-eastern Kansas.</p>
-
-<p>In critical areas where different subspecies of the same species
-occur together in migration, data from samples of this kind should
-prove enlightening. In future analyses, conducted in such areas,
-it might be possible to preserve all specimens of some of the variable
-species, or at least to measure all individuals of species in which<span class="pagenum"><a name="Page_11" id="Page_11">[Pg 11]</a></span>
-size is the most important variable character. Quantitative study
-could then be made of the different geographic variants occurring,
-their proportions in the migrant population determined, and their
-origins deduced. In studying populations of Painted Buntings
-(<i>Passerina ciris</i>) wintering in Mexico, Storer (1951) has provided
-an interesting demonstration of methods which can be applied to
-such samples.</p>
-
-<p>A few bats killed at the tower provided a surprise. They will be
-discussed separately by Richard Van Gelder.</p>
-
-<blockquote>
-
-<p><i>Podilymbus p. podiceps.</i> Pied-billed Grebe.&mdash;Weights: male, 394.8 (all
-weights in grams); females, 332.5, 289.7; all fat.</p>
-
-<p><i>Butorides v. virescens.</i> Green Heron.&mdash;Weight: 1 (unsexed), 168.6.</p>
-
-<p><i>Anas discors.</i> Blue-winged Teal.&mdash;Weights: 4 males, mean 421.2 (391.3-458.1);
-3 females, 367.7, 371.6, 393.2; all fat.</p>
-
-<p><i>Rallus limicola.</i> Virginia Rail.&mdash;Weights: 3 males, 73.7, 83.2, 90.5; 1 female,
-67.3; moderately fat to fat.</p>
-
-<p><i>Porzana carolina.</i> Sora.&mdash;Weights: 4 males, mean 76.8 (68.7-89.9); 3 females,
-62.6, 63.2, 63.5; moderately fat to very fat.</p>
-
-<p><i>Fulica americana.</i> American Coot.&mdash;Weights: 2 females, 385.3, 530.0, both
-fat. None preserved.</p>
-
-<p><i>Zenaidura macroura marginella.</i> Mourning Dove.&mdash;Weights: 2 adult males,
-121.8, 140.2; 3 immature males, 113.1, 126.1, 130.0; 3 adult females, 122.5,
-126.9, 136.0; 2 immature females, 129.4, 132.7; moderately fat to very fat.
-The presence of Mourning Doves in the sample is interesting as these birds
-are not generally regarded as night migrants. Conceivably the specimens were
-local birds going to roost. None preserved.</p>
-
-<p><i>Colaptes auratus luteus.</i> Yellow-shafted Flicker.&mdash;Weights: 2 males, 126.0,
-139.4, little fat. Flickers have several times been recorded as night migrants.</p>
-
-<p><i>Empidonax flaviventris.</i> Yellow-bellied Flycatcher.&mdash;Weight: 1 immature
-male, 11.9, moderately fat. This is a rare species in Kansas, the present being
-the ninth preserved specimen for the State.</p>
-
-<p><i>Troglodytes a&euml;don parkmanii.</i> House Wren.&mdash;Weights: 4 adult males,
-mean 10.5 (9.8-10.9), 2 immature males, 9.0, 11.3; 1 adult female, 9.9, 1 immature
-female, 7.0; no fat (im. &#9792;) to fat.</p>
-
-<p><i>Telmatodytes palustris dissa&euml;ptus.</i> Long-billed Marsh Wren.&mdash;Weights:
-1 adult male, 10.8; 1 adult female, 9.2; both moderately fat. The specimens
-are moderately bright and rufescent above, being typical of the populations of
-the central plains.</p>
-
-<p><i>Cistothorus platensis stellaris.</i> Short-billed Marsh Wren.&mdash;Weights: 1 immature
-male, 8.2; 1 adult female, 8.1; immature female, 8.2; all fat.</p>
-
-<p><i>Dumetella carolinensis.</i> Catbird.&mdash;Weights: 6 adult males, mean 37.5
-(34.1-42.5), little fat to very fat; 14 immature males, mean 37.57 &plusmn; .94 (standard
-error), S. D. (standard deviation) 3.37, little fat to fat; 11 adult females,
-mean 39.09 &plusmn; .94, S. D. 2.97, little fat to fat; 12 immature females, mean
-38.42 &plusmn; .83, S. D. 2.74, moderately fat to fat.</p>
-
-<p><i>Toxostoma r. rufum.</i> Brown Thrasher.&mdash;Weight: 1 immature male, 60.2,
-little fat.</p>
-<p><span class="pagenum"><a name="Page_12" id="Page_12">[Pg 12]</a></span></p>
-<p><i>Hylocichla mustelina.</i> Wood Thrush.&mdash;Weights: 1 adult male, 54.2, moderately
-fat; 2 adult females, 44.6, 45.7, little fat and fat, respectively.</p>
-
-<p><i>Hylocichla ustulata swainsonii.</i> Olive-backed Thrush.&mdash;Weights: 6 immature
-males, mean 31.0 (28.1-33.2), little fat to fat; 6 adult females, mean 29.6 (27.1-35.0),
-moderately fat to fat; 3 immature females, 27.1, 33.8, 35.8, little fat to fat.
-The absence of adult males in our sample of 15 birds is noteworthy but inexplicable
-with our few data.</p>
-
-<p><i>Regulus s. satrapa.</i> Golden-crowned Kinglet.&mdash;Weights: 1 adult male,
-6.7, moderately fat; 2 immature males, 6.5, 7.4, moderately fat and fat; 2 adult
-females, 7.3, 7.4, moderately fat and fat; 1 immature female, 7.2, moderately
-fat.</p>
-
-<p><i>Regulus c. calendula.</i> Ruby-crowned Kinglet.&mdash;Weights: 3 adult males,
-6.2, 7.6, 8.2, little fat to fat; 1 immature male, 6.6, fat; 4 adult females, mean
-6.1 (5.6-6.7), moderately fat to fat; 3 immature females, 5.8, 6.6, 7.0, moderately
-fat to fat.</p>
-
-<p><i>Vireo flavifrons.</i> Yellow-throated Vireo.&mdash;Weight: 1 immature male, 21.5,
-very fat.</p>
-
-<p><i>Vireo s. solitarius.</i> Blue-headed Vireo.&mdash;Weights: 9 adult males, mean
-17.7 (16.6-19.5), little fat to very fat; 17 immature males, mean 17.53 &plusmn; .46,
-S. D. 1.83, no fat (13.8) to very fat (21.3); 7 adult females, mean 17.6 (15.0-21.6),
-moderately fat to very fat; 6 immature females, mean 17.0 (14.5-18.9),
-moderately fat to fat. Surprisingly numerous in the sample.</p>
-
-<p><i>Vireo olivaceus.</i> Red-eyed Vireo.&mdash;Weights: 1 adult male, 16.1, moderately
-fat; 38 immature males, mean 21.21 &plusmn; .43, S. D. 2.60, little fat (1
-specimen) to excessively fat, mostly moderately fat or fat; 2 adult females,
-18.1, 18.1, both fat; 23 immature females, mean 19.28 &plusmn; .46, S. D. 2.16, little
-(2 specimens) to very fat, mostly fat.</p>
-
-<p>Wing length: 1 adult male, 79.1; 38 immature males, mean 78.05 &plusmn; .30,
-S. D. 1.80; 2 adult females, 76.3, 79.0, 23 immature females, mean 75.83 &plusmn;
-.42, S. D. 1.99.</p>
-
-<p>As mentioned below, the presence of only 3 adults in the sample of 64
-Red-eyed Vireos is highly significant and their occurrence only in the earlier
-samples is strong evidence of early migration by the adults.</p>
-
-<p><i>Vireo philadelphicus.</i> Philadelphia Vireo.&mdash;Weights: 2 adult males, 12.1,
-15.9, moderately fat and very fat; 2 immature males, 11.1, 13.2, fat and very
-fat; 2 adult females, 13.1, 14.2, both fat; 5 immature females, mean 14.1
-(12.0-15.2), moderately fat to very fat.</p>
-
-<p>This species previously has been collected in Kansas only twice. Both
-records are from Doniphan County in September, 1922. Field observers occasionally
-record the Philadelphia Vireo in eastern Kansas. Long (1940:450)
-calls it a "very rare migrant in the extreme east." Our sample of 12 birds
-killed on two nights (and probably after the peak of migration of this species)
-leads us to think that this vireo is actually a regular, but overlooked, migrant
-in fair numbers.</p>
-
-<p><i>Vireo g. gilvus.</i> Warbling Vireo.&mdash;Weights: 12 adult males, mean 15.92
-&plusmn; .43, S. D. 1.44, moderately fat to very fat; 8 immature males, mean 16.64
-(14.2-17.8), fat to very fat; 5 adult females, mean 16.1 (13.7-18.0), fat to
-very fat; 5 immature females, mean 15.4 (14.1-17.8), little fat to fat.</p>
-
-<p>Wing length: 12 adult males, mean 73.08 &plusmn; .49, S. D. 1.64; 8 immature
-<span class="pagenum"><a name="Page_13" id="Page_13">[Pg 13]</a></span>males, mean 71.15 (69.9-72.8); 5 adult females, mean 70.0 (69.2-71.0); 5
-immature females, mean 68.4 (67.7-70.3).</p>
-
-<p>Tail length: 12 adult males, mean 53.33 &plusmn; .53, S. D. 1.77; 8 immature
-males, mean 50.03 (47.1-51.3); 4 adult females, mean 48.6 (47.7-49.8); 5 immature
-females, mean 49.2 (47.3-53.0).</p>
-
-<p>There is no indication that western birds (<i>V. g. swainsonii</i>) make up any
-part of this sample.</p>
-
-<p>The sample of 34 Warbling Vireos is too small to show the significance, if
-any, of the 2:1 ratio of males to females in the sample. Adequate samples of
-this species, taken at intervals, would add interesting information on time of
-migration of the four sex- and age-classes.</p>
-
-<p><i>Mniotilta varia.</i> Black and White Warbler.&mdash;Weights: 1 adult male, 12.5,
-fat; 2 adult females, 10.0, 10.0, little fat, fat.</p>
-
-<p><i>Vermivora peregrina.</i> Tennessee Warbler.&mdash;Weights: 1 adult male, 10.9,
-very fat; 1 immature male, 12.9, very fat; 2 adult females, 9.1, 12.5, moderately
-fat and very fat. The relative scarcity of Tennessee Warblers in the
-sample is surprising. They are common in the area in spring.</p>
-
-<p><i>Vermivora c. celata.</i> Orange-crowned Warbler.&mdash;Weights: 9 adult males,
-mean 8.8 (7.7-10.9), little fat to fat; 13 immature males, mean 8.92 &plusmn; .15,
-S. D. .53, little fat to fat; 5 adult females, mean 8.8 (8.3-10.3), little fat to
-moderately fat; 17 immature females, mean 9.13 &plusmn; .08, S. D. .72, little fat to
-fat. Of the 19 Orange-crowned Warblers killed on October 7, 11 had little
-fat, 6 were moderately fat, and only 2 were fat. No one-night sample of any
-other warbler killed at Topeka had less fat than this group of warblers. Furthermore,
-our sample (including 11 males) from October 7 (all sex- and age-classes)
-averaged 8.81 grams; the sample of 13 (including only 4 males) from
-October 1 averaged 9.1 grams. If one can assume, for any one species, that
-individuals undertake nocturnal migration only when they are physiologically
-ready, and this includes a certain amount of fat as a fuel source (Wolfson,
-1954), then this further assumption seems justified: birds killed in migration
-with little fat must have flown longer or farther or both than birds killed with
-more fat. No further speculation on this point is permissible with our data,
-but the possibilities for study of future large kills, especially where actual time
-of death of the birds is known, are obvious.</p>
-
-<p><i>Vermivora r. ruficapilla.</i> Nashville Warbler.&mdash;More Nashville Warblers
-were picked up at Topeka than any other species and they are discussed in
-detail elsewhere in this report. The four sex- and age-classes can be identified
-with fair accuracy on plumage characteristics alone. Adult males have a
-large amount of reddish-brown in the crown, not completely veiled by the
-gray tips of the crown feathers. Immature males have a smaller but distinct
-crown patch, usually completely veiled. All males, compared with females, are
-grayer on the sides of the head, have a more nearly white eye-ring, and show
-clearer yellow on the throat. Adult females differ from immature females in
-that they more often have a trace of rufous in the crown and tend to be
-brighter below than the immatures. Of 177 specimens, 20 were very fat, 108
-were fat, 46 were moderately fat, and 3 had little fat.</p>
-
-<p><i>Parula americana.</i> Parula Warbler.&mdash;Weight: 1 adult female, 7.9, fat.</p>
-
-<p><i>Dendroica petechia aestiva.</i> Yellow Warbler.&mdash;Weights: 1 immature male,
-<span class="pagenum"><a name="Page_14" id="Page_14">[Pg 14]</a></span>10.2, fat; 3 adult females, 8.8, 9.5, 10.1, moderately fat; 2 immature females,
-9.0, 9.4, little fat and fat.</p>
-
-<p><i>Dendroica magnolia.</i> Magnolia Warbler.&mdash;Weights: 1 adult female, 9.0,
-moderately fat; 2 immature females, 7.9, 10.3, moderately fat and fat.</p>
-
-<p><i>Dendroica c. caerulescens.</i> Black-throated Blue Warbler.&mdash;Weights: 2
-immature males, 13.8, 14.1, excessively fat; 1 immature female, 11.4, fat.
-This species is rare in Kansas. Although its breeding range is almost entirely
-east and north of Kansas, records in files at the University of Kansas show
-that more specimens have been taken in western than in eastern Kansas.</p>
-
-<p><i>Dendroica c. coronata.</i> Myrtle Warbler.&mdash;Weight: 1 immature female,
-11.6, fat.</p>
-
-<p><i>Dendroica pensylvanica.</i> Chestnut-sided Warbler.&mdash;Weights: 2 immature
-females, 8.1, 10.0, little fat. Only one specimen from Kansas had been preserved
-previously although the species is a regular transient in small numbers
-throughout the state.</p>
-
-<p><i>Dendroica castanea.</i> Bay-breasted Warbler.&mdash;Weights: 1 adult male, 19.2,
-excessively fat; 1 adult female, 11.7, little fat; 1 immature female, 11.2, moderately
-fat. Only 5 specimens of this warbler have been taken previously in
-Kansas, 4 in spring (Ruth, 1952:18-19) and 1 in fall.</p>
-
-<p><i>Dendroica p. palmarum.</i> Palm Warbler.&mdash;Weights: 2 immature males,
-9.9, 10.9, moderately fat; 2 unsexed immatures, 9.1, 9.4, moderately fat. This
-species has been taken in fall in Kansas only once before (KU 26353, taken
-by Wetmore, at Lawrence, on October 5, 1907), but probably occurs regularly
-in both spring and fall migration.</p>
-
-<p><i>Seiurus a. aurocapillus.</i> Oven-bird.&mdash;Weights: 2 adult males, 22.5, 23.8,
-fat and very fat; 14 immature males, mean 21.89 &plusmn; .66, S. D. 2.46, fat to very
-fat; 8 adult females, mean 21.4 (18.3-25.7), moderately fat to fat; 6 immature
-females, mean 18.2 (15.6-20.0), moderately fat to fat.</p>
-
-<p><i>Seiurus noveboracensis notabilis.</i> Northern Water-thrush.&mdash;Weights: 3
-immature males, 18.1, 18.6, 22.2, moderately fat to fat; 1 immature female,
-22.2, fat. Referring these birds to <i>notabilis</i> is a somewhat arbitrary procedure.
-They display some intermediacy of characters and probably stem from a
-population, intermediate between <i>notabilis</i> and <i>noveboracensis</i>, occupying
-much of central North America (cf. McCabe and Miller, 1933).</p>
-
-<p><i>Oporornis philadelphia.</i> Mourning Warbler.&mdash;Weight data presented elsewhere.
-The birds killed at Topeka provide the latest fall dates for this species
-in Kansas. Fifteen were killed on September 25, 64 on October 1, 2 on
-October 5, 11 on October 6, 2 on October 7, and 1 on October 8. We find
-no other records later than September 15. Of 93 specimens examined, 1 was
-excessively fat, 22 were very fat, 45 were fat, 21 were moderately fat, and 4
-had little fat. The abundance of this secretive species in the sample was a
-great surprise. It had previously been considered a rather rare migrant in
-this area.</p>
-
-<p><i>Geothlypis trichas occidentalis [&gt;brachidactyla?].</i> Yellow-throat.&mdash;Weight
-data presented elsewhere. This species was second in numbers only to the
-Nashville Warbler in the total kill at Topeka. Of 167 birds examined, 29
-were very fat, 114 were fat, 23 were moderately fat, and 1 had little fat.</p>
-
-<p>The Yellow-throats are greatly in need of meaningful and comprehensive
-revision, which to date has been restricted to the western subspecies (Behle,
-1950). Since the appearance of the 1931 A. O. U. Check-List a great deal
-<span class="pagenum"><a name="Page_15" id="Page_15">[Pg 15]</a></span>of scattered taxonomic work on the species, as yet unsynthesized, has made
-the picture of its geographic variation a blurry one so far as the details are
-concerned. Made in the absence of adequate comparative material, the above
-identification is to be regarded as tentative. Also, it is, unfortunately, based
-only on those 6 of our 176 specimens preserved as skins. Five of these are
-adult males, the sixth being an immature female. Compared with a series of
-Kentucky specimens regarded as typical <i>brachidactyla</i>, these birds are paler
-and brighter above (tending toward gray-green rather than brownish olive),
-brighter and more extensively yellow below, with broader, more nearly white
-superciliary stripes above their black masks (in males). In size they are
-close to <i>occidentalis</i> (see Behle, 1950:202). Five males have an average
-wing-length of 56.6 mm. (53-59); one female measures 53. Six males from
-Kentucky: 55.1 (53-56); four females, 51.1 (48-56). Our birds may be assumed
-to have stemmed from a population to the north and west which, if
-not <i>occidentalis</i> (or <i>campicola</i> Behle and Aldrich, of which no comparative
-material is at hand), is intermediate between <i>brachidactyla</i> and more western
-birds. Judging from Behle's map (1950:fig. 32), these birds may have come
-from an area near the confluence of three subspecies (<i>campicola</i>, <i>occidentalis</i>,
-<i>brachidactyla</i>). Long (1940:452) reports three subspecies breeding in Kansas
-(<i>brachidactyla</i>, northeast; <i>occidentalis</i>, west; <i>trichas</i>, southeast). The occurrence
-in Kansas of <i>G. t. trichas</i> as currently understood is completely out of
-the question.</p>
-
-<p><i>Icteria v. virens.</i> Yellow-breasted Chat.&mdash;Weight: 1 unsexed immature,
-29.7, moderately fat.</p>
-
-<p><i>Wilsonia p. pusilla.</i> Wilson Warbler.&mdash;Weights: 2 adult females, 7.5, 7.8,
-fat, moderately fat; 1 unsexed adult, 8.3, fat.</p>
-
-<p><i>Wilsonia canadensis.</i> Canada Warbler.&mdash;Weight: 1 immature female, 10.0,
-little fat. We know of only five other specimens from Kansas, although this
-warbler seems to be a regular migrant in small numbers in the state.</p>
-
-<p><i>Setophaga r. ruticilla.</i> American Redstart.&mdash;Weight: 1 immature female,
-9.1, moderately fat.</p>
-
-<p><i>Dolichonyx oryzivorus.</i> Bobolink.&mdash;Weights: 2 adult females, 39.5, 42.9;
-2 immature females, 38.8, 42.0; all excessively fat. Specimens of the Bobolink
-previously have been taken in fall in Kansas only on September 20 and 24,
-1933, near Lawrence, by Long and Preble (Long, 1934).</p>
-
-<p><i>Pheucticus ludovicianus.</i> Rose-breasted Grosbeak.&mdash;Weights: 1 adult male,
-50.4, fat; one immature male, 54.5, very fat.</p>
-
-<p><i>Passerina cyanea.</i> Indigo Bunting.&mdash;Weights: 1 adult male, 18.4, fat; 2
-immature males, 17.2, 17.2, fat and very fat; 2 adult females, 14.3, 16.9, moderately
-fat and very fat; 1 immature female, 13.4, little fat. The sample was
-carefully checked for Lazuli Buntings (<i>Passerina amoena</i>); none was found.</p>
-
-<p><i>Spiza americana.</i> Dickcissel.&mdash;Weight data presented elsewhere in this
-paper. Dickcissels were picked up at the television tower on October 1 (31),
-5 (1), 6 (3), and 7 (1). These birds, together with an adult female taken
-3 miles east and 3 miles south of Lawrence, on October 11, 1953, by Tordoff,
-are the only specimens of this species taken as late as October in Kansas. The
-Dickcissel becomes inconspicuous in late summer and many observers here and
-elsewhere have thought the species disappeared much earlier than it really
-does (see Ganier, 1949). Of 34 specimens, 20 were very fat and 14 were fat.</p>
-
-<p><span class="pagenum"><a name="Page_16" id="Page_16">[Pg 16]</a></span></p>
-
-<p><i>Passerculus sandwichensis nevadensis.</i> Savannah Sparrow.&mdash;Weights: 1
-adult male, 19.4, fat; 2 immature males, 18.3, 19.0, moderately fat; 5 adult
-females, mean 17.2 (14.8-19.5), little fat to fat; 4 immature females, mean
-18.0 (16.9-19.6), moderately fat to fat. Many of the Savannah Sparrows migrating
-through Kansas have in the past been referred to the subspecies <i>P. s.
-anthinus</i> (= <i>alaudinus</i> of the 1931 A. O. U. Check-List) by various workers
-(see Long, 1940:454). As Peters and Griscom (1938:464-5) have shown,
-true <i>anthinus</i>, breeding in the far northwest, ordinarily occurs in migration
-only in the western part of the country, the breeding Savannah Sparrows of
-a large part of the central continental region (east to southern Wisconsin)
-being <i>P. s. nevadensis</i> as now understood. Migrants of this pale, clay-colored
-subspecies should be abundant in Kansas, and all of the specimens in the
-present sample are referable to it.</p>
-
-<p><i>Ammodramus savannarum perpallidus.</i> Grasshopper Sparrow.&mdash;Weights:
-3 adult males, 16.4, 17.6, 20.6, moderately fat, fat, fat; 5 immature males,
-mean 18.1 (16.0-20.2), little fat to fat; 5 adult females, mean 17.9 (16.8-18.9),
-moderately fat to very fat; 5 immature females, mean 18.1 (16.8-20.6),
-fat to very fat.</p>
-
-<p><i>Passerherbulus caudacutus.</i> Leconte Sparrow.&mdash;Weights: 1 immature male,
-11.2, moderately fat; 1 immature female, 12.2, moderately fat.</p>
-
-<p><i>Ammospiza caudacuta nelsoni.</i> Sharp-tailed Sparrow.&mdash;Weights: 2 adult
-males, 15.2, 17.1, moderately fat and very fat; 1 adult female, 13.3, little fat.
-Five specimens of this species have been taken previously in Kansas, all in
-October in the eastern part of the state. Additionally, several observers have
-reported birds seen but not collected. The three birds from Topeka were
-picked up on October 6, 7, and 10 and are the only specimens taken since
-1907. Possibly our specimens from Topeka struck the tower on the same night.
-Tordoff noticed, upon preparation, that the specimens from October 7 and 10
-showed progressive drying of the extremities and spoilage as compared with the
-bird picked up on October 6.</p>
-
-<p><i>Junco hyemalis cismontanus.</i> Slate-colored Junco.&mdash;Weight: 1 immature
-female, 16.4, little fat. Juncos of hybrid type, whether <i>J. h. hyemalis</i> &times; <i>J.
-oreganus</i> subsp. or true <i>J. h. cismontanus</i>, are fairly common in eastern Kansas.</p>
-
-<p><i>Spizella pallida.</i> Clay-colored Sparrow.&mdash;Weights: 2 adult males, 11.6,
-12.2, both fat; 1 immature male, 11.8, fat; 1 adult female, 12.5, fat; 7 immature
-females, mean 11.1 (9.7-12.5), little fat to fat.</p>
-
-<p><i>Passerella iliaca iliaca.</i> Fox Sparrow.&mdash;Weight: 1 adult female, 29.4, little
-fat. A trifle grayer above than any of several Kentucky specimens, this bird
-nevertheless seems well within the range of variation of <i>iliaca</i>.</p>
-
-<p><i>Melospiza l. lincolnii.</i> Lincoln Sparrow.&mdash;Weights and measurements are
-discussed elsewhere. Of 81 specimens, 15 were very fat, 47 were fat, 12 were
-moderately fat, and 7 had little fat. Interestingly, there is no evidence that
-the large southern montane subspecies (<i>M. l. alticola</i>) has contributed to the
-present sample. No bimodality is evident in the curve of wing-length in our
-birds, the largest of which barely approach, the small extreme recorded for
-<i>alticola</i> by Miller and McCabe (1935:156).</p>
-
-<p><i>Melospiza georgiana ericrypta.</i> Swamp Sparrow.&mdash;Weights: 3 immature
-females, 14.3, little fat, 16.7, 17.0, moderately fat. Swamp Sparrows examined
-were all more or less brightly colored and seem to belong to this northern subspecies.</p>
-
-<p><i>Melospiza melodia juddi.</i> Song Sparrow.&mdash;Weights: 1 adult female, 19.4,
-<span class="pagenum"><a name="Page_17" id="Page_17">[Pg 17]</a></span>little fat; 1 unsexed immature, 16.0, little fat. A large proportion of the migrant
-and wintering Song Sparrows in eastern Kansas probably originate from
-the range of this subspecies in the northern plains. <i>Melospiza melodia euphonia.</i>&mdash;One
-immature female (not weighed) was picked up below the tower
-on October 27, 1954, and thus does not appear in Table 1. The specimen
-proved typical of this generally more eastern subspecies upon comparison with
-a large series from Kentucky. For what it may be worth we refer the single
-specimen to this subspecies. Long (1940:456) reported two eastern subspecies
-from Kansas ("<i>beata</i>," <i>melodia</i>). All Kansas specimens genuinely of
-eastern origin probably originate from the range of <i>euphonia</i>, as now understood.</p></blockquote>
-
-
-
-<hr class="chap" />
-<h2>Randomness of the Sample</h2>
-
-
-<p>The reliability of certain of the conclusions which might be
-drawn from data of the kind presented herein depends largely on
-the randomness of the sample. To what degree does this sample
-provide a true cross-section of the nocturnal migrants present over
-the area on a given night or succession of nights? As far as the
-relative abundance of species in the sample is concerned, there is
-little doubt that it is not at all random. The absence of such species
-as the Gray-cheeked Thrush (<i>Hylocichla minima</i>), among the
-passerines, and many of the shorebirds known to be migrating
-through the area at the time is evidence for this statement. Quite
-possibly many seminocturnal species did not strike the tower at all
-for the simple reason that they could see it, and certain large-eyed
-diurnal species (such as thrushes and shorebirds) may avoid collision
-to some extent, thus not appearing in the sample in proportion
-to their actual numbers. Finally, some or all of the species concerned
-probably migrate partly by day. The sample may to some
-degree reflect the true relative abundance of closely related species.
-For example, there is little doubt that, as shown by the sample,
-Nashville Warblers are more numerous locally at this season than
-Tennessee Warblers, a fact that can readily be corroborated by
-ordinary field observation. Also, the sample is useful in suggesting
-the actual abundance of species which are furtive and/or difficult
-to identify under normal field conditions, for example, the Mourning
-Warbler and Philadelphia Vireo. It is obvious that the sample
-should reflect the true relative abundance at one place and time
-of any two species with equal tendency to migrate by night and
-equal tendency to strike the tower. Since the facts in regard to
-both tendencies are at present unknown for most species, we think
-that interspecific comparisons should be avoided or approached
-with extreme caution.</p>
-
-<p>In respect to the relative abundance of the various sex- and
-age-<span class="pagenum"><a name="Page_18" id="Page_18">[Pg 18]</a></span>classes within a given species, the sample is, we think, as close to
-random as is possible to obtain. Certainly it is greatly superior to
-samples obtained by field collecting, where possible differences in
-habits, wariness, and experience of the birds, and subconscious (if
-not conscious) selection by collectors can all bias the results.
-Dwight (1900:128-9) believed that the greater wariness of adult
-birds was almost entirely responsible for the seemingly disproportionate
-number of immatures in autumn and gave some observational
-evidence in favor of his views. The large percentage of
-adults in some of the samples here treated tends to reinforce
-Dwight's position. To a somewhat lesser extent, this advantage in
-randomness of accidental kills over routine collecting may be supposed
-to apply also in demonstrating the composition by subspecies
-of a single migrant species.</p>
-
-<p>So far as particulars already mentioned are concerned, the present
-sample or other samples of tower-killed birds would seem to be in
-no way superior (that is, more nearly random) to samples obtained
-in connection with lighthouses and other lighted objects, and
-ceilometers. In one important respect, however, it is probably
-somewhat superior to these as the dimly red-lighted structure has
-not been shown to have any important collecting or attracting influence.
-Thus, in computations intended to estimate the over-all
-abundance of migrants, the sample should be more reliable than
-samples involving bright light with its possible attracting effect.</p>
-
-
-
-<hr class="chap" />
-<h2>Number of Migrants</h2>
-
-
-<p>If it can be assumed that nocturnally migrating birds are approximately
-uniformly spaced across the sky and that the red lights did
-not attract birds which would otherwise have missed the tower, it
-is possible to compute the volume of migration from the sample
-killed. In regard to the first assumption, both Stone (1906:250-251)
-and Lowery (1951:409-413) have presented evidence of fairly uniform
-distribution of nocturnal migrants. We have no information
-on the second assumption beyond the facts that birds do not strike
-the high towers on clear nights or lower towers even on stormy
-nights.</p>
-
-<p>On nights when large numbers of birds struck the 950 foot Topeka
-tower, only a few struck a 500 foot radio tower, also lighted
-with red lights, at Lawrence, 24 miles east, under similar weather
-conditions. Most of the birds found at Topeka were fairly close
-to the base of the tower, indicating that they struck the tower itself<span class="pagenum"><a name="Page_19" id="Page_19">[Pg 19]</a></span>
-or that they were flying high enough to strike guy wires only fairly
-close to the tower. The scarcity of birds under the guy wires some
-distance from the tower at Topeka and at the radio tower at Lawrence
-causes us to think that most of the birds were flying more
-than 450 feet above the ground. On this basis, we have computed
-numbers of migrants passing through a plane one mile long and
-500 feet high (2,640,000 square feet), intersecting the assumed path
-of migration at right angles. Vertically, the theoretical plane begins
-at 450 feet above ground and has its top edge at 950 feet. The
-solid (discounting spaces between girders, <i>etc.</i>) cross-sectional area
-of the tower intersecting this plane was computed by actual measurement
-to be 1685 square feet. On the night of September 30-October
-1, 585 birds were killed. By computation (585/1685 =
-X/2,640,000), approximately 916,000 birds passed through the mile-long
-plane that night. On each of the nights of October 5-6 and
-October 6-7, approximately 230,000 birds passed through this plane.
-By comparison, Lowery (1951:436) recorded maximum station
-densities in one night in spring of 63,600 birds at Tampico, Mexico,
-and 51,600 at Lawrence, Kansas, as determined by moon-watching.
-Lowery's figures refer to numbers of birds crossing any part of a
-circle one mile in diameter and are roughly comparable to ours if,
-as we think, most of the birds at Topeka were flying at altitudes between
-450 and 950 feet above the ground.</p>
-
-<p>It must be realized that these figures are only approximations.
-One variable ignored is the frontal extent (or area, viewed from the
-front, subject to damage by striking an obstruction) of the birds
-themselves. Since practically all birds killed showed head or trunk
-injuries, rather than a high proportion with only broken wings, we
-chose to disregard frontal extent of the birds in our calculations.
-If our figures are inaccurate by as much as 50 per cent in either
-direction, which seems unlikely to us, they still give some idea of
-the tremendous volume of nocturnal migration under some conditions.</p>
-
-<p>It may be more meaningful to compute numbers of migrants by
-species. This can be done readily by making appropriate substitutions
-from Table 1 in the equation given above. For example,
-on the night of September 30-October 1, approximately 147,000
-Nashville Warblers passed through the mile-long plane and on the
-same night, 100,000 Mourning Warblers and 14,000 Philadelphia
-Vireos. Neither of the last two species would be judged to be
-abundant migrants in autumn in eastern Kansas by ordinary field<span class="pagenum"><a name="Page_20" id="Page_20">[Pg 20]</a></span>
-observations; the television tower sample, however, indicates that
-these as well as other species must often be overlooked when they
-do stop in Kansas.</p>
-
-
-
-<hr class="chap" />
-<h2>Differential Migration of Sex- and Age-classes</h2>
-
-
-<p><span class="smcap">History of the Subject.</span>&mdash;For a long time it has been known in
-a general way that old and young birds and males and females of
-some species do not always migrate at the same times, by the same
-routes, or even to the same places. This is a subject about which
-much has been written. Reading the summaries of some general
-texts, it is easy to acquire the impression that the facts of the matter
-are well known. On the contrary, they are poorly known and much
-remains to be learned before differential migration is understood.
-This can best be indicated by a brief survey of the literature.</p>
-
-<p>The importance of the subject was emphasized by Meinertzhagen
-(1930:52) in one of the later reviews of differential migration:
-"The main problem concerns the Cause of Migration, the
-Stimulus which compels Migration and the Origin of the Migratory
-Habit.... There is, however, a minor problem which affords
-valuable evidence in helping us to solve the major problem, bearing
-very directly on it, namely, the order of sex and age on migration."</p>
-
-<p>The mystery of how birds, especially the young, find their way
-in migration has fascinated students since the earliest times. The
-quite natural though purely anthropomorphic conclusion of early
-scholars was that the old birds led the young on migration. This
-attractive idea persisted long after ornithology began to grow into
-a science. The classic theory was restated by Palm&eacute;n (1876:267),
-in one of the first thorough reviews of the subject of migration, as
-follows: "Directe Beobachtungen in der Natur ergeben, dass die
-Schaaren von ziehenden V&ouml;geln allgemein &auml;ltere und st&auml;rkere Individuen
-als Anf&uuml;hrer des Zuges haben." Variously modified, this
-view continued to crop up for some time and still found support in
-the 1890's (see Dixon, 1892:69). G&auml;tke (1895:101) correctly questioned
-the credibility of Palm&eacute;n's "direct observations."</p>
-
-<p>With the gradual abandonment of the unsupportable classic
-theory, diametrically opposed views were adopted by workers on
-opposite sides of the Atlantic. The American stand was ably expressed
-by Brewster (1886), who went to great pains to state his
-case and give evidence, and who was later supported by Allen
-(1896:144-147; 1909:17). The Americans held that adult birds
-nearly always preceded the young in migration, and this was based
-on much evidence, whether or not correctly interpreted. Dwight<span class="pagenum"><a name="Page_21" id="Page_21">[Pg 21]</a></span>
-(1900:127) also gave evidence in favor of this theory. Equally
-definite, if, as has later been shown, somewhat vaguely documented,
-was the famous work of G&auml;tke (1895:see pp. 100-113), who after
-many years' observation of migrant birds in Heligoland concluded
-the exact opposite, that young in general precede adults (see critiques
-of Allen, 1896:144-147; Wiegold, 1926:5). G&auml;tke's dissenting
-opinion was for a time supported enthusiastically by British
-workers (Gurney, 1923:579-580).</p>
-
-<p>As so often happens, neither extreme has withstood the test of
-time, and more recent summaries (Meinertzhagen, 1930:55-56;
-Thomson, 1926, 1936:488-489; Wiegold, 1926) have tended to compromise.
-Many exceptions to G&auml;tke's extreme conclusion have been
-detected. Exceptions to the Brewster-Allen stand have also been
-discovered, although work along these lines on the American side
-has lagged somewhat. Rowan (1926) has given further evidence
-on the migration of certain shorebirds, and some evidence has accrued
-in relation to particular species and groups as a result of
-life-history and banding studies (see Pitelka, 1946). Authors of
-major works on migration, however, have either been preoccupied
-with other phases of migration or avoided the issue. In an able
-study (one of several on related subjects) of the composition by sex
-and age of migrant populations in north Germany, Drost (1935:177)
-did not go into the question of order on migration.</p>
-
-<p>One is left with the impression that the whole subject is still unsettled.
-While earlier workers sought to reduce the entire matter
-to law, the latest disclaim the possibility of generalization. After
-summarizing Brewster's and G&auml;tke's opinions, Thomson (1926:79)
-wrote: "It would seem, in any event, that no general rule can be
-laid down." Meinertzhagen's summation (1930:56-57) still represents
-fairly well the status of our knowledge: "Order of sex and
-age on autumn passage is very difficult to arrive at, as evidence is
-conflicting. But, on the whole, it seems that birds flock together,
-old and young, preparatory to moving south, and do in many cases
-initiate migration in company.... But once movement is initiated,
-among birds which do not habitually fraternise in flocks,
-adults, and especially males, will naturally outstrip the less virile
-females and still less virile offspring.... The consequence is
-that any observer at an intermediate station such as Heligoland is,
-in noting birds of the year as first arrivals, has not had an
-opportunity of noting the flocks of adults which have passed without
-alighting. On the other hand, there is very definite evidence to
-show that among certain species, adults follow their offspring on<span class="pagenum"><a name="Page_22" id="Page_22">[Pg 22]</a></span>
-migration. The reason for different behaviour among different types
-of birds remains obscure." We regard much of this as still theory.</p>
-
-<div class="figcenter" style="width: 400px;">
-<img src="images/fig1.jpg" width="400" height="531" alt="" />
-<span class="caption"><span class="smcap">Fig. 1.</span> Composition by age and sex as found in one or more series of
-each of eight species of birds included in the Topeka sample. Each separate
-series is represented by a single histogram, the histograms for a species
-being grouped with the earliest series on the left. Each histogram expresses
-the numbers of adults (left-hand column) and immatures (right-hand
-column) in terms of percentage of the whole series. Thus the two bars
-of each couplet add up to 100 per cent. The hatched portion of each bar
-represents males, the clear portion females.</span>
-</div>
-
-<p>It would be difficult to imagine a better way of resolving the
-problems and uncertainties just reviewed than by the detailed analy<span class="pagenum"><a name="Page_23" id="Page_23">[Pg 23]</a></span>sis
-of large samples of migratory birds killed at random at various
-points and times. An analysis of the sample of birds accidentally
-killed at Topeka is presented here as an initial step in this direction.
-Although the conclusions based on this sample are tentative and
-may in time be altered, the data themselves are definite. If this
-general type of analysis is repeated again and again&mdash;applied to
-samples taken on many dates and in many localities&mdash;a mass of
-hitherto unparalleled evidence for the study of migration will
-emerge.</p>
-
-
-<p><span class="smcap">Differential Migration OF Sex- and Age-classes as shown by
-the Topeka Sample.</span>&mdash;Smaller samples have not been treated.
-Species affording samples seemingly large enough to justify at least
-preliminary analysis were: Catbird, Red-eyed Vireo, Mourning
-Warbler, Dickcissel, Nashville Warbler, Orange-crowned Warbler,
-Yellow-throat, and Lincoln Sparrow (Fig. 1). For all of these except
-the Catbird and Dickcissel, at least two samples from a week
-or more apart were available for comparison in an effort to detect
-trends in migration. Fig. 1 shows the actual ratios of sex- and age-classes
-observed in samples of the species listed above. Each of
-the last four species provided two separate samples, of sufficient
-size to warrant an attempt at measuring the statistical significance
-of the observed changes in adult-immature ratios (Table 2).</p>
-
-<p class="center"><span class="smcap">Table 2.&mdash;Statistics of the Ratios of Adults to
-Immatures in Four Species</span></p>
-
-
-<div class="center">
-<table border="1" cellpadding="4" cellspacing="0" summary="Table 2">
-<tr><td>Species</td><td>Dates of samples</td><td>Total number</td>
- <td>Number and percentage of adults<a name="FNanchor_1_1" id="FNanchor_1_1"></a><a href="#Footnote_1_1" class="fnanchor">[1]</a></td><td>Difference (in %)<a name="FNanchor_2_2" id="FNanchor_2_2"></a><a href="#Footnote_2_2" class="fnanchor">[2]</a></td><td>P<a name="FNanchor_3_3" id="FNanchor_3_3"></a><a href="#Footnote_3_3" class="fnanchor">[3]</a></td></tr>
-<tr><td align="left">Nashville Warbler</td><td>Oct. 1 (93)<br />Oct. 5-7 (63)</td><td>156</td>
- <td>45 (.484)<br />26 (.413)</td><td>.071</td><td>.36</td></tr>
-<tr><td align="left">Orange-crowned Warbler</td><td>Sept. 25-Oct. 1 (19)<br />Oct. 5-9 (25)</td><td>44</td>
- <td>3 (.158)<br />11 (.440)</td><td>.282</td><td>.05</td></tr>
-<tr><td align="left">Yellow-throat</td><td>Oct. 1 (115)<br />Oct. 5-8 (44)</td><td>159</td>
- <td>62 (.540)<br />20 (.455)</td><td>.085</td><td>.34</td></tr>
-<tr><td align="left">Lincoln Sparrow</td><td>Oct. 1-3 (44)<br />Oct. 6-10 (27)</td><td>71</td>
- <td>27 (.614)<br />8 (.296)</td><td>.318</td><td>.01</td></tr>
-</table></div>
-
-<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> Percentage of immatures equals 1.000 minus percentage of adults.</p></div>
-
-<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> Standard error of the difference between ratios was computed by the
-formula</p>
-
-
-<div class="center">
-<table border="0" cellpadding="4" cellspacing="0" summary="formula">
-<tr><td rowspan="2">&#963;<sup>e</sup> =</td>
- <td rowspan="2" align="right"><span style='font-size:300%;font-weight:lighter;margin:0;line-height:1em;text-indent:0;'>&#8730;</span></td>
- <td rowspan="2" class="bor_top">P<sub>e</sub> Q<sub>e</sub> (</td><td class="bor_top">1</td><td rowspan="2" class="bor_top">+</td><td class="bor_top">1</td><td rowspan="2" class="bor_top">)</td>
- <td rowspan="2">,</td></tr>
-<tr><td class="bor_top">N<sub>1</sub></td><td class="bor_top">N<sub>2</sub></td></tr>
-</table></div>
-
-<p>where P<sub>e</sub> equals percentage of adults and Q<sub>e</sub>
-equals percentage of immatures in the entire sample.</p></div>
-
-<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> Probability of error; <i>i. e.</i>, a P of .01 means there is one chance in 100
-that the difference observed does not represent an actual difference in nature.</p></div>
-
-<p><span class="pagenum"><a name="Page_24" id="Page_24">[Pg 24]</a></span></p>
-
-<p>Upon the application of statistical methods it soon became evident
-that, unless changes in ratio between two samples are marked,
-large samples would be required in order to reach conclusions of
-high statistical significance in a single study of the present type. In
-this case (see Table 2), the Lincoln Sparrow and Orange-crowned
-Warbler, though represented by only moderate-sized series, show
-marked changes in age composition over the period studied, and
-the statistical treatment indicates a high degree of probability that
-these changes are real. Assurance that the lesser changes observed
-in the Nashville Warbler and Yellow-throat are real, on the other
-hand, is much less, even though the samples are larger. Few if any
-of the samples here discussed are as large as might be desired.
-Therefore, conclusions based upon them (see below) are to be regarded
-as tentative. Many other, future, samples will perhaps also
-be insufficient in size in themselves. There are, however, statistical
-advantages to repetition which will serve to make the repeated
-analysis even of small samples significant and valuable.</p>
-
-<p>Certain of the samples not treated statistically show ratios that
-can be seen by inspection to be probably significant. For example
-the almost complete absence of adults from the three samples of
-Red-eyed Vireos (Fig. 1E) cannot be disregarded in view of the
-size of the whole sample of the species. The same applies to the
-high percentage of adult females and the near absence of adult
-males in the sample of the Dickcissel (Fig. 1F). The continuity
-in direction of changes observed in the three samples of the Mourning
-Warbler (Fig. 1G) and Red-eyed Vireo is likewise probably
-significant, even though some of the samples compared are small.
-It seems to us that the application of statistical methods to these
-species should await the accumulation of more material. For anyone
-desiring to treat them statistically now, the data are inherent
-in this paper.</p>
-
-<p>We have not computed the standard errors of the ratios of sexes
-within age groups (except experimentally in a few cases). This
-can easily be done, however, and the significance of a given ratio
-determined, on the assumption (perhaps sometimes dubiously justifiable)
-that the sex-ratio in the species concerned is one:one. Obviously
-there is no point in computation of the standard errors of
-adult-immature ratios in single samples (such as that of the Dickcissel)
-until the actual ratio prevailing in the species in nature at
-the season in question is known for comparison with the observed
-ratio. Our formal statistical treatment, therefore, has been limited<span class="pagenum"><a name="Page_25" id="Page_25">[Pg 25]</a></span>
-to an examination of the significance of the <i>changes</i> between adult-immature
-ratios in samples of the same species taken a number of
-days apart.</p>
-
-<p>The samples suggest several patterns of differential migration of
-sex- and age-classes. Indeed, the important consideration brought
-out&mdash;in our opinion not hitherto sufficiently emphasized in literature&mdash;seems
-to be that in generalizing about adults and immatures,
-one must be careful to take sexes into account, and conversely, in
-generalizing about males and females, one must consider also age.
-In other words, there are really four classes to be considered. This
-poses additional problems in analysis and introduces the need for
-still larger samples in order to reach significant conclusions. To
-illustrate: an adult-immature ratio of 40:20 (N = 60) may be satisfactorily
-significant, while within the 40 adults a ratio of 25
-males:15 females may not be. Were the original sample 80:40
-(N = 120) with male adults 50 and female adults 30, it is obvious
-that the significance of the latter ratio would be greater. The same
-applies in reverse if the greater emphasis is placed on sex and the
-lesser on age. Because of the moderate size of the samples this
-problem has been felt in the present study in respect to sex ratios
-within age groups, many of which must at present be regarded as
-of tentative significance.</p>
-
-<p>In short, what the earlier ornithologists regarded as a simple problem
-is in reality a complex one. There are only two patterns in
-what may be called the Brewster-G&auml;tke argument: adults first or
-immatures first (with of course the further possibility of both at the
-same time). Both patterns occur, as is now known, at least to some
-extent. But actual patterns, as suggested by our samples, are more
-complex when all classes are considered. It will readily be seen
-that, if adult males, immature males, adult females, and immature
-females be regarded as units, each with certain migratory characteristics,
-the combinations of these units in various orders of migratory
-precedence are potentially numerous. In fact, of course,
-they do not behave strictly as units (or perhaps very rarely so),
-but our data strongly indicate that the tendency exists in many
-cases. This may be stated another way. The present samples may
-be reduced to two basic patterns, fitting the classic early American
-(adults first) and early European (immatures first) theories. But,
-either such simple arrangement is compounded in some, perhaps in
-truth in all, instances by differential migration of the sexes <i>within</i>
-each age class. This proposition can also be stated backwards:<span class="pagenum"><a name="Page_26" id="Page_26">[Pg 26]</a></span>
-the samples show differential times of migration of the sexes, compounded
-by differential times of migration of the age groups within
-each sex. The order in which these matters are approached depends
-on what one is trying to find out. Influenced by the literature,
-in which most emphasis has been placed on age, we have approached
-the problem from that standpoint. The data and figures
-here given, however, can be juggled if one wishes to place first
-emphasis on the order of sexes in migration.</p>
-
-<p>Bearing in mind what has just been said, particularly in respect
-to sizes of samples necessary for significance, let us consider the
-patterns of migration suggested by the Topeka sample. These are
-as follows:</p>
-
-<p>(1) <i>An early migration largely composed of adults, giving way
-later on to a preponderance of immatures.</i> Regardless of variations
-among them, samples showing this basic pattern are in line with
-the opinions of Brewster (1886) and his followers. This pattern
-is here shown by the Lincoln Sparrow, Yellow-throat, Nashville
-Warbler, Catbird (one sample only), and Red-eyed Vireo (Fig. 1,
-A, B, C, D, E). The evidence of these and all other samples would
-admittedly be more conclusive if the samples were further apart in
-time or, better still, were there more of them. There is evidence
-that differences in migration of the sexes, within age classes, influence
-this pattern, sharply in some instances. In the later samples
-of Lincoln Sparrow, Yellow-throat, and Red-eyed Vireo (Fig. 1,
-A, B, E) there are relatively fewer males, both adult and immature,
-than in the earlier samples and this may be true also of the Catbird,
-judging from the single sample. The Red-eyed Vireo (Fig.
-1, E) is characterized by small number, or absence of, males in
-each sample but the samples are not significantly different, and can
-be regarded as one. Although the samples of the Dickcissel and
-Mourning Warbler (Fig. 1, F, G) show a somewhat different over-all
-pattern and are discussed further on, they also contain few adult
-males. Since these samples are from a period that is near the end
-of the migration of Red-eyed Vireos, Mourning Warblers, and Dickcissels,
-it may be assumed tentatively that the adult males have
-already migrated. Meinertzhagen (1930:56) postulated that in
-many species there is an earlier or more rapid migration of adults,
-particularly males, and the data for the above species in our sample
-tend to support his assumption. But our data suggest in addition
-that in some species <i>immature males</i> migrate earlier, or more rapidly,
-than do <i>immature females</i>, just as adult males precede adult<span class="pagenum"><a name="Page_27" id="Page_27">[Pg 27]</a></span>
-females in some instances. Within this general pattern (adults
-first) another variation is shown by the Nashville Warbler (Fig. 1,
-C) in which the later sample of adults is heavily weighted towards
-males, even though an increasing over-all proportion of immatures
-is evidenced. In this case, and contrary to Meinertzhagen's suggestion,
-it would seem that adult females have preceded or outstripped
-adult males in migration.</p>
-
-<p>(2) <i>An early preponderance of immatures, followed by a preponderance
-of adults.</i> The several species of birds at Topeka that
-display this pattern conform with the conclusions of G&auml;tke and
-other early Old World ornithologists that in most species immatures
-precede adults in migration. In the present sample two variations
-of this pattern occur.</p>
-
-<p>(a) In the Dickcissel (Fig. 1, F) and the Mourning Warbler
-(Fig. 1, G), immatures decrease more markedly than adults (visible
-in samples of Mourning Warbler; inferred in Dickcissel), leaving
-the adults in the majority in the closing phase of migration. The
-distinctive and interesting feature in each of these two species is
-the ascendancy in numbers of adults <i>despite</i> the almost complete
-disappearance of adult males. The relative increase of adults is
-here caused by a retarded migration of adult females, which linger
-conspicuously behind all other classes. Something of this nature
-was suggested, in theory, by Dixon (1892:70) who thought that
-adult females are delayed by "maternal duties." It was hinted at
-also by Dwight (1900:127) who thought that in some species females
-molted later than males as a result of prolongation of parental
-responsibilities. As mentioned already, there is need for caution
-in interpreting the present samples because the Dickcissel is represented
-only by one sample and two of the three samples of Mourning
-Warblers are small. In the case of the Mourning Warbler, the
-samples may be regarded as one, nearly lacking in adult males.
-The progressive increase of adult females, however, may be significant;
-at least there are enough of these to make division of the
-birds into three samples enlightening. There is, of course, some
-chance that the majority of adult males have not yet migrated, or
-are migrating by a different route. This seems unlikely in both
-cases. October 1 is late in the migration of the Dickcissel and it
-seems that large-scale migration would not occur much later, and
-in the case of the Mourning Warbler adult males are rare in all
-three samples, extending over a considerable period and reaching
-late into the probable migration period of the species. It is
-inter<span class="pagenum"><a name="Page_28" id="Page_28">[Pg 28]</a></span>esting to conjecture just when and where adult male Mourning
-Warblers do migrate in autumn. Brewster (1886:16) wrote: "This
-species arrives at Cambridge [Massachusetts] about September 12,
-and during the remainder of the month is ... abundant....
-The adults, however, are so very uncommon that I have
-never known them [to] represent more than five per cent of the
-total number of individuals. They do not seem to be more numerous
-in the earlier flights than towards the close of the month,
-and I am very sure that they cannot be found in this locality before
-the young begin to appear." While the present samples show an
-abundance of adult <i>females</i> of this species (could Brewster have
-failed to recognize these as adults?) the whereabouts of the adult
-males remains a mystery.</p>
-
-<p>(b) Another variation is displayed by the Orange-crowned Warbler
-(Fig. 1, H). Here also there is an increase of adults towards
-the end of migration, but this increase is marked by a growing percentage
-not of females but of males. Locally this species is a late
-migrant compared with most others of the Parulidae. Thus the first
-sample, composed of birds taken September 25-October 1, may be
-regarded as fairly early in the fall migration. Immature birds compose
-84.2 per cent of this sample, there being no adult males at all.
-By October 5-9 the picture has changed markedly, the sample being
-composed of 44 per cent adults (82 per cent of which, in turn,
-are males) and 56 per cent immatures. In view of this trend one
-can not help suspecting that a still later sample would show a majority
-of adults, perhaps nearly all males. This of course does not
-necessarily follow; the migration of immatures could simply be
-more protracted, and could have commenced earlier, than that of
-adults.</p>
-
-<p>Little imagination is required to see how enlightening it might be
-could we analyze thoroughly the patterns of all migrating species.
-When the detailed facts are available, it seems likely that general
-trends will emerge which may be of great significance to the study
-of migration in general. A final point which must eventually be
-clarified is determination of the extent of variability in the pattern
-of each species from year to year and locality to locality.</p>
-
-<p>Once patterns of precedence in migration of different classes are
-established, search into the life-histories of the species concerned
-may help to explain the peculiarities discovered. In the present
-case, for instance, we find a possible clue to the reason for the high
-proportion of adult females of the Dickcissel late in migration, as<span class="pagenum"><a name="Page_29" id="Page_29">[Pg 29]</a></span>
-shown by our sample. Gross (1921:14-15) presented evidence that
-adult female Dickcissels molt considerably later than their mates,
-and we have independent evidence that individuals of this species
-are at times almost flightless when molting the remiges!</p>
-
-
-
-<hr class="chap" />
-<h2>Molt in Relation to Migration</h2>
-
-
-<p><span class="smcap">General comment.</span>&mdash;The exact relationship between molt and
-migration seems not to have been definitely established. The subject
-has received cursory attention in the literature and conflicting
-opinions have been expressed. Dwight (1900:126-128) believed
-that molt is completed or nearly completed before migration in
-nearly all passerine species that occur in New York save for certain
-swallows and flycatchers. Molt has since been found to precede
-migration of at least one of the flycatchers (<i>Empidonax virescens</i>)
-considered by Dwight to be an exception to this rule (Mengel,
-1952). In Great Britain the subject of molt in migration was considered
-in some detail by Rintoul and Baxter (1914) and Ticehurst
-(1916), who arrived at conclusions similar to Dwight's. These
-workers also found certain swallows to be exceptions to the rule.</p>
-
-<p>The above authors and others have shown that, at least among
-passerines, some body molt is frequently found in migrating individuals
-but that molt of tail feathers is much less often found and
-molt of remiges almost nonexistent. Baxter and Rintoul noted only
-four cases of molting wing feathers among hundreds of migrants.
-Among the diverse non-passerine orders the picture seems to be
-more complicated, as might be expected. We do not, however,
-comprehend the reasoning which led Meinertzhagen (1930:56) to
-summarize: "... on the whole it can be said that though birds
-seldom migrate when flight feathers are in quill, moult in general
-does not influence migration." This seems to us an obvious <i>non
-sequitur</i>. Meinertzhagen (<i>loc. cit.</i>) went on to say: "Males and
-females of one species are believed to moult simultaneously [see,
-however, Dwight, 1900:127], and there is no doubt that in some
-cases the two sexes migrate at slightly different times, and occasionally
-prefer different winter quarters. Birds of the year never
-moult their quills previous to their first autumn migration [Consultation
-of Dwight, 1900, who gives many examples of this, would
-have spared Meinertzhagen this error.], and yet they frequently
-follow adults on passage and sometimes precede them. There are
-no grounds for believing that adults have moulted their quills before
-birds of the year are prepared to migrate [but there are, in
-<span class="pagenum"><a name="Page_30" id="Page_30">[Pg 30]</a></span>many cases; <i>cf.</i> Dwight, 1900:127], in the case where adults precede
-the latter. Neither is there any evidence to show that adults
-have not moulted their quills till after their offspring are ready for
-passage, in the case where they follow their offspring. It does not,
-therefore, appear that moult is an important factor."</p>
-
-<p>Comments interpolated above show our feeling that this summary
-is inadequate and misleading. To us it seems that the extreme
-rarity in migration of birds with remiges in molt is strong
-evidence that molt <i>does</i> influence at least the time of migration. It
-is immaterial whether this molt occurs before or after migration,
-although in the majority of cases it seems to take place before.
-Much more needs to be known of the migration pattern and molt
-of each species before generalizations can safely be made.</p>
-
-<p>Analysis of samples of migrants can show only the presence and
-nature or the absence of molt in birds actually migrating. In the
-present instance shortage of time and manpower for preserving
-some and processing all of the sample resulted in incomplete data
-being kept on molt. We include this section to emphasize uncertainties
-still prevalent and to stimulate further work.</p>
-
-
-<p><span class="smcap">Molt in the Topeka sample.</span>&mdash;Our limited findings coincide with
-those of Rintoul and Baxter (1914). Body molt was noted in a
-number of individuals and species. When present, this molt almost
-invariably was in its final stages. One immature male Rose-breasted
-Grosbeak (October 1) was in heavy body molt. It is perhaps
-worthy of mention here that this grosbeak evidently migrates at
-times in extensive molt. An adult male (RMM 1102) taken by
-Mengel near Henderson, Kentucky, on September 9, 1949, was
-molting plumage of body, wings, and tail, no feather of the last
-being longer than one half inch. This remarkable specimen had
-only five primaries on one side and four on the other fully functional.
-The outermost on the left and two outermost on the right
-were from the previous plumage, not yet dropped; the three innermost
-of each wing were new and full-length.</p>
-
-<p>In the present sample molt of remiges was noted in one specimen,
-an adult female Indigo Bunting (October 1) with outer primaries
-sheathed and with molt in progress in the body plumage. The one
-(immature) Yellow-breasted Chat in the sample (October 1) had
-all of its tail feathers nearly full-length but in quill, possibly as a
-result of accident, and two feathers were being replaced also in the
-tail of an immature Clay-colored Sparrow (October 6), which was
-also in body molt and had some, juvenal feathers on the belly and
-flanks.</p>
-
-<p><span class="pagenum"><a name="Page_31" id="Page_31">[Pg 31]</a></span></p>
-
-<p>Body molt near completion was further noted as follows: immature
-male Yellow-throated Vireo (October 1), adult male Blue-headed
-Vireo (October 1), immature female Leconte Sparrow (October
-23), several Lincoln Sparrows (various dates).</p>
-
-
-
-<hr class="chap" />
-<h2>Size Differences according to Sex and Age</h2>
-
-
-<p><span class="smcap">Linear measurements.</span>&mdash;Taxonomists long have recognized in
-many species that males differ in size from females. Less attention,
-until recently, has been paid to the relative sizes of adult and immature
-birds. Many taxonomists, however, seem to have had an
-uneasy suspicion that immature birds are "untrustworthy" in comparison
-with adults, and immatures have often been excluded from
-samples when recognizable. Since, however, there are still relatively
-few reliably aged specimens in collections, for the most part
-only those immature birds immediately recognizable as such by
-obvious plumage differences (which are often present only in
-juvenal plumage) have been excluded from series. The majority
-of birds in first winter plumage so closely resemble adults that the
-two ages have been included in series for measurement. In most
-passerines these younger birds still bear the juvenal feathers in
-wing and tail and are, in size of these important parts, quite as
-"untrustworthy" as birds still in juvenal body plumage. Even if a
-complete postjuvenal molt occurs we still should not assume that
-first winter feathers are as long as adult winter feathers without
-first determining that this is so. Although aware of this problem,
-systematists until recently seemingly have been more or less content
-to disregard it, or forced to do so for practical reasons. Miller
-(1941:179) had little choice but to hope that size differences between
-adult and immature juncos were unimportant. Behle (1942:217)
-wrote of Horned Larks, <i>Eremophila alpestris</i>: "... the
-plumages of first-year birds and adults seem indistinguishable,
-though I have never quite satisfied myself that there are no differences
-in lengths of rectrices and remiges." He added, with
-logic confusing to us: "Since it is a difficult problem to determine
-the ages of horned larks that have passed the postjuvenal molt, this
-similarity of plumages is fortunate for the systematist."</p>
-
-<p>In recent years, some workers have analyzed size differences between
-adults and immatures. Sibley (1950:115) showed that adult
-Red-eyed Towhees (<i>Pipilo erythrophthalmus</i>) had notably longer
-wings and tails than immatures, and the same was demonstrated in
-Red Crossbills (<i>Loxia curvirostra</i>) by Tordoff (1952). In work
-<span class="pagenum"><a name="Page_32" id="Page_32">[Pg 32]</a></span>with jays (<i>Aphelocoma</i>), Pitelka (1951:199) found that: "...
-in comparisons of dimensions of sex and age groups within a given
-sample, although magnitude of difference varies from one character
-to another, most of the averages are successively smaller for first-year
-males and adult and first-year females." He listed exceptions
-and concluded: "Segregation [of sex and age classes] has proved
-to be of extreme significance in an interpretation of individual and
-geographic variation."</p>
-
-<p>Much along these lines can be learned by examination of large
-random samples such as that afforded by the Topeka accident. Although
-only a few species in this sample were measured, the results
-secured seem to show further the need for segregation of
-age classes in taxonomic work with some species.</p>
-
-<p>Figure 2 shows the variation in the lengths of wing and tail in
-the Nashville Warbler. It is evident from the figure that in both
-of these characters the four sex- and age-classes differ significantly.
-An accurate idea of geographic variation in this species could not
-be obtained without separating these classes in comparisons. Age
-classes in spring and summer, long after the skull is completely
-ossified, can be segregated only if differences in plumage can be
-found. In the Nashville Warbler, such differences occur in fall
-(see annotated list) but these probably are obliterated by the partial
-prenuptial molt. These facts emphasize the importance, for
-taxonomic studies, of aged specimens collected in late summer or
-early fall on their breeding ground and in fresh winter plumage.
-Figure 3 shows the variation in length of wing in the Lincoln Sparrow
-in which age seems to be of much less importance than in the
-Nashville Warbler. Males and females of the Lincoln Sparrow
-differ significantly in wing-length, but adults and immatures are
-of nearly the same size. It would seemingly not be necessary to
-separate age classes in studies of geographic variation in the Lincoln
-Sparrow. Size data for some other species are given in the
-annotated list.</p>
-
-
-<p><span class="smcap">Weights.</span>&mdash;Little seems to have been done to determine the correlation
-of weights with sex- and age-classes. Weight may be the
-best single index of over-all size and is especially useful to students
-of the physiology of migration. Weight, however, is subject to the
-considerable variable imposed by fat condition. In large and comparable
-series, this variable is probably removed insofar as comparability
-of means is concerned, but the high variability of weight
-in most cases naturally results in more overlap (less separability)
-between populations with means close together than is found with
-most linear measurements.</p>
-
-<p><span class="pagenum"><a name="Page_33" id="Page_33">[Pg 33]</a></span></p>
-
-<div class="figcenter" style="width: 400px;">
-<img src="images/fig2.jpg" width="400" height="707" alt="" />
-<span class="caption"><span class="smcap">Fig. 2.</span> Statistics of variation in length of wing and tail in the Nashville
-Warbler. The solid lines represent the observed ranges in millimeters.
-The stippled boxes represent two standard errors (&#963;m) to each side of
-the means (vertical lines). The open boxes represent one standard deviation
-(&#963;) to each side of the means.</span>
-</div>
-
-<p><span class="pagenum"><a name="Page_34" id="Page_34">[Pg 34]</a></span></p>
-
-<div class="figcenter" style="width: 400px;">
-<img src="images/fig3.jpg" width="400" height="350" alt="" />
-<span class="caption"><span class="smcap">Fig. 3.</span> Statistics of variation in length of wing in the Lincoln Sparrow.</span>
-</div>
-
-<p>Figures 4-6 show variation in weight in the samples of Nashville
-Warbler, Mourning Warbler, Yellow-throat, Dickcissel, and Lincoln
-Sparrow. Each figure is essentially self-explanatory. It will
-be seen that in the Nashville Warbler and Lincoln Sparrow, weight
-is roughly proportional to wing-length (shown in figs. 2 and 3),
-giving about equally good separation of sex- and age-classes in the
-latter and poorer separation in the former. Data for these and
-other species indicate a generally greater weight of males than of
-females, but less difference according to age. In some other species,
-for example the Yellow-throat, immatures seem to be a little heavier
-on the average than adults. It is unfortunate that wing-lengths are
-not at present available for these.</p>
-
-<div class="figcenter" style="width: 400px;">
-<img src="images/fig4.jpg" width="400" height="655" alt="" />
-<span class="caption"><span class="smcap">Fig. 4.</span> Statistics of variation in weight in the Nashville Warbler and
-Mourning Warbler.</span>
-</div>
-
-<div class="figcenter" style="width: 400px;">
-<img src="images/fig5.jpg" width="400" height="677" alt="" />
-<span class="caption"><span class="smcap">Fig. 5.</span> Statistics of variation in weight in the Yellow-throat and Dickcissel.</span>
-</div>
-
-<div class="figcenter" style="width: 400px;">
-<img src="images/fig6.jpg" width="400" height="346" alt="" />
-<span class="caption"><span class="smcap">Fig. 6.</span> Statistics of variation in weight in the Lincoln Sparrow.</span>
-</div>
-
-<p>These comments on weight suggest an additional factor which
-may play a part in rate of migration and which some day may be
-profitably studied. Suppose that in some species adults and immatures
-are nearly the same in weight but that immatures have
-shorter wings. In such a species the immatures are relatively
-shorter-winged for their weight than adults. In aerodynamic terms,
-<span class="pagenum"><a name="Page_35" id="Page_35">[Pg 35]</a><br /><a name="Page_36" id="Page_36">[Pg 36]</a><br /><a name="Page_37" id="Page_37">[Pg 37]</a></span>they would have a higher "wing-loading." (Wing-loading is the
-result obtained by dividing area of effective wing by total weight;
-it is here assumed that in a single species wing area is directly proportional
-to wing length.) This being the case, immatures with
-higher "wing-loading" would require more energy (derived from
-fat) to fly the same distance as adults, or with the same amount of
-fat they would fly a shorter distance. Thus they might tend to be
-outstripped in migration by adults starting at the same time. The
-reverse, of course, would also be theoretically true, if adults possessed
-a higher wing-loading than immatures. Physical factors
-such as these rather than the differential "virility" postulated by
-Meinertzhagen (1930:56) might account for the arrival of certain
-classes of some species on the wintering grounds in advance of
-others. There are, of course, many other factors which must be
-taken into account before the effect, if any, of the wing-loading
-factor can be evaluated. Data for illuminating calculations will
-become available, however, with the accumulation of abundant information
-on weights, measurements, and migration patterns.</p>
-<hr class="chap" />
-
-<p><span class="pagenum"><a name="Page_38" id="Page_38">[Pg 38]</a></span></p>
-
-
-
-<h2>Computations of Longevity and Survival</h2>
-
-
-<p>Tanner (ms., and letter, April 21, 1955) recently devoted considerable
-ingenuity to computing by actuarial methods the longevity
-of the Oven-bird, using the adult-immature ratio in samples killed
-at a ceilometer at Knoxville, Tennessee. Tanner's computations
-were based on the simple assumption that</p>
-
-
-
-<div class="center">
-<table border="0" cellpadding="4" cellspacing="0" summary="formula">
-<tr><td rowspan="2">S (survival rate) =</td>
- <td>Number of adults in population (or sample)</td></tr>
-<tr><td class="bor_top">Total size of population (or sample).</td></tr>
-</table></div>
-
-<p>Further application of such techniques may prove desirable and
-rewarding. It would seem at present, however, to be a risky procedure,
-as it has been abundantly shown (see above) that adults
-and immatures often do not migrate at the same times and rates,
-and the ratios of adults to immatures in samples of migrants are
-likely to be far from representative of the true proportions in the
-populations concerned. It should be added that Tanner is perfectly
-aware of this objection.</p>
-
-
-
-<hr class="chap" />
-<h2>Processing of Samples</h2>
-
-
-<p>Thorough processing of large samples of birds killed accidentally
-is time-consuming. We were fortunate in having considerable assistance;
-even so, all desirable data could not be obtained from the
-1090 birds of the present sample. As aids to others conducting
-studies of this kind we should mention a few points which may be
-of assistance.</p>
-
-<p>Birds should be picked up as soon as possible after death, certainly
-by the end of the day after the accident and preferably much
-sooner. They should be weighed as soon as possible after collection
-(weights decrease rapidly, even under refrigeration), and the
-weights (in grams, to one tenth of a gram) written on tags attached
-to a leg of each specimen. The sample should then be
-sorted by species or groups of species of approximately equal size
-(to avoid crushing of smaller birds by larger ones), placed in boxes,
-paper bags, or better, air-tight containers clearly marked with date,
-locality, and other necessary particulars, and relegated to a deep-freeze
-compartment. For all but the smallest samples, such freezing
-units are indispensable to complete study. Once frozen, the birds
-may be selected for study at leisure, but time is still important, as,
-even when frozen, gonads may eventually deteriorate, and birds<span class="pagenum"><a name="Page_39" id="Page_39">[Pg 39]</a></span>
-eventually become desiccated which is a disadvantage if skins are
-to be made.</p>
-
-<p>In the cases of large kills, or limited manpower, or both, it may
-be impossible to process all birds, however desirable this might be.
-If possible, however, all should be collected, identified, the numbers
-and species recorded, and rarities saved. Further, partial analysis,
-or more properly, complete analysis of a partial sample, can be
-made. Analyses which for any reason require randomness of
-sample pose a special problem. We think that in very large kills
-the best way to solve this problem is probably to make one or more
-transects across the area where dead birds are found. These
-transects should cross both the areas of greatest and least density
-(to allow for fast and slow flying species). Their width may be
-adjusted to give the desired number of birds, that is, the number
-that can be adequately processed. Another alternative would be
-to decide to study certain abundant species and pick up all of these.
-There are other possibilities, but in any event the method of
-sampling should be thoroughly described wherever all birds have
-not been processed.</p>
-
-
-
-<hr class="chap" />
-<h2>Summary</h2>
-
-
-<p>The foregoing paper discusses accidents in which large numbers
-of night-migrating birds are killed. A brief historical review of
-ornithological interest in such occurrences is given, and the types
-of data provided by the accidents are listed and discussed. In particular,
-recent accidents occurring in early October, 1954, through
-much of eastern United States are cited, and detailed analysis is
-presented of a sample of 1090 birds killed one mile west of Topeka,
-Shawnee County, Kansas, between September 25 and October 23,
-1954.</p>
-
-<p>At Topeka during the period mentioned, 1090 birds representing
-61 species were collected and were processed at the University of
-Kansas. For all specimens, weight, sex, age, and fat condition were
-recorded, and certain species were measured as well. Some notes
-on molt were taken. A total of 193 birds was preserved as study
-skins, and 49 as skeletons. Comments on weight, size, sex, age,
-subspecific identity, and status in Kansas are presented in an annotated
-list.</p>
-
-<p>Randomness of this and other similar samples is discussed. A
-theoretical computation is given for several nights of the numbers<span class="pagenum"><a name="Page_40" id="Page_40">[Pg 40]</a></span>
-of migrants passing through a plane one mile in width, from 450
-to 950 feet above ground level, and oriented to face the arriving
-migrants. The computed totals give some idea of the tremendous
-volume of nocturnal migration under some conditions. Potentialities
-of further study of such events are discussed and a comparison
-is made with lunar observations.</p>
-
-<p>Differential migration of sex- and age-groups as shown by the
-larger samples taken at Topeka (Catbird, Red-eyed Vireo, Nashville
-Warbler, Yellow-throat, Mourning Warbler, Dickcissel, Lincoln
-Sparrow) is discussed and the history of theories on this subject
-reviewed. It is shown that age and sex must both be taken
-into account in studies of differential migration. Several patterns
-of migration are displayed by the species analyzed, adults migrating
-earlier than immatures in some instances, young earlier than adults
-in others, but all seemingly being complicated to varying degrees
-by differential migration of sexes within age groups. It is suggested
-that explanations of these patterns may be sought in the
-life histories of the species involved.</p>
-
-<p>Molt in relation to migration is discussed briefly, and it is held
-that there is an important relationship between molt and time of
-migration. Specimens noted to be in molt are listed.</p>
-
-<p>Size differences, in wing length, tail length, and weight are discussed
-in relation to sex and age, and variation in one or more of
-these characters is analyzed as found in the Topeka samples of
-Nashville Warbler, Mourning Warbler, Yellow-throat, Dickcissel,
-and Lincoln Sparrow. It seems that in some instances significant
-size differences prevail between adults and immatures and that
-these age classes should be separated in taxonomic work with species
-where differences in size are known to exist. When the facts
-are not known they should be determined, and the large samples
-collected in accidents to nocturnal migrants present excellent opportunities
-for ascertaining the facts.</p>
-
-<p>Other uses of material obtained in large migration accidents are
-discussed, such as computations of longevity and the problems of
-processing large, accidentally-killed samples. Care should be taken
-to select samples as nearly random as possible when all birds cannot
-be processed.</p>
-
-<p>Repeated and thorough analysis of accidental kills should provide
-a mass of valuable data bearing on many questions and problems
-which have thus far been little studied.</p>
-
-<hr class="chap" />
-
-<p><span class="pagenum"><a name="Page_41" id="Page_41">[Pg 41]</a></span></p>
-
-
-
-
-<h2>Literature Cited</h2>
-
-
-<p><span class="smcap">Allen, J. A.</span></p>
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-<blockquote>
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-<p>1880. Destruction of birds by light-houses. Bull. Nuttall Orn. Club,
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-<p>1896. G&auml;tke's 'Heligoland.' Auk, 13(2):137-153, April.</p>
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-<p>1901. Barrington's 'The Migration of Birds at Irish Light Stations.' [Review.]
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-<p>1909. An American's views of bird migration. Brit. Birds, 3(1):12-19,
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-<p><span class="smcap">Barrington, R. M.</span></p>
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-<p>1900. The migration of birds as observed at Irish lighthouses and lightships
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-some sources list pagination as XXV + 667, possibly in error.)</p></blockquote>
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-<p><span class="smcap">Behle, W. H.</span></p>
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-<p>1942. Distribution and variation of the Horned Larks (Otocoris alpestris)
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-<p>1950. Clines in the Yellow-throats of western North America. Condor,
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-<p>1886. Bird migration. Mem. Nuttall Orn. Club, No. I, pp. 1-22, Cambridge,
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-<p><span class="smcap">Carson, L. B.</span></p>
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-<blockquote>
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-<p>1954a. [Destruction of birds at a television tower at Topeka, Kansas.]
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-October]. [Unsigned, unpaged, and untitled article by
-L. B. Carson, ed.]</p>
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-<p>1954b. [Further destruction of birds at a television tower at Topeka, Kansas.]
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-[published December]. [Unsigned, unpaged, and untitled
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-<p>1954c. New records for fall migrants in eastern Kansas. Kansas Orn. Soc.
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-<blockquote>
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-<p>1892. The migration of birds. London, Chapman and Hall. Pp. XVI +
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-<p><span class="smcap">Dobben, W. W. H. van</span>, and <span class="smcap">M. F. Bruyns</span></p>
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-<blockquote>
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-<p>1939. Zug nach Alter und Geschlecht an niederl&auml;ndischen Leuchtt&uuml;rmen.
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-<p><span class="smcap">Drost, R.</span></p>
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-<blockquote>
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-<p>1935. Ueber das Zahlenverh&auml;ltnis von Alter und Geschlecht auf dem
-Herbst-und Fr&uuml;hjahrszuge. Vogelzug, 6(4):177-182, October.</p></blockquote>
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-<p><span class="smcap">Dwight, J., Jr.</span></p>
-
-<blockquote>
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-<p>1900. The sequence of plumages and moults of the passerine birds of
-New York. Annals New York Acad. Sci., 13(1):73-360, October 19.</p></blockquote>
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-<p><span class="pagenum"><a name="Page_42" id="Page_42">[Pg 42]</a></span></p>
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-<p><span class="smcap">Ganier, A. F.</span></p>
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-<blockquote>
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-<p>1949. The late summer Dickcissel departure. Migrant, 20(3):52-53,
-September.</p></blockquote>
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-<p><span class="smcap">G&auml;tke, H.</span></p>
-
-<blockquote>
-
-<p>1895. Heligoland as an ornithological observatory. Translated by Rudolph
-Rosenstock. Edinburgh, David Douglas. Pp. X + II +
-599 + 11 II. (advt.).</p></blockquote>
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-<p><span class="smcap">Gross, A. O.</span></p>
-
-<blockquote>
-
-<p>1921. The Dickcissel (Spiza americana) of the Illinois prairies. Auk,
-38(1):1-26, January 18 (first of two parts).</p></blockquote>
-
-<p><span class="smcap">Gurney, J. H.</span></p>
-
-<blockquote>
-
-<p>1923. Bird migration as observed on the east coast of England. Ibis, 11th
-ser., 5(4):573-603, October 3.</p></blockquote>
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-<p><span class="smcap">Howell, J. C.</span>, <span class="smcap">Laskey, A. R.</span>, and <span class="smcap">J. T. Tanner</span></p>
-
-<blockquote>
-
-<p>1954. Bird mortality at airport ceilometers. Wilson Bull., 66(3):207-215,
-September [published October 29].</p></blockquote>
-
-<p><span class="smcap">Howell, J. C.</span>, and <span class="smcap">J. T. Tanner</span></p>
-
-<blockquote>
-
-<p>1951. An accident to migrating birds at the Knoxville airport. Migrant,
-22(4):61-62, December.</p></blockquote>
-
-<p><span class="smcap">Laskey, A. R.</span></p>
-
-<blockquote>
-
-<p>1951. Another disaster to migrating birds at the Nashville airport. Migrant,
-22(4):57-60, December.</p></blockquote>
-
-<p><span class="smcap">Long, W. S.</span></p>
-
-<blockquote>
-
-<p>1934. Notes from eastern Kansas. Auk, 51(2):255, April 4.</p>
-
-<p>1940. Check-list of Kansas birds. Trans. Kansas Acad. Sci., 43:433-456.</p></blockquote>
-
-<p><span class="smcap">Lovell, H. B.</span></p>
-
-<blockquote>
-
-<p>1952. Catastrophe to birds at a Louisville airport. Kentucky Warbler,
-28(1):5-6, February.</p></blockquote>
-
-<p><span class="smcap">Lowery, G. H., Jr.</span></p>
-
-<blockquote>
-
-<p>1951. A quantitative study of the nocturnal migration of birds. Univ.
-Kansas Publ., Mus. Nat. Hist., 3(2):361-472, June 29.</p></blockquote>
-
-<p><span class="smcap">McCabe, T. T.</span></p>
-
-<blockquote>
-
-<p>1943. An aspect of collectors' technique. Auk, 60(4):550-558, October 7.</p></blockquote>
-
-<p><span class="smcap">McCabe, T. T.</span>, and <span class="smcap">A. H. Miller</span></p>
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-<blockquote>
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-<p>1933. Geographic variation in the Northern Water-thrushes. Condor,
-35(5):192-197, September-October (September 15).</p></blockquote>
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-<p><span class="smcap">Meinertzhagen, R.</span></p>
-
-<blockquote>
-
-<p>1930. Nicoll's birds of Egypt. Vol. I (of 2). London, Hugh Rees Ltd.
-Pp. XVI + 348.</p></blockquote>
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-<p><span class="smcap">Mengel, R. M.</span></p>
-
-<blockquote>
-
-<p>1952. Certain molts and plumages of Acadian and Yellow-bellied flycatchers.
-Auk, 69(3):273-283, July 7.</p></blockquote>
-
-<p><span class="smcap">Miller, A. H.</span></p>
-
-<blockquote>
-
-<p>1941. Speciation in the avian genus Junco. Univ. California Publ. Zo&ouml;l.,
-44(3):173-434, May 24.</p></blockquote>
-
-<p><span class="pagenum"><a name="Page_43" id="Page_43">[Pg 43]</a></span></p>
-
-<p><span class="smcap">Miller, A. H.</span>, and <span class="smcap">T. T. McCabe</span></p>
-
-<blockquote>
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-<p>1935. Racial differentiation in Passerella (Melospiza) lincolnii. Condor,
-37(3):144-160, May-June (May 15).</p></blockquote>
-
-<p><span class="smcap">Palm&eacute;n, J. A.</span></p>
-
-<blockquote>
-
-<p>1876. Ueber die Zugstrassen der V&ouml;gel. Leipzig, Wilhelm Engelmann.
-Pp. VI + 292 + I.</p></blockquote>
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-<p><span class="smcap">Peters, J. L.</span>, and <span class="smcap">L. Griscom</span></p>
-
-<blockquote>
-
-<p>1938. Geographical variation in the Savannah Sparrow. Bull. Mus. Comp.
-Zo&ouml;l., 80(13):445-478, January.</p></blockquote>
-
-<p><span class="smcap">Pitelka, F. A.</span></p>
-
-<blockquote>
-
-<p>1946. Age in relation to migration in the Blue Jay. Auk, 63(1):82-84,
-January 25.</p>
-
-<p>1951. Speciation and ecologic distribution in American jays of the genus
-Aphelocoma. Univ. California Publ. Zo&ouml;l., 50(3):195-464, July 20.</p></blockquote>
-
-<p><span class="smcap">Rintoul, L. J.</span>, and <span class="smcap">E. V. Baxter</span></p>
-
-<blockquote>
-
-<p>1914. Notes on some passerine birds found migrating in moult. Scottish
-Naturalist, no. 35, pp. 245-252, November.</p></blockquote>
-
-<p><span class="smcap">Rowan, W.</span></p>
-
-<blockquote>
-
-<p>1926. Notes on Alberta waders included on the British list. Part II.
-Brit. Birds, 20(2):34-42, June 1.</p></blockquote>
-
-<p><span class="smcap">Ruth, E. L.</span></p>
-
-<blockquote>
-
-<p>1952. The Bay-breasted Warbler in Kansas. Kansas Orn. Soc. Bull.,
-3(3):18-19.</p></blockquote>
-
-<p><span class="smcap">Sibley, C. G.</span></p>
-
-<blockquote>
-
-<p>1950. Species formation in the Red-eyed Towhees of Mexico. Univ. California
-Publ. Zo&ouml;l., 50(2):109-194, November 24.</p></blockquote>
-
-<p><span class="smcap">Spofford, W. R.</span></p>
-
-<blockquote>
-
-<p>1949. Mortality of birds at the ceilometer of the Nashville airport. Wilson
-Bull., 61(2):86-90, June.</p></blockquote>
-
-<p><span class="smcap">Stone, W.</span></p>
-
-<blockquote>
-
-<p>1906. Some light on night migration. Auk, 23(3):249-252, July.</p></blockquote>
-
-<p><span class="smcap">Storer, R. W.</span></p>
-
-<blockquote>
-
-<p>1951. Variation in the Painted Bunting (<i>Passerina ciris</i>), with special
-reference to wintering populations. Occas. Papers Mus. Zool.,
-Univ. Michigan, no. 532, pp. 1-12, June 29.</p></blockquote>
-
-<p><span class="smcap">Thomson, A. L.</span></p>
-
-<blockquote>
-
-<p>1926. Problems of bird-migration. Boston and New York, Houghton
-Mifflin Company. Pp. XV + I + 350.</p>
-
-<p>1936. Recent progress in the study of bird-migration: a review of the
-literature, 1926-35. Ibis, 13th ser., 6(3):472-530, July 1.</p></blockquote>
-
-<p><span class="smcap">Ticehurst, C. B.</span></p>
-
-<blockquote>
-
-<p>1916. Notes on migrants and moult, with special reference to the moults
-of some of our summer visitants. Scottish Naturalist, no. 50, pp.
-29-38, February.</p></blockquote>
-
-<p><span class="pagenum"><a name="Page_44" id="Page_44">[Pg 44]</a></span></p>
-
-<p><span class="smcap">Tordoff, H. B.</span></p>
-
-<blockquote>
-
-<p>1952. Notes on plumages, molts, and age variation of the Red Crossbill.
-Condor, 54(4):200-203, July-August.</p></blockquote>
-
-<p><span class="smcap">Wiegold, H.</span></p>
-
-<blockquote>
-
-<p>1926. Masse, Gewichte und Zug nach Alter und Geschlecht bei Helgol&auml;nder
-Zugv&ouml;geln. Wissenschaftliche Meeresuntersuchungen, Abt.
-Helgoland, Neue Folge, 15ter Band, Heft 3, No. 17, Lipsius &amp;
-Tischer, Kiel und Leipzig, pp. 1-73.</p></blockquote>
-
-<p><span class="smcap">Wolfson, A.</span></p>
-
-<blockquote>
-
-<p>1954. Weight and fat deposition in relation to spring migration in transient
-White-throated Sparrows. Auk, 71(4):413-434.</p></blockquote>
-
-<p><i>Transmitted June 30, 1955.</i></p>
-
-
-
-<hr class="chap" />
-
-<div class="transnote">
-<h2>Transcriber's Notes:</h2>
-
-<p>Obvious printer's errors have been repaired, other inconsistent
-spellings have been kept, for example inconsistent use of hyphen (e.g.
-"age-classes" and "age classes") and diacritical mark (e.g. "Zool."
-and "Zoöl.")</p>
-</div>
-
-
-
-
-
-
-
-
-<pre>
-
-
-
-
-
-End of the Project Gutenberg EBook of Studies of Birds Killed in Nocturnal
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