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*** START OF THE PROJECT GUTENBERG EBOOK 40112 ***

Transcriber's Notes

Text Emphasis
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_Text_ - Italics
=Text= - Bold

Symbolic Representations
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[F]   Female symbol
[M]   Male symbol
[BC]  Circle Black
[TW]  Circle Top White
[RW]  Circle Right White
[BW]  Circle Bottom White
[LW]  Circle Left White
[RTW] Circle Right Third White
[LTW] Circle Left Third White




UNIVERSITY OF KANSAS PUBLICATIONS

MUSEUM OF NATURAL HISTORY

Vol. 16, No. 8, pp. 777-837, 17 figures in text

December 16, 1968


Systematics of Megachiropteran Bats in the Solomon Islands

BY

CARLETON J. PHILLIPS


UNIVERSITY OF KANSAS

LAWRENCE

1968




UNIVERSITY OF KANSAS PUBLICATIONS, MUSEUM OF NATURAL HISTORY

Editorial Committee: E. Raymond Hall, Chairman; Frank B. Cross, Editor;
Henry S. Fitch; J. Knox Jones, Jr.


Volume 16, No. 8, pp. 777-837, 17 figs.

Published December 16, 1968


UNIVERSITY OF KANSAS

Lawrence, Kansas


PRINTED BY

ROBERT R. (BOB) SANDERS, STATE PRINTER

TOPEKA, KANSAS

1968

[Illustration]

31-9490




Systematics of Megachiropteran Bats in the Solomon Islands

BY

CARLETON J. PHILLIPS




CONTENTS


                                                                  PAGE

  INTRODUCTION                                                     781

  GAZETTEER                                                        783

  METHODS AND MATERIALS                                            786

  ACKNOWLEDGMENTS                                                  786

  KEY TO GENERA                                                    787

  _Rousettus amplexicaudatus hedigeri_ Pohle                       788

  _Pteralopex atrata atrata_ Thomas                                791

  _Pteralopex atrata anceps_ Andersen                              792

  _Pteropus hypomelanus luteus_ Andersen                           796

  _Pteropus admiralitatum solomonis_ Thomas                        796

  _Pteropus admiralitatum colonus_ Andersen                        796

  _Pteropus admiralitatum goweri_ Tate                             797

  _Pteropus howensis_ Troughton                                    797

  _Pteropus tonganus geddiei_ MacGillivary                         798

  _Pteropus rayneri rayneri_ Gray                                  800

  _Pteropus rayneri grandis_ Thomas                                801

  _Pteropus rayneri rubianus_ Andersen                             802

  _Pteropus rayneri lavellanus_ Andersen                           802

  _Pteropus rayneri monoensis_ Lawrence                            803

  _Pteropus rayneri cognatus_ Andersen                             803

  _Pteropus rayneri rennelli_ Troughton                            804

  _Pteropus woodfordi_ Thomas                                      804

  _Pteropus mahaganus_ Sanborn                                     806

  _Dobsonia inermis inermis_ Andersen                              808

  _Dobsonia inermis_ new subspecies                                809

  _Macroglossus lagochilus microtus_ Andersen                      813

  _Melonycteris aurantius_ Phillips                                816

  _Melonycteris woodfordi_ Thomas                                  816

  _Nyctimene albiventer bougainville_ Troughton                    818

  _Nyctimene albiventer_ new subspecies                            819

  _Nyctimene_ new species                                          822

  _Nyctimene major scitulus_ Andersen                              825

  ZOOGEOGRAPHY AND SPECIATION                                      825

  LITERATURE CITED                                                 834




INTRODUCTION


The Solomon Islands constitute an archipelago east of the large island
of New Guinea and more than a thousand miles off the northeastern coast
of Australia. This archipelago, which is principally of volcanic origin
although sedimentary layers of calcareous rocks occur on many islands
(Lever, 1934; Belkin, 1962), consists of a double chain of islands
having a northwest-southeast axis of more than 600 miles. The
archipelago is more or less an extension of New Guinea and in fact is
connected to it in stepping-stone fashion by New Britain, New Ireland,
and numerous smaller islands (see Fig. 1).

Australia and New Guinea have many kinds of mammals but the only
terrestrial mammals in the Solomon Islands are a species of the genus
_Phalanger_ (order Marsupialia), and several species of four genera of
rodents, one genus of which probably was introduced by man.
Additionally, several kinds of bats have reached and colonized the
Solomon Islands.

    [Illustration: FIG. 1. Showing the Solomon Islands in relation to
       major adjacent land masses.]

In the past 100 years at least 43 species and subspecies of Chiroptera
of 16 genera have been recorded from the Solomon Islands; of these 27
species and subspecies of seven genera are in the suborder
Megachiroptera. At least one genus of Megachiroptera is endemic as are
numerous species of other genera, and subspecies of still other species.

In 1963 and 1964, the Bernice P. Bishop Museum sent several collecting
parties to the British Solomon Islands Protectorate and the Australian
Trust Territory of New Guinea. In the Solomons, J. Linsley Gressitt,
Philip Temple, Peter Shanahan, and Ray Straatmann visited many of the
larger and more accessible islands and collected a wealth of zoological
materials. I have had the opportunity to study and report on specimens
of mammals, especially bats, collected by the persons named and
deposited in the Bishop Museum. This report is the third in a series on
bats from the Solomons (Phillips, 1966; 1967). Other specimens, mostly
obtained in 1944 by personnel of United States military units, are
stored in the United States National Museum and have been available for
study. Aims of the following report are to (1) identify the
megachiropteran bats to species and subspecies and (2) discuss
distribution of these bats in the Solomon Islands.

In all, 27 kinds (subspecies and monotypic species) of the order
Megachiroptera are known from the Solomon Islands. These pertain to
three subfamilies of the one family Pteropodidae.

The 43 Solomon Islands, having a total land area of more than 15,300
square miles (see Belkin, 1962:42-43), are listed in the gazetteer (see
also Figure 2). Politically, all of the Solomon Islands except Buka and
Bougainville, which are included in New Guinea Trust Territory under
mandate to Australia, are in the British Solomon Islands Protectorate.

The Solomons are within 300 to 700 miles of the equator and have a
fairly constant tropical climate, except at high elevations. The
temperature varies little; monthly mean temperature is between 81° and
83° F. and at sea level ranges from about 70° to 93° F. yearly (Belkin,
1962:42).

Southeast tradewinds are relatively constant from May to October and
this period, in general, is a dry season except at higher elevations on
windward coasts. From December to March prevailing winds are from the
north and precipitation throughout the island group is especially heavy.
Rainfall on the island of Tulagi averages about 120 inches per year
(Bryan, Edwin H., 1941; MS, p. 2, at Pacific Sci. Information Center,
Bishop Museum) and up to 300 inches have been recorded on the north
coast of Guadalcanal (Belkin, 1962:42-43). Occasional dry periods occur
even in the period of December to March.

Most islands of the Solomon Group support dense tropical rain forest.
Much of it has been modified by man. Some clearings and scattered
coconut plantations are found along coasts. On some of the larger
islands (for example, Guadalcanal) coastal scrub (especially on leeward
coasts) and extensive grassy areas are to be found. Additional notes on
vegetation are in the gazetteer.

The 165,000 persons living on the Solomon Islands are mostly
Melanesians but some are mixed Papuan, Malay, and Polynesian. These
native peoples are notorious for their cannibalistic tendencies; the
eating of human flesh usually was related to warfare, although
malefactors and human sacrifices accounted for some of the cannibalism
(Cranstone, 1961:29). Prior to the Second World War few Europeans
visited the Solomons and several islands still remain beyond reach of
modern-day technology. For example, Rennell and Bellona islands, south
of the main part of the archipelago, are visited only rarely, and then
only by a medical officer or the Resident Commissioner. According to
Troughton (1936:341), the islanders in the interior of Bougainville as
late as 1935, were prone to kill and feast upon strangers. In 1932,
Lewis (1951:37) felt that the natives of Malaita Island were especially
resistant to outside interference by Caucasians and reported that no
"white man or foreigner" was safe on Malaita.

Troughton (1936), who listed Melanesian names for mammals, indicated
that the native peoples distinguished between kinds of bats that closely
resembled one another. Of these, the only bats that seem to be used as
food belong to the genus _Pteropus_.




GAZETTEER


In the following list, currently-used names of islands are given; when
available, older names and variant spellings are indicated in
parentheses. For certain islands, especially those visited by field
parties from the Bishop Museum or those frequently mentioned in previous
literature on bats, some descriptive and ecological information also is
provided.

Latitude and longitude of islands are from publication no. 881 of the
Hydrographic Office of the United States Navy Department (Anonymous,
1944); names of islands were checked against a list by Brigham (1900);
descriptive information mostly is from reports by Temple and Straatmann
(1964, field notes, at the Department of Entomology, Bishop Museum).

ALU.--7° 07' S, 155° 54' E.

BANIKA.--9° 05' S, 155° 13' E.

BARA (Gera).--9° 31' S, 160° 31' E.

BELLONA (Bello).--11° 18' S, 159° 48' E.

BOUGAINVILLE (Mamamolimo).--6° 12' S, 155° 15' E. This is the largest
island in the Solomon Group, being 127 miles long (northwest to
southeast) and about 59 miles across at the widest place. The highest
elevations are 9850 and 10171 feet, at the tops of active volcanoes.
Ecologically, Bougainville is mostly dense rain forest, which is less
dense on the summits of higher mountains.

BUKA.--5° 15' S, 154° 38' E.

CHOISEUL.--7° 04' S, 157° 01' E. This island, formed along a
northwest-southeast line of low mountains (maximum elevation of 3500
feet), is about 90 miles long and 20 miles wide. Most collecting was at
Malangona (Sasamunga on some maps) on the southwestern coast.

FAURO.--6° 55' S, 156° 07' E. This small island, about 14 miles long
(north-south) and six miles wide (east-west), lies about 10 miles south
and east of Bougainville. Fauro is formed around a volcanic cone having
a maximum elevation of 1925 feet; it has considerable dense mangrove
swamp along the west coast, and mature rain forest with little
understory growth. Most collecting was at Toumoa, on one of two southern
peninsulas.

FLORIDA (Nggela).--9° 05' S, 160° 16' E. Florida, the main island in the
Nggela Island Subgroup, is mountainous and except for some small grassy
areas, supports dense rain forest. It is nearly 25 miles long
(east-west) and nine miles wide (north-south), with a maximum
elevation, at Mount Barnett, of about 1366 feet. Most collecting was at
Haleta, on the southwestern coast. At this locality there were scattered
mangrove swamps, rain forest, and gardens inland.

GANONGGA (Ronogo, Ronongo).--8° 03' S, 156° 35' E.

GATUKAI.--8° 47' S, 158° 12' E.

GHIZO (Gizo, Keso).--8° 05' S, 156° 59' E.

GOWER (N'dai).--7° 54' S, 160° 34' E.

GUADALCANAL (Guadalcanar).--9° 15' S, 159° 35' E. Guadalcanal is mostly
of volcanic origin and has an irregular chain of mountains along the
southern coast. The highest elevation is 8005 feet at Mount Popomanasiu.
This large island is nearly 80 miles long (east-west) and 25 miles wide
(north-south). Most of the northwestern part of Guadalcanal supports
_alang-alang_ grass. The remainder of the island is heavily wooded.

KILINAILAU (Cartaret).--4° 44' S, 155° 28' E.

KOLOMBANGARA (Duki, Kulambangara).--8° 00' S, 157° 05' E. Kolombangara,
formed from an extinct volcano, is about 18 miles in diameter and nearly
circular. The highest peaks, rising as precipitous cliffs in some
places, reach a maximum elevation of about 5000 feet. The vegetation is
mostly virgin rain forest. Mangrove swamp and small coconut groves occur
along the coast. Field parties from the Bishop Museum were able to reach
the highest elevations, and concentrated their work along the
southwestern side of the island.

    [Illustration: FIG. 2. Solomon Islands. Principal islands
       are named.]

MALAITA (Mala, Malanta, Malayta).--9° 00' S, 161° 00' E. This long (104
miles northwest to southeast), narrow (about 23 miles at its widest
spot) island, between Santa Ysabel and San Cristobal islands, is
basically of volcanic origin with some limestone (coral) deposits along
the coast. Mount Kolovrat, having an elevation of 4275 feet, is the
highest point. The Bishop Museum field party lived at Dala, in dense
rain forest about 12 miles north of Auki on the northwestern coast of
Malaita.

MALAPA.--9° 49' S, 160° 53' E.

MONO (Treasury).--7° 22' S, 155° 35' E. This is a small island (maximum
elevation 1150 feet) in the Treasury Island Subgroup just south of
Bougainville. Mono is about nine miles long (east-west) and five and one
half miles wide (north-south). The basic volcanic core is described in
field notes as topped with coral limestone.

NEW GEORGIA (Kausagi).--8° 20' S, 157° 30' E. The New Georgia Subgroup
is composed of 11 moderate-sized islands and islets. New Georgia Island,
the main member of the subgroup, is 50 miles long (northwest to
southeast) and from five to 30 miles wide. On the northern side several
volcanic peaks attain an elevation of about 3000 feet. The entire island
is forested.

NGGELA (Florida Islands).--4° 31' S, 154° 11' E. This subgroup consists
of several small to medium-sized islands between Guadalcanal and
Malaita. Florida is the main island.

NISSAN (Green, Sir Charles Hardy's).--4° 31' S, 154° 11' E.

NUKUMANU (Le Maira, Tasman).--4° 32' S, 159° 25' E.

ONTONG JAVA (Lord Howe Atoll, Liuniuwu).--5° 25' S, 159° 30' E.

PAVUVO.--9° 04' S, 159° 08' E.

RAMOS.--8° 16' S, 160° 11' E.

RENNELL.--11° 38' S, 160° 14' E. This island, of limestone (coral)
origin, along with Bellona, is nearly 100 miles southwest of any other
member of the Solomons and has been regarded, because of this distance,
as an oceanic island instead of a continental island. It is about 50
miles long (east-west) and nine miles wide (north-south); its highest
elevation is 500 feet.

ROVIANA (Rendova, Rovianna, Rubiana).--8° 21' S, 157° 20' E.

RUSSELL.--9° 04' S, 159° 12' E.

SAN CRISTOBAL (San Christoval, Bauro, Makira, Arussi).--11° 33' S, 161°
43' E. This island is composed mostly of ancient volcanic rock, has a
maximum elevation of 4100 feet, is nearly 70 miles long (northwest to
southeast) and 24 miles wide, and supports a dense rain forest.

SANTA YSABEL (George, Ysabel, San Isabel, Isbel, Mahaga).--8° 00' S,
159° 07' E. Santa Ysabel is a long (90 miles from northwest to
southeast), narrow (19 miles at the widest spot), forested island,
consisting of a single chain of volcanic mountains. The numerous bays
and mouths of rivers provide excellent anchorages. Collecting was at
Tatamba approximately two miles south of Tanambuli where the
considerable area of forest was dense and bamboo thickets were abundant.

SAVO (Savu).--9° 08' S, 159° 49' E.

SHORTLAND.--7° 03' S, 155° 47' E.

SIKAIANA (Stewart).--8° 22' S, 162° 44' E.

SIMBO (Narovo, Naorovo, Naravo, Navoro, Sembo).--8° 16' S, 156° 31' E.

STIRLING.--7° 25' S, 155° 35' E.

TANABULI (Tanambuli, Tunnibili, Tunnibilis, Tunnibul, Tunnivula).--8°
24' S, 159° 35' E.

TAUU (Marqueen, Mortlock).--4° 48' S, 157° 32' E.

TELIPARI.--8° 15' S, 157° 32' E.

UGI.--10° 14' S, 161° 44' E.

VANGUNO (Vangunu).--8° 39' S, 158° 00' E.

VELLA LAVELLA.--7° 43' S, 156° 40' E. The coastline is rugged and
indented by numerous small bays. Some peaks are 3000 feet high. The
southeastern half of Vella Lavella is said to consist of uplifted coral,
and to be thickly planted to coconut palms. The native population is
concentrated here. The northwestern half of the island is rain forest
and is nearly uninhabited. Most of the collecting was at Pusisama, on
the southern beach and on Ulo Crater, an extinct volcano at the middle
of the island.

YANNTA.--10° 20' S, 161° 20' E.




METHODS AND MATERIALS


The phylogenetic arrangement and nomenclature in the text beyond are
mainly that of Laurie and Hill (1954). The synonymies for accounts of
genera are as follows: (1) first use of the generic name employed along
with the original description, and (2) original proposals, in
chronological order, of other generic names subsequently applied to the
bat in the Solomons. The synonymies in accounts of species and
subspecies are as follows: (1) first use of the accepted name, followed
by its type locality, followed, in chronological order, by other
references to the first name-combination, (2) first use of the
name-combination employed herein (if different from the original
combination), followed, in chronological order, by other references to
the present name-combination, and (3) other name-combinations, in
chronological order, employed for the bat in the Solomons. The word
"part" is used in parentheses after a name if some specimens listed
under that name are from the Solomon Islands and are referable to the
species or subspecies being written about.

Unless noted otherwise, specimens listed as examined were prepared
originally as museum skins with skulls. Approximately 70 per cent of
bats collected in the Solomons were preserved in formalin and now are
stored in alcohol. Because it was necessary to obtain dimensions and
examine various morphological characteristics of skulls, many crania
were extracted from bats preserved in alcohol.

Although all specimens in the Bishop Museum from the Solomon Islands
have been catalogued with the prefix BBM-BSIP, catalogue numbers without
prefixes in the lists of specimens examined refer to this museum.
Catalogue numbers with the prefix USNM refer to specimens in the U. S.
National Museum and those with the prefix AM-M refer to specimens in the
Australian Museum.

Unless indicated otherwise, all measurements in this paper are in
millimeters and are of adults. Cranial measurements, and external
measurements of specimens stored in alcohol, were taken by me. The
cranial measurements were taken with dial calipers using techniques
described by Hall (1946:672-685). External measurements (except length
of forearm) of specimens originally prepared as dried study skins, were
transcribed from specimen labels.

Capitalized color nomenclature is from Ridgway (1912). Noncapitalized
color terms are from published reports that did not use Ridgway's
terminology.




ACKNOWLEDGMENTS


Financial support for this investigation was from (1) a United
States Army Medical Research and Development Command grant
(DA-MD-49-193-62-G65) to the Entomology Department of the Bernice P.
Bishop Museum, and (2) a National Science Foundation grant (2185-4703)
to the author, through the Committee on Systematics and Evolutionary
Biology of The University of Kansas. I am grateful to many individuals
who have helped me in various ways throughout the course of this study.
Dr. J. Linsley Gressitt, Chairman of the Entomology Department, Bernice
Bishop Museum, allowed me to study specimens collected by his
expeditions; Professors E. Raymond Hall and J. Knox Jones, Jr., of the
Museum of Natural History and the Department of Zoology, The University
of Kansas, offered advice and guidance and constructively reviewed the
manuscript. Other persons who have given me assistance and, in some
cases, arranged for loans of comparative materials, are: Dr. David H.
Johnson, Division of Mammals, United States National Museum; Mr. Hobart
M. Van Deusen and Dr. Richard G. Van Gelder, Archbold Expeditions and
Department of Mammalogy, American Museum of Natural History; Messrs.
Ellis LeG. Troughton and Basil Marlow, Mammal Department, The Australian
Museum; Dr. Joseph Curtis Moore, Department of Mammalogy, Field Museum
of Natural History; Mr. John Edwards Hill, Mammal Room, British Museum
(Natural History); Prof. William B. Davis, Department of Zoology, Texas
A & M University; Miss Barbara Lawrence, Museum of Comparative Zoology,
Harvard University. Messrs. Jerry R. Choate and H. H. Genoways, two
colleagues in zoology at The University of Kansas, have assisted me in
many ways, for which I am grateful. Linda Anne Phillips, my wife,
prepared many of the figures and tables used herein. I thank also
Setsuko Nakata, Edwin H. Bryan, Robert Bowan, and Ilse Koehler, who, as
staff members of the Bishop Museum, were especially helpful to me. Most
of the specimens reported herein were collected by Philip Temple and
Peter Shanahan.




Key to Genera

  1.     Uropatagium lacking, or, if present, deeply indented in
         center; tail vertebrae absent, or if present, free          2

  1'.    Uropatagium present, not indented; tail vertebrae present,
         free or in uropatagium                      MICROCHIROPTERA 1

  2(1).  External tail-vertebrae lacking, or, if present, less
         than 3 mm long                                              3

  2'.    External tail-vertebrae more than 3 mm long                 6

  3(2).  Small or medium-sized (forearm less than 50);
         tongue long, extensile                                      4

  3'.    Large (forearm more than 80); tongue not long and
         extensile                                                   5

  4(3).  Uropatagium present; small claw present on second phalanx
         of second digit; tail short (about 3 mm)
                                                =Macroglossus=, p. 812

  4'.    Uropatagium absent; no claw on second phalanx of second
         digit; no tail                         =Melonycteris=, p. 814

  5(3'). Entire back set with hair; wing membranes not meeting
         at middle of back                          =Pteropus=, p. 793

  5'.    Back naked; wing membranes meeting at middle of back,
                                                  =Pteralopex=, p. 790

  6(2'). Nostrils having definite tubelike extensions
                                                   =Nyctimene=, p. 817

  6'.    Nostrils lacking tubelike extensions                        7

  7(6'). Forearm less than 80; large, sharp claw on second phalanx
         of second digit; four upper incisors      =Rousettus=, p. 787

  7'.    Forearm more than 90; small, blunt claw on second phalanx
         of second digit; two upper incisors        =Dobsonia=, p. 807


Family PTEROPODIDAE


Subfamily Pteropodinae


Rousettus Gray

   1821. _Rousettus_ Gray, London Medical Repository, 15:299, April 1.

   1843. _Xantharpyia_ Gray, List of species ... British Museum, p. 37.

   1852. _Cynonycteris_ Peters, Reise nach Mossambique, p. 25.

The genus _Rousettus_ occurs throughout the tropical regions of the Old
World, and in the Solomons is readily distinguished from all other
megachiropteran genera by having both a small claw on the second digit
and free caudal vertebrae. The oriental species have been divided into
two groups on the basis of size (Tate, 1942:344). The subspecies
_Rousettus amplexicaudatus hedigeri_ appears to be the sole
representative of this genus in the Solomon Islands. Prior to 1953,
several workers (Thomas, 1887b:323, 1888b:475; Matschie, 1899:68;
Sanborn, 1931:11) used the name _Rousettus amplexicaudatus brachyotis_
for it, but Pohle (1953) suggested that the specimens from the Solomons
recorded by earlier workers were _R. a. hedigeri_ named by him on the
basis of the specimen that he saw from Bougainville.


=Rousettus amplexicaudatus=

_Rousettus amplexicaudatus_ has at least three subspecies, one of which
is endemic to the Solomon Islands. The species is wide-ranging, being
known from as far west as Thailand (Ellerman and Morrison-Scott,
1966:93) and as far east as the Solomons.

    [Illustration: FIG. 3. Distribution of _Rousettus amplexicaudatus
       hedigeri_. For names of islands see Fig. 2.]


=Rousettus amplexicaudatus hedigeri= Pohle

   1953. _Rousettus amplexicaudatus hedigeri_ Pohle, Z. Säugetierk.,
         17:127, October 27, type from Bougainville.

   1887. _Cynonycteris brachyotis_, Thomas, Proc. Zool. Soc. London,
         p. 323, March 15; 1888, Thomas, Proc. Zool. Soc. London, p. 475,
         December 4, from Fauro.

   1889. _Xantharpyia brachyotis_, Matschie, Die Megachiroptera ...
         naturkunde, p. 68, from Guadalcanal.

   1912. _Rousettus brachyotis_, Andersen, Catalogue of the Chiroptera
         ... British Museum, 1:809; 1931, Sanborn, Publ. Field Mus. Nat.
         Hist., Zool. Ser., 18:11, February 12, from Santa Ysabel.

_Specimens examined_ (20 males and 21 females; all in alcohol; ten
crania extracted and cleaned).--Guadalcanal in May, 23863, 23915; Fauro
in April, 23804-5; Malaita in June, 24079; Choiseul in March, 23563-4,
23616, 23627, 23630, 23632-3, 23642, 23658, 23663-4, 23680, 23692-3,
23713, 23722; Kolombangara in January and February, 23343, 23366,
23382-4, 23389-90, 23408-9, 23424, 23455, 23471-4, 23501.

_Measurements._--Average and extreme external measurements of 13 males
and 18 females are, respectively, as follows: Length of head and body,
104.4 (99-118), 108.6 (104-117); tail vertebrae, 16.8 (13-19), 17.6
(15-24); hind foot, 18.0 (16-19), 16.2 (12-18); ear, 15.9 (15-17), 15.0
(14-16); length of forearm, 70.1 (66.0-74.1), 68.1 (65.0-69.1). Average
and extreme measurements of skulls of five males and five females are,
respectively, as follows: Greatest length of skull, 33.2 (33.0-33.7),
31.5 (30.9-32.1); condylobasal length, 31.3 (30.9-31.9), 30.1
(29.3-30.8); palatal length, 14.0 (13.3-14.8), 13.3 (13.0-13.7);
zygomatic breadth, 20.8 (19.8-21.8), 19.4 (18.7-20.8); length of
maxillary tooth-row, 11.0 (10.9-11.3), 10.3 (10.1-10.6); length of
mandibular tooth-row, 12.6 (12.4-12.9), 11.8 (11.7-12.2).

_Remarks._--The specimens from Choiseul, Kolombangara, and Malaita
islands provide new records of distribution for _Rousettus
amplexicaudatus hedigeri_ (Fig. 3). It was described as smaller than _R.
a. brachyotis_ Dobson, which is known from New Guinea, Amboina, and the
Bismarck Archipelago (Pohle, 1953:127-128). Andersen (1912:809) gave the
range of length of forearm in _R. a. brachyotis_ as 73-81, whereas Pohle
(1953:127) gave the length of forearm of the type specimen of _R. a.
hedigeri_ (adult male) as 67. Measurements of specimens examined by me
indicate that _hedigeri_ occurs throughout the Solomon Islands. Cranial
measurements of my specimens and Pohle's type are less than those of _R.
a. brachyotis_ (see Andersen, 1912:48).

Sanborn (1931:11) noted that the forearms of three males examined by him
were longer than that of a female. Mean and range for length of forearm
of males and females listed herein, respectively, are 70.1 (66.0-74.1)
and 68.1 (65.0-69.1). Also, each of seven cranial measurements taken by
me averaged more in males than in females. Sagittal and lambdoidal
crests are more prominent in males than in females.

    TABLE 1. A Summary of Breeding Data for Females of _Rousettus
       amplexicaudatus hedigeri_ Collected December to June.

  ===========+===========+==============+===========+=============
             |   Total   |    Number    |           |  Number of
     MONTH   |  number   | adult [F][F] |  Number   |  immature
             | collected |   collected  | lactating | individuals
  -----------+-----------+--------------+-----------+-------------
  December   |     3     |       3      |     3     |      0
  January    |    11     |      11      |     8     |      0
  February   |     6     |       0      |    --     |      1
  March      |    16     |       1      |     0     |      9
  April      |     2     |       2      |     0     |      0
  June       |     1     |       1      |     0     |      0
  -----------+-----------+--------------+-----------+-------------

As shown in Table 1, adult females obtained in December and January were
lactating when captured whereas those obtained in March, April, and
June were not. More than half of the individuals collected in March were
immature (judging from small size, unfused epiphyses, and lack of wear
on teeth). The immature individuals probably had been nursing in
December and January.



=Pteralopex= Thomas

   1888. _Pteralopex_ Thomas, Ann. Mag. Nat. Hist., ser. 6, 1:155,
         February 1.

   1762. _Pteropus_ Brisson, Regnum animale ..., ed. 2, p. 153.

_Pteralopex_, with one species and two subspecies, is the only
megachiropteran genus endemic to the Solomons. Thomas (1888b:475)
considered this unusual bat a relic, isolated from the time when
pteropodids had cuspidate cheek-teeth. Although two workers (Matschie,
1899:11; Simpson, 1945:54) have synonymized _Pteralopex_ with
_Pteropus_, I regard _Pteralopex_ as a morphologically distinct genus.

Individuals of _Pteralopex_ can be distinguished from all species of
_Pteropus_ in the Solomon Islands by the following features: wing
membranes originate along dorsal midline; braincase diminutive relative
to rest of skull; sagittal crest pronounced; cheek-teeth cuspidate,
broad and massive; i2 about 10 times larger than i1; upper canines with
well-developed secondary cusp; postorbital process fused with zygomatic
arch, forming complete bony ring around orbit.

Andersen (1909a:216; 1912:436) considered the relationships of
_Pteralopex_ and _Pteropus_ and concluded that _Pteropus pselaphon_ Lay,
1829, from the Sulphur Islands east of Taiwan, and _Pteropus samoensis_
Peale, 1848, from the Samoan Islands, were the "closest" living
relatives of _Pteralopex_. He stated further that _Pteralopex_ "presents
in fact scarcely a single character which is not either developed to a
certain extent or at least distinctly foreshadowed in _Pteropus
pselaphon_, _pilosus_, _tuberculatus_, or _leucopterus_." In summary,
Andersen thought several species of _Pteropus_ had undergone
evolutionary development resembling that in _Pteralopex_, and that the
latter, with its massive, cuspidate cheek-teeth, could be considered a
highly modified _Pteropus_. For this hypothesis to be plausible, one
must assume that the originally complex cheek-teeth of pteropodids
became simple and, at least in the case of _Pteralopex_, secondarily
became complex once again. According to present-day theory of
evolutionary development, his hypothesis is improbable. Thomas
(1888b:475) probably was correct when he considered _Pteralopex_ an
isolated relic.

Although _Pteralopex_ usually is listed after _Pteropus_ in phylogenetic
arrangements (see, for example, Sanborn, 1931:21; Pohle, 1953:129;
Laurie and Hill, 1954:40), I have placed _Pteralopex_ before _Pteropus_.


=Pteralopex atrata=

Two subspecies of _Pteralopex atrata_ (_P. a. atrata_ and _P. a.
anceps_) have been named; specimens of both are rare in museum
collections. Thomas (1888_a_:155) described adults of _atrata_. Sanborn
(1931:21) examined the one additional specimen known to me and reported
that it agreed with Thomas' description.

Andersen (1909_b_:266) used a subadult female ("nearly fully grown") as
the holotype of _anceps_. At least five additional specimens, all
adults, of _anceps_ now are housed in various collections. Judging from
these individuals, the holotype of _anceps_ was only four-fifths grown
and because he used an immature individual, Andersen's (1912:437)
criteria for distinguishing the two subspecies mostly are invalid.

    [Illustration: FIG. 4. Distribution of _Pteralopex atrata_;
       _P. atrata atrata_ ([RW]) and _P. atrata anceps_ ([BC]).
       For names of islands see Fig. 2.]


Key to Subspecies of _Pteralopex atrata_

  1.  Length of forearm 139-144 mm.; dorsal surface of distal
      one-fourth of tibia and entire metatarsus naked; known only
      from Guadalcanal and Santa Ysabel islands
                                            _Pteralopex atrata atrata_

  1'. Length of forearm 162-166 mm.; dorsal surface of distal
      one-fourth of tibia and entire metatarsus furred; known only
      from Bougainville and Choiseul islands
                                            _Pteralopex atrata anceps_


=Pteralopex atrata atrata= Thomas

   1888. _Pteralopex atrata_ Thomas, Ann. Mag. Nat. Hist., ser. 6,
         1:155, February, type from Guadalcanal; 1888, Thomas, Proc. Zool.
         Soc. London, p. 475, December 4; 1896, Heude, Mém. Hist. Nat.
         Emp. China, 3:179; 1897, Trouessart, Catalogus Mammalium ...,
         1:83; 1907, Miller, Bull. U. S. Nat. Mus., 57:60, June 29; 1912,
         Andersen, Catalogue of the Chiroptera ... British Museum, 1:439;
         1931, Sanborn, Publ. Field Mus. Nat. Hist., Zool. Ser., 18:21,
         February 12, from Santa Ysabel.

   1954. _Pteralopex atrata atrata_, Laurie and Hill, List of land
         mammals of New Guinea, Celebes and adjacent islands, p. 40,
         June 30.

   1899. _Pteropus (Pteralopex) atrata_, Matschie, Die Megachiroptera
         ... naturkunde, p. 11; 1904, Trouessart, Catalogus Mammalium ...,
         Suppl., p. 49.

_Specimens examined._--None.

_Remarks._--_Pteralopex atrata atrata_ is known from four specimens from
Guadalcanal and one from Santa Ysabel (Sanborn, 1931:21).

Sanborn (_loc. cit._) reported that a specimen wounded at night, while
feeding on young green coconuts, was the only fruit bat that attempted
to attack the collectors. Troughton (1936:348) has suggested, on the
basis of his experiences with _Pteropus_, that this behavior probably
was a reaction from fear rather than an indication of general
aggressiveness on the part of _Pteralopex_.


=Pteralopex atrata anceps= Andersen

   1909. _Pteralopex anceps_ Andersen, Ann. Mag. Nat. Hist., ser. 8,
         3:266, March, type from Bougainville; 1912, Andersen, Catalogue
         of the Chiroptera ... British Museum, 1:437; 1936, Troughton,
         Rec. Australian Mus., 14:348, April 7; 1953, Pohle,
         Z. Säugetierk., 17:129, October 27.

   1954. _Pteralopex atrata anceps_, Laurie and Hill, List of land
         mammals of New Guinea, Celebes and adjacent islands, p. 40,
         June 30.

_Specimens examined_ (three males, two females; one skull-only and one
in alcohol).--Choiseul in March, 23682; Bougainville in July, USNM
276973-74, USNM 276928, USNM 277112.

_Measurements._--Measurements of three males and one female are,
respectively, as follows: Length of head and body, 280, 271, 261, 255;
hind foot, 50, 54, 52, 59; ear, 23, 23, 26, 22; length of forearm, 160,
162, 166, 171; greatest length of skull, 77.6, 77.9, 78.9, 77.0;
condylobasal length, 74.3, 74.3, 75.5, 73.8; zygomatic breadth, 42.2,
45.4, 43.1, 42.6; breadth across upper canines, 18.7, 21.1, 19.0, 19.0;
breadth across first upper molars, 22.2, 25.3, 22.9, 22.0; length of
maxillary tooth-row, 29.3, 29.8, 28.9, 28.2; length of mandibular
tooth-row, 32.8, 32.8, 32.1, 31.4.

_Remarks._--Heretofore, _Pteralopex atrata anceps_ was not known from
Choiseul. The specimen from that island agrees well with specimens in
the U. S. National Museum from Cape Torokina, Bougainville.

The type specimen of this subspecies is a subadult and is smaller than
the specimens examined by me; Andersen (1912:440) gave length of forearm
of the type as 137 (as opposed to 164 in adults). He (1912:438) figured
the dentition of _anceps_ and described the ways in which it differed
from the dentition of _atrata_. Although he (1912:437) concluded that
_anceps_ and _atrata_ represented "two stages of specialization of ...
dentition," there apparently are few, if any, dental differences between
the two subspecies. Teeth of adults of _anceps_ differ from teeth of the
immature type of _anceps_ as follows: in adults the anterior basal ledge
of P4 extends onto the labial surface, whereas in the type it does not;
and maxillary and mandibular teeth in adults are spaced as in the
subspecies _atrata_ (see Andersen, 1912:438, fig. 22) and not crowded as
in the type of _anceps_. Distance between individual cheek-teeth
apparently increases with growth of the cranium and mandible.

Adults of _P. a. anceps_ that I examined are darker than the subadult
type. The mantle in these adults is black, whereas it is seal-brown in
the type (Andersen, 1912:439).

An adult female was lactating when obtained on Bougainville in July
(USNM 276928).


Key to _Pteropus_ in the Solomon Islands

  1.       Premolars having distinct basal ledges;
           molars 2.5-4.0 wide                                       2

  1'.      Premolars lacking definite basal ledges;
           molars 1.0-2.4 wide                                      14

  2(1).    Rostrum unshortened (orbit to anterior tip of nasals
           about one-third greatest length of skull); dorsal
           surface of tibiae nearly naked                            3

  2'.      Rostrum shortened (orbit to anterior tip of nasals less
           than one-third greatest length of skull); dorsal surface
           of tibiae usually at least partially furred               8

  3(2).    Forearm more than 128                                     4

  3'.      Forearm less than 128                                     5

  4(3).    Forearm about 155; venter and dorsum nearly black,
           mantle pale yellow            =P. tonganus geddiei=, p. 798

  4'.      Forearm 128-136; venter and dorsum near Mars Brown,
           mantle Ochraceous or Cream-Buff
                                       =P. hypomelanus luteus=, p. 796

  5(3').   Mantle dark, russet or cinnamon, not strongly
           contrasting with color of back
                                  =P. admiralitatum solomonis=, p. 796

  5'.      Mantle pale, Ochraceous-Buff or Cream-Buff, strongly
          contrasting  with color of back                            6

  6(5').   Mantle Ochraceous-Orange to Ochraceous-Buff, hairs pale
           basally; forearm 108-111  =P. admiralitatum goweri=, p. 797

  6'.      Mantle Ochraceous to Cream-Buff, but hairs dark brown
           basally; forearm 110-122                                  7

  7(6').   Length of forearm 110-112
                                    =P. admiralitatum colonus=, p. 796

  7'.      Length of forearm about 122           =P. howensis=, p. 797

  8(2').   Forearm more than 145                                     9

  8'.      Forearm less than 144                                    12

  9(8).    Forearm more than 162                                    10

  9'.      Forearm less than 162                                    11

  10(9).   Forearm 167-173                =P. rayneri grandis=, p. 801

  10'.     Forearm about 164             =P. rayneri rubianus=, p. 802

  11(9').  Flanks and lower belly brightly colored, Burnt Sienna
           to Sanford's Brown; forearm less than 150
                                        =P. rayneri monoensis=, p. 803

  11'.     Flanks and lower belly darker, near tawny; forearm
           more than 150,              =P. rayneri lavellanus=, p. 802

  12(8').  Pelage of dorsum tricolored; rump brightly colored;
           forearm 139-141,               =P. rayneri rayneri=, p. 800

  12'.     Pelage of dorsum bicolored; rump dark; forearm
           less than 135                                            13

  13(12'). Mantle tawny with some Ochraceous-Buff; forearm
           about 130,                    =P. rayneri rennelli=, p. 804

  13'.     Mantle russet, lacking Ochraceous-Buff; forearm
           about 121,                    =P. rayneri cognatus=, p. 803

  14(1').  Forearm more than 131; dorsum Tawny Olive
                                                =P. mahaganus=, p. 806

  14'.     Forearm less than 100; dorsum dark brown
                                                =P. woodfordi=, p. 804


=Pteropus= Brisson

   1762. _Pteropus_ Brisson, Regnum animale ..., ed. 2, p. 153.

_Remarks._--More species (seven) and subspecies (12) of _Pteropus_ occur
in the Solomon Islands than of any other chiropteran genus. Other kinds
of _Pteropus_, as yet unknown, may live there.

The relationships among the species of these large fruit-eating bats,
commonly termed "flying foxes," are obscure and the genus is in need of
revision. The basic, definitive work is still that of Andersen (1912).
Tate (1942) and Felten (1964_a_, 1964_b_) have offered some additional
remarks but groupings and suggested relationships of species of
_Pteropus_ almost entirely are the products of Kund Andersen. According
to present-day concepts of variation and speciation, Andersen's criteria
are artificial.

Basically, there are three "species-groups" of _Pteropus_ in the Solomon
Islands. The first is composed of species in which the rostrum is
"unshortened" (its length about one third of greatest length of skull),
and the cheek-teeth are of moderate size (M1 is 2.8-3.2 wide). The
species are _P. hypomelanus_, _P. admiralitatum_, _P. tonganus_, and _P.
howensis_. The first and second species were placed in the _Pteropus
hypomelanus_ group by Andersen (1912:98).

In the second group the rostrum is "shortened" (its length less than one
third of greatest length of skull) and the cheek-teeth are of moderate
to large size (M1 3.3-4.1 wide). _Pteropus rayneri_, endemic to the
Solomons and represented there by at least seven subspecies, fits into
this category.

The third group is represented by _P. mahaganus_ and _P. woodfordi_.
Both species are endemic to the Solomon Islands. In these species the
rostrum is unshortened but the cheek-teeth are greatly reduced,
especially in width (M1 is 1.0-2.2 wide). Both _P. mahaganus_ and _P.
woodfordi_ can be included in the _Pteropus scapulatus_ group of
Andersen (1912:402).


=Pteropus hypomelanus=

_Pteropus hypomelanus_ is a wide-ranging species of flying fox having at
least seven subspecies; three occur in southeastern Asia, two on and
near Celebes, and two in New Guinea and islands adjacent to the
southeastern coast of New Guinea, including one island in the Solomons
(Ellerman and Morrison-Scott, 1966:95; Laurie and Hill, 1954:32-33).

    [Illustration: FIG. 5. Distribution of _Pteropus hypomelanus luteus_
       ([TW]), _Pteropus admiralitatum solomonis_ ([RW]), _Pteropus a.
       colonus_ ([BW]), _Pteropus a. goweri_ ([LW]), _Pteropus tonganus
       geddiei_ ([RTW]), and _Pteropus howensis_ ([BC]). For names of
       islands see Fig. 2.]


=Pteropus hypomelanus luteus= Andersen

   1908. Pteropus hypomelanus luteus Andersen, Ann. Mag. Nat. Hist.,
         ser. 8, 2:362, October, type from Kiriwini Island, Trobriand
         Islands; 1912, Andersen, Catalogue of the Chiroptera ... British
         Museum, 1:128; 1947, Sanborn and Beecher, Jour. Mamm., 28:388,
         November 19, from Banika Island, Russell Islands.

_Specimens examined._--None.

_Remarks._--Andersen (1908:362) identified specimens of _Pteropus
hypomelanus_ from eastern New Guinea and three nearby islands (Conflict
Islands, Trobriand Islands, and Woodlark Island) as _P. hypomelanus
luteus_. Sanborn and Beecher (1947:388) identified a female from Banika
Island in the Solomons as of this subspecies although this specimen was
darker and had a slightly smaller skull than typical _P. hypomelanus
luteus_. They noted that the pelage of the venter of the female was
uniformly dark rather than the typical Ochraceous-Buff to Cream-Buff;
the specimen was regarded as a dark phase of the subspecies. Although
not recorded previously for _luteus_, other subspecies of _P.
hypomelanus_ were known in dark phase as well as pale and intermediate
phases of coloration (Andersen, 1912:122). The reported occurrence of
_P. h. luteus_ on Banika Island extended the known geographic range
about 450 miles eastward from Woodlark Island.


=Pteropus admiralitatum=

Three subspecies, all about the same size but differing in coloration,
have been described from the Solomon Islands. _P. a. goweri_ is known
only from Gower (Ndai) Island, notably removed from the western chain of
islands inhabited by _P. a. colonus_ and _P. a. solomonis_. Only one
other subspecies, from the Admiralty Islands, is known.


=Pteropus admiralitatum solomonis= Thomas

   1904. _Pteropus solomonis_ Thomas, Novit. Zool., 11:597, type from
         Ghizo Island; 1912, Andersen, Catalogue of the Chiroptera ...
         British Museum, 1:149; 1931, Sanborn, Publ. Field Mus. Nat. Hist.,
         Zool. Ser., 18:12, February 12, from Ronongo (Ganongga), Vella
         Lavella, and Narovo (Simbo) islands; 1947, Sanborn and Beecher,
         Jour. Mamm., 28:389, November 19, from Banika and Guadalcanal
         islands.

   1954. _Pteropus admiralitatum solomonis_, Laurie and Hill, List of
         land mammals of New Guinea, Celebes and adjacent islands, p. 33,
         June 30.

_Specimens examined._--None.

_Remarks._--Andersen (1912:149) considered _Pteropus admiralitatum_, and
especially the subspecies _P. a. solomonis_, to be the easternmost
"representative" of _Pteropus hypomelanus_. In comparison with _P.
hypomelanus luteus_, _P. a. solomonis_ differs mostly in size, being
much smaller (length of forearm about 110 rather than 134). It is now
known that both species occur on Banika Island in the Solomons.

The subspecies _P. a. solomonis_ has been recorded from a "chain" of
islands that included Vella Lavella, Simbo, Ghizo, Ganongga, Banika, and
Guadalcanal (see Fig. 5).


=Pteropus admiralitatum colonus= Andersen

   1908. _Pteropus colonus_ Andersen, Ann. Mag. Nat. Hist., ser. 8,
         2:363, October, type from Shortland Island; 1912, Andersen,
         Catalogue of the Chiroptera ... British Museum, 1:150; 1931,
         Sanborn, Publ. Field Mus. Nat. Hist., Zool. Ser., 18:12,
         February 12, from Mono Island.

   1954. _Pteropus admiralitatum colonus_, Laurie and Hill, List of
         land mammals of New Guinea, Celebes and adjacent islands, p. 33,
         June 30.

   1887. _Pteropus hypomelanus_ (part), Thomas, Proc. Zool. Soc.
         London, p. 471, December 4; 1898, Trouessart, Catalogus Mammalium
         ..., 1:82, from "I. Salomonis."

   1899. _Pteropus (Spectrum) hypomelanus_ (part), Matschie, Die
         Megachiroptera ... naturkunde, p. 24.

_Specimens examined._--None.

_Remarks._--_Pteropus admiralitatum colonus_ is the largest of the three
subspecies that occur in the Solomon Islands. It closely resembles _P.
hypomelanus luteus_, except in being smaller throughout (see Andersen,
1912:151-152, for measurements) and darker on the underparts.

This bat has been found in a group of small islands (Alu, Mono, and
Shortland) about 30 miles south of Bougainville. Because of this
proximity and because yet another subspecies of this species occurs
northward of Bougainville, it is interesting that neither Troughton
(1936) nor Pohle (1953) included the species in their faunal lists for
Bougainville.

Andersen (1912:152) indicated that the M1 in _P. admiralitatum colonus_
is smaller than in _P. a. solomonis_, the subspecies found in islands to
the southeast (4.4-4.5 and 5.2, respectively), but Sanborn (1931:13)
studied specimens of these two subspecies that overlapped in size of M1.


=Pteropus admiralitatum goweri= Tate

   1934. _Pteropus goweri_ Tate, Amer. Mus. Novit., 718:1, May 4, type
         from Gower (Ndai) Island.

   1954. _Pteropus admiralitatum goweri_, Laurie and Hill, List of
         land mammals of New Guinea, Celebes and adjacent islands, p. 33,
         June 30.

_Specimens examined._--None.

_Remarks._--_Pteropus admiralitatum goweri_ was described from six
specimens collected in 1930 by the Whitney South Sea Expedition (Tate,
1934:1). This subspecies closely resembles the other two subspecies of
_P. admiralitatum_ (_colonus_ and _solomonis_) found in the Solomon
Islands. Color and length of forearm (see key on p. 793) seem to be the
only reliable criteria for distinguishing between these subspecies. The
longitude of Gower Island, 160° 34' E, was incorrectly listed in Laurie
and Hill (1954:152) as 159° 34' E.


=Pteropus howensis= Troughton

   1931. _Pteropus howensis_ Troughton, Proc. Linn. Soc. New South
         Wales, 56:204, June 24, type from Lord Howe Islands (Ontong Java);
         1950, Sanborn and Nicholson, Fieldiana:Zool., 31:329, August 31.

_Specimens examined_ (one male, three females, and two sex unknown; two
embryos in alcohol).--Liuniuwu, Lord Howe Islands (Ontong Java) in
August, USNM 278703-6, USNM 279715-6.

_Measurements._--Average and extreme measurements of one male and three
females are as follows: Length of head and body, 185.2 (176-196); hind
foot, 34.5 (33-36); ear, 21.5 (21-23); forearm not measured [broken in
all specimens examined]. Cranial measurements of a male and a female
are, respectively, as follows: Greatest length of skull, 55.3, 53.8;
condylobasal length, 54.2, 52.8; palatal length, 26.7, 26.0; zygomatic
breadth, 30.6, 29.9; breadth of braincase, 19.9, 19.2; breadth across
first upper molars, 14.3, 14.3; length of maxillary tooth-row, 20.7,
19.6; length of mandibular tooth-row, 23.1, --.

_Remarks._--Apparently _Pteropus howensis_ is confined to Ontong Java
(Lord Howe Islands) located northeastward of the main body of islands
that constitute the Solomon Archipelago (see Fig. 5). According to A. J.
Nicholson, who collected the specimens listed above, _P. howensis_ is
not abundant in Ontong Java. He related this circumstance to the fact
that these small islands are nothing more than parts of a coral atoll
used almost entirely for the production of coconuts (see Sanborn and
Nicholson, 1950:329).

Specimens of _Pteropus howensis_ deposited in the U. S. National Museum
agree well in most ways with the original description of the species by
Troughton (1931:204-205). Slight variation in color is evident; in two
specimens, the mantle, just posterior to the ears, is Ochraceous-Buff.

The relationship of this species to other kinds of _Pteropus_ known from
Melanesia is not clear. Troughton (1931:204, 206) compared _P. howensis_
with _P. hypomelanus_ and _P. admiralitatum_ and found that it resembled
each of them. Tate (1934:2) noted that the skull of _P. admiralitatum
goweri_ was similar to that of _P. howensis_ in structure. The latter
species is, however, larger (length of forearm 122 according to
Troughton, 1931:205) than any subspecies of _P. admiralitatum_ (length
of forearm 108-112). Also, the cheek-teeth of _P. howensis_ that I have
studied are relatively larger than those of either _P. hypomelanus_ or
_P. admiralitatum_. Furthermore, in _P. howensis_ there is a small but
distinct cusp located medio-posteriorly on P4 (most noticeable in young
individuals) that is more reduced or undeveloped in specimens of the
other two species. Cheek-teeth of _P. howensis_ resemble those in a
dull-colored specimen of _P. tonganus_ from Fiji Island with which I
compared the specimens listed above.

Weights and crown-rump lengths of the two embryos (in an advanced stage
of development) examined were 20 and 29 grams and 43 and 51 mm.
(apparently these are the specimens listed by Sanborn and Nicholson,
1950:329).


=Pteropus tonganus=

_Pteropus tonganus_ has at least three subspecies, one of which has been
recorded from the Solomons. The species ranges from a small island off
the eastern coast of New Guinea, where there is an endemic subspecies,
eastward to Tonga and the New Hebrides (Laurie and Hill, 1954:33-34).
Felten (1964_a_) recently has reported on the species in the New
Hebrides.


=Pteropus tonganus geddiei= MacGillivary

   1860. _Pteropus geddiei_ MacGillivary, Zoologist, 18:7134,
         September, type from Aneitum Island, New Hebrides; 1912,
         Andersen, Catalogue of the Chiroptera ... British Museum,
         1:189; 1931, Sanborn, Publ. Field Mus. Nat. Hist., Zool.
         Ser., 18:13, February 12, from Rennell Island in the Solomons.

   1914. _Pteropus tonganus geddiei_, Revilliod, _in_ Sarasin and
         Roux, Nova Caledonia (A), 1:341; 1954, Laurie and Hill, List
         of land mammals of New Guinea, Celebes and adjacent islands,
         p. 34, June 30.

_Specimens examined._--None.

_Remarks._--_Pteropus tonganus geddiei_, as far as is known, is the
widest ranging subspecies of this genus. It is the only megachiropteran
in the Solomon Islands having affinities with bats to the southeast (the
New Hebrides, Santa Cruz Islands, Samoan Islands and Fiji Islands)
rather than with those to the west (New Guinea). The subspecies _P.
tonganus geddiei_, which ranges from the Solomons to the New Hebrides
(about 500 miles straight-line distance), is said to be remarkably
uniform throughout its range. Sanborn (1931:14) compared color and size
in specimens from the Solomon Islands and the New Hebrides and found
little variation. Another subspecies, _P. t. bascilicus_ Thomas 1915,
apparently closely related to _geddiei_, is known from Dampier [=
Kar-kar] Island off the northeastern coast of New Guinea and therefore
farther westward from the New Hebrides than are the Solomon Islands.
Additional remarks on the distribution of this species are in the
section on Zoogeography and Speciation.


=Pteropus rayneri=

_Pteropus rayneri_ is endemic to the Solomon Islands. It is divisible
into seven subspecies (see Fig. 6), which, excepting _P. r. rennelli_
and _P. r. cognatus_, are strikingly colored--the mantle, back, and rump
being of different colors. Differences in color and size provide
characters differentiating the subspecies (see key, p. 793). Recorded
lengths of forearms do not overlap between any two subspecies. _P. r.
grandis_, northernmost in distribution, has the longest (about 170)
forearm and _P. r. cognatus_, known from two of the southernmost
islands, has the shortest (about 121).

    [Illustration: FIG. 6. Distribution of _Pteropus rayneri_: _P. r.
       rayneri_ ([RTW]); _P. r. grandis_ ([RW]); _P. r. lavellanus_ ([BC]);
       _P. r. monoensis_ ([BW]); _P. r. rubianus_ ([TW]); _P. r. cognatus_
       ([LW]); _P. r. rennelli_ ([LTW]). For names of islands see Fig. 2.]

Adult males of _Pteropus rayneri_ have well-developed tufts of hair on
each side of the neck where a gland is located (see Andersen,
1912:259). Apparently these glands are not present in females as none
were found in specimens studied by me or those reported by Sanborn
(1931:16). Evidently, these glands are associated with sexual maturity
in males because neither Sanborn nor I found them in subadult males.


=Pteropus rayneri rayneri= Gray

   1870. _Pteropus rayneri_ (part), Gray, Catalogue of monkeys, lemurs
         and fruit-eating bats ... British Museum, p. 108, cotypes from
         Guadalcanal; 1878, Dobson, Catalogue of the Chiroptera ...
         British Museum, p. 33; 1879, Trouessart, Rev. Mag. Zool., 6:204;
         1879, Trouessart, Ann. Sci. Nat. Zool, 8:16; 1887, Thomas, Proc.
         Zool. Soc. London, p. 322, March 15; 1888, Thomas, Proc. Zool.
         Soc. London, p. 472, December 4; 1898, Trouessart, Catalogus
         Mammalium ..., 1:78; 1912, Andersen, Catalogue of the Chiroptera
         ... British Museum, p. 254; 1931, Sanborn, Publ. Field Mus. Nat.
         Hist., Zool. Ser., 18:15, February 12, from Guadalcanal
         and Malaita.

   1954. _Pteropus rayneri rayneri_, Laurie and Hill, List of land
         mammals of New Guinea, Celebes and adjacent islands, p. 35,
         June 30.

   1899. _Pteropus (Spectrum) rayneri_ (part), Matschie, Die
         Megachiroptera ... naturkunde, p. 22; 1904, Trouessart,
         Catalogus Mammalium ..., Suppl., p. 51.

_Specimens examined_ (four males and one female; one embryo in
alcohol).--Guadalcanal in July and November, USNM 278700-02, USNM
278142, USNM 278714.

_Measurements._--Measurements of three males and one female are,
respectively, as follows: Length of head and body, --, 210, 214, 215;
hind foot, --, 33, 39, 42; ear, --, 23, 23, 23; length of forearm, --,
138, 136, 134; greatest length of skull, 61.5, 59.2, 61.6, 61.2;
condylobasal length, 61.4, 58.2, 60.3, 60.0; zygomatic breadth, 36.6,
35.3, 35.4, 36.5; breadth of braincase, 23.7, 22.5, 22.6, 24.1; breadth
across first upper molars, --, 16.9, 16.7, 16.8; width of M1, 3.4, 3.5,
3.5, 3.5; length of maxillary tooth-row, 22.4, 22.1, 23.6, 23.2; length
of mandibular tooth-row, 26.4, 25.5, 25.9, 25.6.

_Remarks._--_Pteropus rayneri_ was named on the basis of two specimens
(cotypes) obtained on Guadalcanal and listed as "male" and "female";
according to Andersen (1912:254), however, both are females.

_P. r. rayneri_ is known from Guadalcanal and Malaita (see Fig. 6), and
is of almost the same size as _P. r. cognatus_, which is known from San
Cristobal and Ugi, only about 40 miles to the southeast. In the latter
subspecies the back and rump are the same color (Prouts Brown), whereas
in _P. r. rayneri_ the rump is brightly colored and therefore contrasts
strongly with the dark brown back. A specimen of _rayneri_ from Malaita
was reported by Sanborn (1931:15) as unusually small and having a
dark-colored rump patch. In the specimens examined from Guadalcanal,
there is noticeable variation in color of the mantle that does not seem
related to age or sex. In two specimens (adult male and female) the
mantle is Cinnamon-Rufous tinged with Russet, strongly contrasting with
the crown, which is Ochraceous-Tawny and has scattered silvery hairs.
Another specimen has a darker mantle (near Chestnut-Brown) and a crown
of about the same color, but with a few scattered Ochraceous-Tawny
hairs.

The skull of one adult male bears an extra peglike tooth posterior to M3
on the right side.

An embryo, in an advanced stage of development, in the collection of the
U. S. National Museum, measures: Length of head and body, 98; hind foot,
30; ear, 8.5; length of forearm, 48 (this may be the same specimen
listed by Sanborn and Nicholson, 1950:329).


=Pteropus rayneri grandis= Thomas

   1887. _Pteropus grandis_ Thomas, Ann. Mag. Nat. Hist., ser. 5,
         19:147, March, type from Shortland; 1887, Thomas, Proc. Zool.
         Soc. London, p. 320, March 15, from Alu and Shortland; 1897,
         Trouessart, Catalogus Mammalium ..., 1:80, from "I. Salomonis";
         1899, Matschie, Die Megachiroptera ... naturkunde, p. 15; 1904,
         Trouessart, Catalogus Mammalium ..., Suppl., p. 49; 1907, Miller,
         Bull. U. S. Nat. Mus., 57:58, June 29; 1912, Andersen, Catalogue
         of the Chiroptera ... British Museum, 1:259, from Bougainville;
         1931, Sanborn, Publ. Field Mus. Nat. Hist., Zool. Ser., 18:16,
         February 12, from Choiseul, and Santa Ysabel; 1936, Troughton,
         Rec. Australian Mus., 19:348, April 7; 1953, Pohle, Z.
         Säugetierk., 17:128, October 27.

   1954. _Pteropus rayneri grandis_, Laurie and Hill, List of land
         mammals of New Guinea, Celebes and adjacent islands, p. 35,
         June 30.

_Specimens examined_ (six males and 10 females; five in
alcohol).--Choiseul in March, 23580, 23644, 23593; Bougainville in July,
August, September, and October, USNM 276926-7, USNM 276968, USNM
277091-9.

_Measurements._--Average and extreme measurements of four males and
seven females are as follows: Length of head and body, 281 (260-302);
hind foot, 52.3 (50-58); ear, 33.1 (31-37); length of forearm, 173
(168-180). Average and extreme measurements of skulls of three males and
six females are as follows: Greatest length of skull, 73.7 (71.3-77.7);
condylobasal length, 73.1 (70.5-77.4); zygomatic breadth, 40
(36.4-41.5); breadth across first-upper molars, 20.9 (18.3-22.1); length
of maxillary tooth-row, 28.1 (26.9-29.9); length of mandibular
tooth-row, 31.8 (29.7-32.7).

_Remarks._--_Pteropus rayneri grandis_ is the largest subspecies of the
species. It is also the widest ranging subspecies, being found on six
islands (see Fig. 6).

Although the specimens listed above agree well with descriptions of
color given by Thomas (1887_a_:147) and Andersen (1912:259, 263-264),
some individual variation is noticeable. In bats not yet fully grown
(judging from small size, unfused epiphyses, and lack of wear on teeth),
numerous scattered hairs on the sides of the face and crown are buffy.
In adults the face and crown are blackish. With regard to individual
variation in color of mantle and rump patch, specimens with the
following combinations were noted (1) mantle Brick Red, rump patch
bright, basal three-quarters of hairs white, tips Warm Buff (2) mantle
darker, near Hessian Brown, rump patch dark, Chestnut along edges,
center Ochraceous-Tawny (3) mantle Brick Red, rump patch intermediate
between the two other types. Size of rump patch also is variable. In
some specimens it extends onto the upper parts of the thighs whereas in
other specimens it does not.

Sanborn (1931:16) reported an extra tooth, behind the last lower molar,
in a specimen from Choiseul. In one of three specimens in the Bishop
Museum, m3 is lacking. Judging from Troughton's (1936:346) remarks, size
of individuals varies considerably. Specimens that he examined from
Bougainville had longer forearms (up to 177) and larger hind feet
(54-57) than those examined by me from Choiseul. On the other hand,
specimens listed above from Bougainville agree well with those from
Choiseul. In many specimens in the U. S. National Museum, length of the
right- and left-forearm differ. For example, in No. 276926 the right
forearm measures 180 whereas the left is 174; in No. 277098 the right is
172 and the left is 167. Troughton (1936:346) gave standard ear
measurement in _P. r. grandis_ as ranging from 29.5 to 31.5. Ears of
specimens that I examined varied from 31.0 to 37.0.


=Pteropus rayneri rubianus= Andersen

   1908. _Pteropus rubianus_ Andersen, Ann. Mag. Nat. Hist., ser. 8,
         2:366, October, type from Rubiana; 1912, Andersen, Catalogue
         of the Chiroptera ... British Museum, 1:255; 1931, Sanborn,
         Publ. Field Mus. Nat. Hist., Zool. Ser., 18:15, February 12,
         from Narovo (Simbo).

   1954. _Pteropus rayneri rubianus_, Laurie and Hill, List of land
         mammals of New Guinea, Celebes and adjacent islands, p. 35,
         June 30.

   1888. _Pteropus grandis_ (part), Thomas, Proc. Zool. Soc. London,
         p. 470, December 4, from Rubiana; 1899, Matschie, Die
         Megachiroptera ... naturkunde, p. 15; 1904, Trouessart,
         Catalogus Mammalium ..., Suppl., p. 49.

_Specimens examined_ (two males and one female).--Kolombangara, in
February, 23458-60.

_Measurements._--Measurements of two males and one female are,
respectively, as follows: Length of head and body, 253, 265, 251; hind
foot, 53, 50, 50; ear, 30, 31, 32; length of forearm, 158, 161, 160;
greatest length of skull, 70.2, 67.4, --; condylobasal length, 67.0, --,
68.4; zygomatic breadth, 40.0, 39.4, 40.7; breadth across first upper
molars, 19.4, 20.4, 19.9; length of mandible, 53.9, 49.4, 51.3.

_Remarks._--Kolombangara Island is a new locality for _Pteropus rayneri
rubianus_; heretofore this subspecies was known only from Rubiana and
Narovo islands (Andersen, 1908:366; Sanborn, 1931:15). The coloration of
a specimen from Narovo Island was described as between that of _P. r.
rubianus_ and _P. r. lavellanus_. Sanborn (1931:16) allocated it to the
subspecies _rubianus_ on the basis of length of forearm.

Andersen's descriptions (1908:366; 1912:256) of _rubianus_ were of a
specimen stored in alcohol. Coloration of the museum skins examined by
me is as follows: Dorsum from shoulders to rump near Vandyke Brown;
crown and mantle Brick Red; face close to Mummy Brown; rump patch and
thighs close to Warm Buff, strongly contrasting with back and mantle;
base of hairs dark, Seal Brown; venter dark; chest about same as back
but paler laterally (to Ochraceous Tawny); throat Brick Red.


=Pteropus rayneri lavellanus= Andersen

   1908. _Pteropus lavellanus_ Andersen, Ann. Mag. Nat. Hist., ser. 8,
         2:366, October, type from Vella Lavella; 1912, Andersen,
         Catalogue of the Chiroptera ... British Museum, 1:259; 1931,
         Sanborn, Publ. Field Mus. Nat. Hist., Zool. Ser., 18:16,
         February 12, from Ghizo and Ronongo.

   1954. _Pteropus rayneri lavellanus_, Laurie and Hill, List of land
         mammals of New Guinea, Celebes and adjacent islands, p. 36,
         June 30.

_Specimens examined_ (one male and one female).--Vella Lavella in
November, 23192, 23142.

_Measurements._--Measurements of a male and a female are, respectively,
as follows: Length of head and body, 286, 282; hind foot, 55, 56; ear,
30, 30; length of forearm, 156, 155; greatest length of skull, 72.9,
67.6; condylobasal length, 71.8, 64.2; zygomatic breadth, 38.4, 37.9;
breadth across first upper molars, 19.9, 19.8; length of mandible, 54.6,
50.8.

_Remarks._--_Pteropus rayneri lavellanus_ inhabits islands
geographically near those from which _P. r. rubianus_ is known (see Fig.
6) and in most respects the two subspecies closely resemble each other.
_P. r. lavellanus_ is slightly the smaller (average length of forearm
about 156 instead of 160) and darker. A bat from Narovo [Simbo] Island,
only a few miles from Vella Lavella, identified by Sanborn (1931:16) on
basis of its size as _P. r. rubianus_, resembled the subspecies
_lavellanus_ in color and probably represents an intergrade between the
two populations.

The color of _P. r. lavellanus_ is close to that of _P. r. rubianus_
except that the crown, mantle, and foreneck are near Chestnut-Brown, the
basal portions of hair black, and the fur of the venter, from sternum to
pectoral region, is dark, almost black (compare with description of _P.
r. rubianus_ under account of that subspecies).

Measurements of the male examined are greater than those of the female
studied. Andersen (1912:259) noted that the canine teeth are heavier in
males than in females.


=Pteropus rayneri monoensis= Lawrence

   1945. _Pteropus rayneri monoensis_ Lawrence, Proc. New England
         Zool. Club, 23:63, March 26, type from Mono (Treasury); 1954,
         Laurie and Hill, List of land mammals of New Guinea, Celebes
         and adjacent islands, p. 36, June 30.

_Specimens examined._--None.

_Remarks._--_Pteropus rayneri monoensis_ is the most recently described
subspecies of _P. rayneri_. Lawrence (1945:63) judged that in most ways
this bat is intermediate between _P. r. grandis_ and _P. r. lavellanus_.
Coloration of _monoensis_ indicates affinity with the former, whereas
length of forearm (145-148) approaches that in the latter. The small
skull, narrow palate, and whitish rump patch of _monoensis_ are
differences that distinguish it from _grandis_ and _lavellanus_. The
relatively isolated position of Mono Island may have been important in
establishment of the distinctive features of this bat.

Lawrence (1945:65) quoted a collector as stating: "They [individuals of
_P. r. monoensis_] rest quietly during the day in the tops of
heavy-leaved, tall jungle trees, and start flying about dusk, looking
for feeding spots. There is usually quite a flight for fifteen to twenty
minutes at twilight...."

No additional specimens of this subspecies have been collected on small
adjacent islands and _monoensis_ may therefore be confined to Mono
Island.


=Pteropus rayneri cognatus= Andersen

   1908. _Pteropus cognatus_ Andersen, Ann. Mag. Nat. Hist., ser. 8,
         2:365, October 1, type from San Cristobal; 1912, Andersen,
         Catalogue of the Chiroptera ... British Museum, 1:251; 1931,
         Sanborn, Publ. Field Mus. Nat. Hist., Zool. Ser., 18:15,
         February 12, from San Cristobal and Ugi; 1954, Laurie and Hill,
         List of land mammals of New Guinea, Celebes and adjacent islands,
         p. 35, June 30.

   1962. _Pteropus rayneri cognatus_, Hill, The natural history of
         Rennell Island, British Solomon Islands, 4:9, February.

   1870. _Pteropus rayneri_ (part), Gray, Catalogue of monkeys, lemurs
         and fruit-eating bats ... British Museum, p. 108, from San
         Cristobal; 1878, Dobson, Catalogue of the Chiroptera ... British
         Museum, p. 33.

   1904. _Pteropus_ (_Spectrum_) _rayneri_ (part), Trouessart,
         Catalogus Mammalium ..., Suppl., p. 51.

_Specimens examined._--None.

_Remarks._--Specimens of _Pteropus rayneri cognatus_ first were reported
under the name _Pteropus rayneri_ based on three specimens (one from San
Cristobal and two from Guadalcanal). Because the description was based
mostly on the two specimens from Guadalcanal, the name _rayneri_ is
applicable to the bats from that island. Andersen (1908:365) thought
that specimens that he studied, from San Cristobal, were specifically
distinct from _P. rayneri_ and he proposed the name _Pteropus cognatus_
for them. Later, Hill (1962:9) reduced _cognatus_ to subspecific status
under _P. rayneri_.

Presently _P. r. cognatus_ is known only from San Cristobal and the
small adjacent island of Ugi (see Fig. 6).


=Pteropus rayneri rennelli= Troughton

   1929. _Pteropus rennelli_ Troughton, Rec. Australian Mus., 17:193,
         September 4, type from Rennell Island; 1954, Laurie and Hill,
         List of land mammals of New Guinea, Celebes and adjacent islands,
         p. 35, June 30.

   1962. _Pteropus rayneri rennelli_, Hill, The natural history of
         Rennell Island, British Solomon Islands, 4:7, February.

_Specimens examined._--None.

_Remarks._--Until recently, _Pteropus rayneri rennelli_ was known from
but a single specimen. Hill (1962:7) reported two additional specimens
and pointed out that _P. r. cognatus_ and _P. r. rennelli_ probably
represent the extremes of an east-west cline in size. _P. r. rennelli_
and _P. r. cognatus_ differ from other subspecies of the species in
lacking tricolored pelage on the dorsum, but their short rostrum clearly
indicates affinity with other members of this complex group in the
Solomon Islands (Hill, 1962:8).

The relationship of the subspecies _rennelli_ and _cognatus_ is close,
both geographically and genetically. Longer forearm, longer metacarpals,
and longer mandibular tooth-row serve to differentiate _rennelli_ from
_cognatus_.


=Pteropus woodfordi= Thomas

   1888. _Pteropus woodfordi_ Thomas, Ann. Mag. Nat. Hist., ser. 6,
         1:156, February, type from Guadalcanal; 1888, Thomas, Proc. Zool.
         Soc. London, p. 472, December 4; 1898, Trouessart, Catalogus
         Mammalium ..., 1:78; 1907, Elliot, Field Columbian Mus., Zool.
         Ser., 8:491; 1912, Andersen, Catalogue of the Chiroptera ...
         British Museum, 1:410, from New Georgia and Guadalcanal; 1931,
         Sanborn, Publ. Field Mus. Nat. Hist., Zool. Ser., 18:19, February
         12, from Kolombangara; 1947, Sanborn and Beecher, Jour. Mamm.,
         28:389, November 19, from Banika and Guadalcanal; 1954, Laurie
         and Hill, List of land mammals of New Guinea, Celebes and
         adjacent islands, p. 39, June 30.

   1899. _Pteropus (Sericonycteris) woodfordi_, Matschie, Die
         Megachiroptera ... naturkunde, p. 83; 1904, Trouessart, Catalogus
         Mammalium ..., Suppl., p. 54.

   1945. _Pteropus austini_ Lawrence, Proc. New England Zool. Club,
         23:59, March 26, from Florida.

_Specimens examined_ (four males and three females; five in alcohol and
two skin-onlys).--Fauro, in April, 23727, 23790; Guadalcanal in May and
June, 23823, 23931; Pavuvo (Russell Islands) in August and October, USNM
277887, USNM 283872-3.

_Measurements._--External measurements of two males and two females are,
respectively, as follows: Length of head and body, 152, 128, 132, 155;
hind foot, 29, 26, 31, 28; ear, 16, 14, 14, 17; length of forearm, 79,
76, 86, 90.

_Remarks._--Heretofore, _Pteropus woodfordi_ was known from New Georgia,
Guadalcanal, Kolombangara, and Banika (see Fig. 7); specimens from Fauro
and Pavuvo islands, listed above, provide new northern localities of
record for this species.

Judging by small size and unfused epiphyses, a bat obtained in April and
another obtained in June are subadults. Specimens of adults, examined by
me, agree well with the descriptions of _P. woodfordi_ by Thomas
(1888_a_:156) and Andersen (1912:407-409), but are slightly smaller than
specimens listed by Sanborn and Beecher (1947:389). Color of pelage in
this species seems to vary. Adults seen have a pale head and mantle,
contrasting strongly with the dark back. Andersen (1912:409) and
Lawrence (1945:61) discussed individuals that had scattered silvery
hairs mixed with dark fur dorsally and darker mantles that did not
contrast noticeably with the rest of the dorsum.

Lawrence (1945:389) named _Pteropus austini_ as a new species closely
related to _P. woodfordi_ and other species of the _P. scapulatus_ group
of Andersen (1912:402) and Tate (1942:336). Sanborn and Beecher
(1947:389), studied a series of _P. woodfordi_ from Banika and
Guadalcanal and found that skulls of two subadults agreed well with
cranial characteristics ascribed to _P. austini_, which was based on two
subadults. Lawrence (1945:61) stated also that "the interfemoral
membrane is entirely absent medially in _austini_, while in _woodfordi_
it is present as a barely discernible ridge 8 mm. wide." Andersen
(1912:408) had earlier reported that in the type of _woodfordi_ the
interfemoral membrane was "undeveloped in [the] centre." In 13 adults
(in alcohol) studied by Sanborn and Beecher (1947:389), as well as in
adults examined by me, the uropatagium is not present. In size, however,
these specimens agree with dimensions given for _woodfordi_ by Thomas
(1888_a_:156) and Andersen (1912:410); for example, length of forearm is
93-99. According to Lawrence (1945:59) _austini_, in which the
interfemoral membrane is lacking, is smaller than _woodfordi_ and has a
forearm of about 84. In two juveniles of _P. woodfordi_ in the U. S.
National Museum, the medially-developed interfemoral membrane is about 7
wide. One specimen has small but distinct calcars whereas the other
(slightly larger) apparently lacks calcars. This suggests individual
variation in the presence or absence, as well as in the size, of the
uropatagium in _Pteropus woodfordi_.

Sanborn and Beecher (1947:389) decided that "until fully adult specimens
showing the characters of _austini_ are available, it best be considered
a synonym of _woodfordi_." For the following reasons I agree with these
authors: (1) _austini_ is known from only two specimens, both of which
are apparently subadults; (2) _austini_ is reported to have a forearm 84
long and no interfemoral membrane, whereas _woodfordi_ has a forearm
about 96 long and an interfemoral membrane that is only slightly
developed; (3) specimens that agree in size and cranial characters with
the type of _woodfordi_ but that lack an interfemoral membrane have been
obtained; and (4) skulls of subadults of _woodfordi_ agree with the
description of skulls of _austini_.

Sanborn (1931:19) reported that specimens of _Pteropus woodfordi_ were
obtained at night, while feeding on young green coconuts. Lawrence
(1945:62) reported that in the late afternoon a collector found
individuals of _austini_ [= _woodfordi_] in the fronds of a coconut
tree, apparently feeding on pollen shoots. Sanborn and Beecher
(1947:388) have reported malaria (_Plasmodium_) in _P. woodfordi_
obtained on Guadalcanal. They suggested that malaria might have rendered
one individual helpless because when it was found, on the ground, no
wounds were evident and parasites were present in the blood.

    [Illustration: FIG. 7. Distribution of _Pteropus woodfordi_ ([BW])
       and _P. mahaganus_ ([BC]). For names of islands see Fig. 2.]


=Pteropus mahaganus= Sanborn

   1931. _Pteropus mahaganus_ Sanborn, Publ. Field Mus. Nat. Hist.,
         Zool. Ser., 18:19, February 12, type from Santa Ysabel, also
         reported from Bougainville; 1954, Laurie and Hill, List of
         land mammals of New Guinea, Celebes and adjacent islands,
         p. 39, June 30.

_Specimens examined_ (one male and two females; one in
alcohol).--Bougainville, in August and October, USNM 276972, USNM
277104-5.

_Measurements._--Measurements of one male and two females are,
respectively, as follows: Length of head and body, 180, 204, 198; hind
foot, 42, 38, 44; ear, 25, 23, 22; length of forearm, 134, 138, 140.
Measurements of the skull of the male and one female are, respectively,
as follows: Greatest length of skull, 52.5, 55.8; condylobasal length,
50.9, 54.3; palatal length, 24.1, 26.0; zygomatic breadth, 28.9, 32.5;
breadth across first upper molars, 14.4, 15.0; width of M1, 2.2, 2.2;
length of maxillary tooth-row, 17.4, 18.4; length of mandibular
tooth-row, 20.1, 21.4.

_Remarks._--Sanborn (1931:19-21) described _Pteropus mahaganus_ on basis
of six specimens, five from Santa Ysabel and one from Bougainville. The
latter was in poor condition and only provisionally allocated to this
species. The specimens examined by me (listed above) confirm the
occurrence of _P. mahaganus_ on Bougainville.

Sanborn (1931:20) described _mahaganus_ as "similar to and about the
size of [_Pteropus scapulatus_] from Australia, but lighter in color,"
and considered it, along with _P. woodfordi_, a member of the _Pteropus
scapulatus_ group of Andersen (1912:402) and Tate (1942:336). I would
judge, however, that _P. mahaganus_ and _P. woodfordi_ are much more
closely related to one another than to _P. scapulatus_ of Australia. The
only significant characteristic that the latter has in common with the
two species from the Solomons is small cheek-teeth. In fact, teeth of
_scapulatus_ are relatively smaller than teeth of either _mahaganus_ or
_woodfordi_. Also, in _scapulatus_ the upper canines are widely
separated due to lateral expansion of the palate at that point, whereas
in _mahaganus_ and _woodfordi_ the width across the upper canines is
relatively much less.


=Dobsonia= Palmer

   1898. _Dobsonia_ Palmer, Proc. Biol. Soc. Washington, 12:114,
         April 30.

   1810. _Cephalotes_ (part) É. Geoffroy, Ann. du Mus. d'Hist. Nat.,
         15:104.

_Dobsonia_, a genus of large to medium-sized fruit bats, occurring from
Celebes to the Solomon Islands, contains at least nine species. One
species and its two subspecies are endemic to the Solomons.

_Dobsonia_ differs from all other genera of megachiropteran bats in the
Solomons by combining absence of a small claw on the second digit and
presence of external tail vertebrae.

The cranium of _Dobsonia_ resembles, in some ways, the cranium of
_Rousettus_ as well as that of _Pteropus_. Even so, in _Dobsonia_ the
rostrum is shorter and the cheek-teeth, especially in the upper jaw, are
more crowded. The anterior part of the mandible is narrow and the lower
incisors are diminutive and often concealed by the flesh of the gum.


=Dobsonia inermis=

In a review of the genus _Dobsonia_, Andersen (1909_c_:532) named and
described _D. inermis_ and _D. nesea_ from the Solomons. Specimens of
_Dobsonia inermis_ from San Cristobal and Ugi were said to differ from
specimens of _D. nesea_ from Alu, Shortland, and Rubiana in having
perpendicular as opposed to anteriorly slanted upper canines. Andersen
(1909_c_:532) reported that the two species were of "... the same
general size." Troughton (1936:348-349) studied specimens of _Dobsonia_
from Bougainville and Santa Ysabel and, because of individual variation
in proclivity of the upper canines, concluded that _D. nesea_ was
conspecific with _D. inermis_. He (p. 349) noted that the ears were
shorter in _inermis_ than in _nesea_, but the size of teeth showed
insular variation and a "... confusing amount of intergradation ...
[that obscures] ... diagnostic importance."

Specimens of _Dobsonia_ from Choiseul are smaller (externally and
cranially) than those from Alu, Shortland, Rubiana, Bougainville, Fauro,
Vella Lavella, Guadalcanal, Florida, Ugi, San Cristobal, and Rennell.
Specimens from Santa Ysabel (see Fig. 8) are intermediate in size
between those from Choiseul and the other islands listed. Judging from
available specimens, two subspecies of _Dobsonia inermis_ occur in the
Solomons. Specimens from Choiseul (see A, Fig. 8), which are smaller
than those from other islands, represent one subspecies (heretofore
unrecognized), whereas specimens from other islands (except Santa
Ysabel) represent a second subspecies. Specimens from Santa Ysabel are
slightly larger than those on Choiseul and are regarded as intergrades
between the two subspecies. Specimens from Rennell, Ugi, San Cristobal,
Florida, Fauro, and Guadalcanal are slightly smaller than those from
Bougainville, Vella Lavella, Shortland, and Rubiana, but the differences
are not great enough to warrant recognition of two subspecies.
Therefore, the subspecific name _nesea_ is arranged as a synonym of
_inermis_, which has priority, and the latter name is used for specimens
of _Dobsonia inermis_ from the Solomon islands other than Choiseul and
Santa Ysabel. Additional remarks on the distribution of this species are
in the section on Zoogeography and Speciation.

Pohle (1953:130) suggested that _Dobsonia inermis_ (as well as _D.
crenulata_ and _D. praedatrix_) is conspecific with _D. viridis_, but
Laurie and Hill (1954:41) did not adopt his suggestion. I have not seen
adequate series of _crenulata_, _praedatrix_, and _viridis_ (none of
which occurs in the Solomons) to judge systematic relationships of these
kinds; therefore I follow Laurie and Hill.


=Dobsonia inermis inermis= Andersen

   1909. _Dobsonia inermis_ Andersen, Ann. Mag. Nat. Hist., ser. 8,
         4:532, December, type from San Cristobal; 1912, Andersen,
         Catalogue of the Chiroptera ... British Museum, 1:475.

   1936. _Dobsonia inermis inermis_, Troughton, Rec. Australian Mus.,
         14:349, April 7, from Santa Ysabel; 1954, Laurie and Hill, List
         of land mammals of New Guinea, Celebes and adjacent islands,
         p. 41, June 30; 1956, Hill, The natural history of Rennell
         Island, British Solomon Islands, 1:74, November 28, from
         Rennell Island.

   1878. _Cephalotes peroni_ (part), Dobson, Catalogue of the
         Chiroptera ... British Museum, p. 91; 1879, Trouessart, Rev.
         Mag. Zool., 3:208; 1887, Thomas, Proc. Zool. Soc. London,
         p. 323, March 15, from Ugi and San Cristobal; 1888, Thomas,
         Proc. Zool. Soc. London, p. 476, December 4; 1897, Trouessart,
         Catalogus Mammalium ..., 1:87.

   1899. _Dobsonia peroni_ (part), Trouessart, Catalogus Mammalium
         ..., 2:1278.

   1909. _Dobsonia nesea_ Andersen, Ann. Mag. Nat. Hist., ser. 8,
         4:532, December 1, type from Shortland Island; 1912, Andersen,
         Catalogue of the Chiroptera ... British Museum, 1:476, from
         Shortland and Rubiana; 1931, Sanborn, Publ. Field Mus. Nat.
         Hist., Zool. Ser., 18:22, February 12, from San Cristobal.

   1936. _Dobsonia inermis nesea_, Troughton, Rec. Australian Mus.,
         14:348, April 7, from Bougainville; 1953, Pohle, Z. Säugetierk.,
         17:130, October 27; 1954, Laurie and Hill, List of land mammals
         of New Guinea, Celebes and adjacent islands, p. 41, June 30,
         from New Georgia.

_Specimens examined_ (13 males and three females; three in alcohol,
crania extracted and cleaned).--Fauro in April, 23728, 23740, 23751;
Vella Lavella in November, 23134-36, 23141, 23145, 23147, 23149, 23151,
23153; Guadalcanal in May and June, 23865, 23914, 24008; Florida in
October, 24416.

_Measurements._--See tables 2 and 3.

_Remarks._--Heretofore, _Dobsonia inermis inermis_ was unreported from
Fauro, Vella Lavella, Guadalcanal, and Florida. Apparently the
subspecies occurs on most islands of the archipelago (see Fig. 9).

In coloration and most cranial dimensions the specimens listed above
agree with specimens of _D. i. inermis_ from Alu, Shortland, and Rubiana
(Andersen, 1909_c_:532; 1912:475, 476), Bougainville (Troughton,
1936:348, 349), and Rennell (Hill, 1963:74). The forearm in the adult
male holotype of "_nesea_," from Shortland, is 109.5 as opposed to 109.0
in an adult female topotype of _inermis_ from Ugi (Andersen, 1912:478)
in the southeastern part of the archipelago (see Fig. 9). Forearms of
specimens of _D. i. inermis_ from Vella Lavella are 107 to 112
(measurements from labels because forearms of these specimens were
broken and therefore could not be remeasured). Forearms of specimens
from Fauro, Florida, Guadalcanal, and Rennell are 103.6 to 110.0 (see
Hill, 1956:74). Variation in length of forearm probably is not
significant because no cline is evident (see Fig. 9 and Table 2).

In 1964, 11 specimens of _Dobsonia inermis_ were collected on Choiseul.
They are smaller, externally and cranially, than specimens of _D.
inermis_ from Sun Cristobal, Ugi, Rennell, Guadalcanal, Florida,
Rubiana, Vella Lavella, Shortland, Alu, Bougainville, and Fauro, and may
be named and described as follows:


=Dobsonia inermis minimus=, new subspecies

_Type._--Adult male skin and skull, in good condition (originally stored
in 70 per cent alcohol for about one year), no. BBM-BSIP 23716, Bernice
P. Bishop Museum; from Choiseul Island, British Solomon Islands
Protectorate; obtained on 20 March 1964 by Philip Temple, original
number 1524.

_Distribution._--Choiseul Island (type locality); intergrades from Santa
Ysabel also assigned to this subspecies.

_Diagnosis._--Size small for species; wing membranes, feet, and ears
black; dorsal surface of interfemoral membrane sparsely set with silvery
hairs, other membranes naked; hair soft, medium length (10 on mantle, 5
on crown), black hairs and scattered white hairs on face and crown; fur
of dorsal surface of mantle composed of whitish hairs having faint olive
cast imparting general color of Buffy-Citrine; hair of venter short
(about 5), soft, and fine; general coloration Buffy-Citrine; cranium
delicate; rostrum narrow in dorsal aspect (nasals not expanded
laterally); forehead (junction of nasals and frontals) pronounced in
lateral aspect; teeth resembling those of other subspecies of _D.
inermis_ but slightly smaller.

_Comparisons._--From adults of _Dobsonia inermis inermis_, which occurs
on Rennell, San Cristobal, Ugi, Malaita, Florida, Guadalcanal, Rubiana,
Vella Lavella, Shortland, Alu, Bougainville, and Fauro, _minimus_
differs in being smaller. Average length of mandible 31.2 and 33.4. For
other measurements see Table 2.

From _Dobsonia praedatrix_, which occurs on New Britain, New Ireland,
and Duke of York (northward of the Solomons), _minimus_ differs in being
smaller in all dimensions; length of forearm averaging 100.5 as opposed
to 116.0, and greatest length of skull 42.4 as opposed to 50.0.

    [Illustration: FIG. 8. Greatest length of skull plotted against
       zygomatic breadth for two subspecies of _Dobsonia inermis_. Symbols
       represent _D. i. inermis_ ([BW]), _D. i. minimus_ ([TW]), and
       intergrades assigned to _minimus_ ([BC]). Capital letters are used
       to relate groups of specimens to the island or islands from which
       they were collected; spatial distribution of specimens indicated in
       the scatter diagram thus is shown in the inset map. Specimens from
       Santa Ysabel and Bougainville are deposited in the Australian
       Museum. The type specimen of _D. i. inermis_ is labeled "E." For
       names of islands see Fig. 2.]

    [Illustration: FIG. 9. Distribution of _Dobsonia inermis inermis_
       ([BC]) and _D. inermis minimus_ ([RW]). For names of islands see
       Fig. 2.]

    TABLE 2. Average and Extreme Measurements of Two Subspecies of
       _Dobsonia inermis_.

  ================+====================================+===================
                  |           _D. i. minimus_          | _D. i. inermis_
                  +------------------+-----------------+-------------------
                  |                  |                 |  Guadalcanal,
    MEASUREMENT   |     Choiseul     |  Santa Ysabel   |  Fauro, Vella
                  |     4 [M], 2 [F] |  1 [M], 2 [F]   |Lavella, Florida
                  |                  |                 |    9 [M], 2 [F]
  ----------------+------------------+-----------------+-------------------
  Length of head  |                  |                 |
    and body      |174.5 (170 -180  )|                 |174.5 (160  -190  )
  Tail vertebrae  | 28.5 (24  - 33  )| 23.0 (21.5-24.0)| 30.4 ( 25  - 35  )
  Hind foot       | 25.3 (25.1- 25.9)| 23.5 (23.5-24.0)| 29.3 ( 26.0- 31.6)
  Ear             | 21.5 (21.0- 22.9)| 21.1 (21.0-21.5)| 23.1 ( 19  - 25  )
  Length of       |                  |                 |
    forearm       |100.5 (98.1-104.0)|105.3 (104 -107 )|108.4 (105  -112  )
  2nd metacarpal  | 43.6 (42.6- 45.2)|                 | 48.1 ( 45.9- 50.9)
  3rd metacarpal  | 61.5 (59.8- 62.9)|                 | 67.6 ( 65.2- 68.5)
  4th metacarpal  | 57.5 (56.5- 58.5)|                 | 62.5 ( 58.7- 65.5)
  5th metacarpal  | 59.0 (57.0- 60.5)|                 | 64.4 ( 61.8- 66.0)
                  |                  |                 |
  Greatest length |                  |                 |
    of skull      | 42.4 (42.1- 43.5)| 44.0 (43.0-45.6)| 45.9 ( 45.2- 47.4)
  Condylobasal    |                  |                 |
    length        | 40.4 (39.5- 41.3)| 41.7 (41.1-42.6)| 43.6 ( 43.1- 45.0)
  Zygomatic       |                  |                 |
    breadth       | 25.6 (24.9- 26.8)| 26.5 (25.7-27.6)| 27.9 ( 27.2- 28.5)
  Breadth of      |                  |                 |
    braincase     | 16.8 (16.5- 17.4)| 18.0 (17.1-19.9)| 17.9 ( 16.7- 19.0)
  Breadth across  |                  |                 |
    upper canines |  8.4 ( 8.1-  8.7)|                 |  9.2 (  9.2-  9.5)
  Breadth across  |                  |                 |
    first upper   |                  |                 |
    molars        | 12.1 (11.8- 12.6)|                 | 13.1 ( 12.6- 13.3)
  Length of       |                  |                 |
    maxillary     |                  |                 |
    tooth-row     | 15.6 (15.5- 15.8)| 16.4 (16.0-17.2)| 16.4 ( 15.9- 17.0)
  Length of       |                  |                 |
    mandibular    |                  |                 |
    tooth-row     | 17.1 (16.8- 17.6)| 17.8 (17.3-18.4)| 18.2 ( 17.8- 19.4)
  ----------------+------------------+-----------------+-------------------

_Measurements._--Comparative measurements of the subspecies _inermis_
and _minimus_ are given in Table 2. Some measurements of the type are as
follows: Length of head and body, 147; tail vertebrae, 31; hind foot,
25; ear, 21; length of forearm, 99.5; 2nd metacarpal, 42.8; 3rd
metacarpal, 62.7; 4th metacarpal, 58.5; 5th metacarpal, 59.1; greatest
length of skull, 42.2; condylobasal length, 40.6; zygomatic breadth,
25.8; breadth of braincase, 16.8; length of maxillary tooth-row, 15.8;
length of mandible, 31.2.

_Remarks._--_Dobsonia inermis minimus_ is the smallest subspecies of
_Dobsonia inermis_. Specimens from Santa Ysabel, southeastward of
Choiseul, are slightly larger than the type and paratypes of _minimus_.
As can be seen in the scatter diagram (Fig. 8), a male from Santa Ysabel
is as large as one male and most females of _D. i. inermis_. The other
three specimens from Santa Ysabel also are slightly larger than
specimens of _minimus_ from Choiseul, but are much smaller than
specimens of _D. i. inermis_, and, therefore, are referred to _D. i.
minimus_.

Although there is a cline in size of _Dobsonia inermis_ from Choiseul to
Florida (generally southward; Fig. 9), no cline in size is apparent
between Choiseul and Fauro (generally westward). Specimens of _D.
inermis_ from Fauro are average for the subspecies _inermis_; there is
no evidence, in the small series available, of intergradation between
_minimus_ on Choiseul and _inermis_ on Fauro.

_Specimens examined_ (eight males and three females, all originally in
alcohol; seven crania, all adults, extracted and cleaned).--Choiseul in
March, 23565, 23628, 23637, 23665-67, 23640, 23714, 23716 (holotype),
23717, 23720. Ellis LeG. Troughton kindly examined and measured nos.
AM-M. 3693[M], AM-M. 3694[M], AM-M. 3937[F], and AM-M. 3940[F], from
Santa Ysabel in the Australian Museum.


Subfamily Macroglossinae


=Macroglossus= F. Cuvier

   1824. _Macroglossus_ F. Cuvier, Des dents des mammiferes ...
         zoologiques, p. 248.

   1840. _Kiodotus_ Blyth, _in_ Cuvier's animal kingdom ..., p. 69.

   1891. _Carponycteris_ Lydekker, _in_ Flower and Lydekker, mammals
         living and extinct, p. 654.

   1902. _Odontonycteris_ Jentink, Notes Leyden Mus., 23:140, July 15.

_Macroglossus_, the widest-ranging genus of macroglossine bats, occurs
from southeastern Asia to the southern islands of the Solomon
Archipelago (see Ellerman and Morrison-Scott, 1966:101; Laurie and Hill,
1954:44). One species, known also from Celebes and New Guinea, occurs in
the Solomons and is represented there by an endemic subspecies.

Numerous generic names have been applied, at one time or another, to
bats now considered as _Macroglossus_. Trouessart (1904:65) and Miller
(1907:70) listed the one bat of this genus occurring in the Solomons
under _Carponycteris_ and _Kiodotus_, respectively. Andersen (1911:642;
1912:767) and, later, Sanborn (1931:22) identified this bat as
_Macroglossus lagochilus microtus_. Troughton (1936:350), reporting an
extension of range of this species in the Solomons, used the generic
name _Odontonycteris_ without explanation. Andersen (1912:754) pointed
out that Jentink originally established the name _Odontonycteris_ on the
basis of an extra premolar in each upper jaw as opposed to the usual two
in _Macroglossus_, and arranged _Odontonycteris_ as a synonym of
_Macroglossus_ because "in no genus of Megachiroptera are dental
anomalies of so frequent occurrence as in _Macroglossus_, and on no
point of the jaws are these anomalies ... so often met with as on that
occupied by the molar series." Sanborn (1931:22) and Phillips (1966:27)
noted variation in number of incisors in _Macroglossus_ as well as in
_Melonycteris_, another macroglossine genus. All of the more recent
workers (Ellerman and Morrison-Scott, 1966; Pohle, 1953; Laurie and
Hill, 1954) use the name _Macroglossus_.


=Macroglossus lagochilus=

_Macroglossus lagochilus_ has at least three subspecies, one of which is
endemic to the Solomons. The species ranges from Celebes on the west to
the Solomon Islands on the east, occurring not only in New Guinea but
also on many of the small adjacent islands (see Laurie and Hill,
1954:44).

    [Illustration: FIG. 10. Distribution of _Macroglossus lagochilus
       microtus_. For names of islands see Fig. 2.]


=Macroglossus lagochilus microtus= Andersen

   1911. _Macroglossus lagochilus microtus_ Andersen, Ann. Mag. Nat.
         Hist., Ser. 8, 7:642, June, type from Guadalcanal, additional
         specimens from Florida; 1912, Andersen, Catalogue of the
         Chiroptera ... British Museum, 1:767; 1931, Sanborn, Publ.
         Field Mus. Nat. Hist., Zool. Ser., 18:22, February 12, from San
         Cristobal; 1953, Pohle, Z. Säugetierk., 17:130, October 27,
         from Bougainville; 1954, Laurie and Hill, List of land mammals
         of New Guinea, Celebes and adjacent islands, p. 44, June 30.

   1888. _Macroglossus australis_ (part). Thomas, Proc. Zool. Soc.
         London, p. 476, December 4, from Guadalcanal.

   1904. _Carponycteris nana_ (part), Trouessart, Catalogus Mammalium
         ..., Suppl., p. 65.

   1907. _Kiodotus_ sp., Miller, Bull. U. S. Nat. Mus., 57:70,
         June 29.

   1936. _Odontonycteris lagochilus microtus_, Troughton, Rec.
         Australian Mus., 14:350, April 7, from Bougainville.

_Specimens examined_ (14 males and 16 females; in alcohol).--Choiseul in
March, 23654-57, 23614, 23629, 23643, 23645, 23647, 23677-79, 23684;
Vella Lavella in December, 23277-79, 23283-84; Fauro in April, 23765;
Guadalcanal in May and June, 23830, 23864, 23935; Kolombangara in
January, 23385, 23399, 23397, 23407, 23420-21; Santa Ysabel in June,
24067; Malaita in June, 24067.

_Measurements._--Average and extreme external measurements of 14 males
and 15 females are as follows: Length of head and body, 68.3 (63-72);
tail vertebrae present but scarcely perceptible and therefore not
measured; hind foot, 11.4 (9.0-12.9); ear, 12.0 (10.0-12.9); length of
forearm, 37.6 (36.2-39.9).

_Remarks._--The distribution of _Macroglossus lagochilus microtus_ has
not been well known. Specimens herein reported from Choiseul, Fauro, and
Vella Lavella provide new records of distribution. As shown on Figure
10, the subspecies occurs throughout the Solomon Islands.

_Macroglossus lagochilus microtus_ differs slightly from _M. l. nanus_
Matschie, the subspecies of the Bismarck Archipelago and Admiralty
Islands to the north of the Solomons. _M. l. nanus_ averages slightly
larger than _microtus_ (see Andersen, 1912:768-769, for comparative
measurements) but otherwise closely resembles it.

Individual variation is evident in several measurements of the specimens
at hand (in length of forearm, for example) but no clines are apparent.
Four females obtained in March were lactating, as was one taken in
December and one taken in January.


=Melonycteris= Dobson

   1877. _Melonycteris_ Dobson, Proc. Zool. Soc. London, p. 119,
         June 1.

   1877. _Cheiropteruges_ Ramsay, Proc. Linn. Soc. New South Wales,
         2:19, July.

   1887. _Nesonycteris_ Thomas, Ann. Mag. Nat. Hist., ser. 5, 14:147,
         February.

The genus _Melonycteris_ is known from three species, two apparently
endemic to the Solomon Islands and the third occurring in eastern New
Guinea and the Bismarck Archipelago (Laurie and Hill, 1954:45).

Heretofore, the generic name _Nesonycteris_ has been applied to the
species in the Solomons, whereas _Melonycteris_ has been restricted to
the one species in the Bismarck Archipelago and New Guinea. Andersen
(1912:792) judged that _Nesonycteris_ was clearly distinct from
_Melonycteris_ on the basis of two characters (loss of a claw on the
second digit and loss of the inner, lower incisors). On the other hand,
he noted striking similarities in general cranial features, dentition,
palatal ridges, tongue, and external appearance of the two genera. Pohle
(1953:131) synonymized the two but Laurie and Hill (1954:45) considered
them distinct. I have suggested previously (Phillips, 1966:26, 27) that
characteristics used to distinguish between _Melonycteris_ and
_Nesonycteris_ are of less than generic value. Variability of number of
incisors in the upper jaw of specimens of _Melonycteris_ (and in other
macroglossine genera, as well) indicates a lack of selective pressure
for either increase or decrease in number of incisors. Furthermore, the
loss of the small claw on the second digit might not be important
because, as Bader and Hall (1960:15) have pointed out, limbs of bats
vary more in phenotypic expression than do other parts of the skeletal
structure.

The discovery of a new species (_Melonycteris aurantius_) in the Solomon
Islands sheds additional light on the problem. Although _M. aurantius_
possesses the distinguishing characteristics of the genus
"Nesonycteris," the species closely resembles _Melonycteris_ in other
features. Similarity in structure of hair of _Melonycteris_ and
_Nesonycteris_, as first reported by Benedict (1957:293), also supports
the argument for synonymy (see Phillips, 1966:26).

_Melonycteris aurantius_ lacks a small claw on the second digit and has
only two lower incisors. In these ways this species is like _woodfordi_,
which also is restricted to the Solomons. On the other hand, the
structure of the skull of _M. aurantius_ is like that of _M. melanops_,
which is the species found in the Bismarck Archipelago.

Although _melanops_ is not yet known from the Solomon Islands, I have
included it in the following key.

Key to Known Species of _Melonycteris_

  1.    Ventral surface darker than dorsum, but not strongly
        contrasting with it; lacking a small claw on the
        second digit,                                                2

  1´.    Ventral surface nearly black, strongly contrasting with
         dorsum; small claw on second digit,
                                               =Melonycteris melanops=

  2(1´). Pelage bright, Cinnamon-Rufous; postorbital region of
         skull expanded (about 8.3 wide),
                                      =Melonycteris aurantius=, p. 816

  2´.    Pelage dark, near Wood-Brown or Cinnamon; postorbital
         region of skull constricted (about 7.5),
                                      =Melonycteris woodfordi=, p. 816

    [Illustration: FIG. 11. Distribution of _Melonycteris aurantius_
       [BC] and _M. woodfordi_ [LW]. For names of islands see Fig. 2.]


=Melonycteris aurantius= Phillips

   1966. _Melonycteris aurantius_ Phillips, Jour. Mamm., 47:23-27,
         March 12, type from Florida Island, additional specimens from
         Choiseul Island.

_Specimens examined_ (six females; three in alcohol).--Florida in
October, 24440; Choiseul in March, 23615, 23617, 23558, 23694, 23681.

_Measurements._--Average and extreme measurements of six females are as
follows: Length of head and body, 80.8 (77-106); hind foot, 17.2
(16.0-18.7); ear, 12.7 (11.5-14.0); length of forearm, 49.3 (42.9-53.8).
Average and extreme measurements of skulls of five females are as
follows: Greatest length of skull, 31.8 (30.8-33.3); condylobasal
length, 29.7 (28.6-32.4); zygomatic breadth, 18 (17.2-20.0); breadth of
braincase, 12.6 (12.4-13.2); postorbital breadth, 8.3 (8.0-8.9); length
of maxillary tooth-row, 10.1 (9.4-10.4); length of mandibular tooth-row,
11.7 (10.8-12.2).

_Remarks._--On Choiseul Island _Melonycteris aurantius_ was taken at the
same locality as its congener, _Melonycteris woodfordi_.

Externally, _M. aurantius_ resembles _M. woodfordi_. These species are
the same size, but the former is brighter in color (nearly orange in
adults) than the latter, which is Wood-Brown dorsally. Internally,
differences between _M. aurantius_ and _M. woodfordi_ are more obvious.
In the skull of _M. aurantius_, the postorbital region is expanded
(measuring about 8.3), whereas in _M. woodfordi_ the postorbital region
is constricted. Furthermore, in lateral aspect the posterior portion of
the skull of _M. aurantius_ is down-turned and the angle of the facial
axis with the basicranial axis is much more acute than in _M.
woodfordi_.

The number of upper incisors is highly variable in the six specimens of
_M. aurantius_ that I have examined. In two specimens an extra tooth has
erupted just anterior to I2 and there is a total of six upper incisors.
In two other specimens an extra tooth has erupted in front of I2 on one
side but not the other. I could find no trace of an extra tooth in the
remaining two specimens.

Practically nothing is known about the natural history of _M.
aurantius_, or, indeed, that of either of the other two species of this
genus. One field collector (Temple, _in litt._) for the Bishop Museum
reported that he obtained both _M. aurantius_ and _M. woodfordi_ in the
same mist net in one night. The holotype, an adult female, was lactating
when obtained in October.


=Melonycteris woodfordi= (Thomas)

   1887. _Nesonycteris woodfordi_ Thomas, Ann. Mag. Nat. Hist., ser.
         5, 14:147, February, type from Shortland Island; 1887, Thomas,
         Proc. Zool. Soc. London, p. 324, March 15; 1888, Thomas, Proc.
         Zool. Soc. London, p. 476, December 4; 1898, Trouessart,
         Catalogus Mammalium ..., 1:90; 1899, Matschie, Die Megachiroptera
         ... naturkunde, p. 91; 1904, Trouessart, Catalogus Mammalium ..,
         Suppl., p. 66; 1907, Miller, Bull. U. S. Nat. Mus., 57:74,
         June 29; 1912, Andersen, Catalogue of the Chiroptera ... British
         Museum, 1:792, from Alu, Shortland, Fauro, and Guadalcanal; 1931,
         Sanborn, Publ. Field Mus. Nat. Hist., Zool. Ser., 18:23, February
         12, from Russell Island (Pavuvo); 1954, Laurie and Hill, List of
         land mammals of New Guinea, Celebes and adjacent islands, p. 45,
         June 30.

   1953. _Melonycteris woodfordi_, Pohle, Z. Säugetierk., 17:130,
         October 27, from Bougainville Island; 1966, Phillips, Jour. Mamm.,
         47:23, March 12, from Choiseul.

_Specimens examined_ (three males and one female; in
alcohol).--Choiseul, in April, 23413-14, 23434, 23275.

_Measurements._--Average and extreme measurements of three males and one
female are as follows: Length of head and body, 86.1 (83.1-91.0); hind
foot, 19.6 (17.2-22.2); ear, 11.3 (10.8-11.7); length of forearm, 54.4
(52.1-57.7).

_Remarks._--Specimens of _Melonycteris woodfordi_ from Choiseul
constitute a new locality of occurrence for the species. Apparently _M.
woodfordi_ occurs throughout the Solomons (see Fig. 11).

Thomas (1887_a_:147) named _Nesonycteris woodfordi_ in a preliminary
report that appeared before the publication of the more detailed
description of the genus and species (1887_b_:323-324). In the second
paper he stated that the anterior projections of the premaxillary bones
are separated distinctly in both _Nesonycteris_ and _Melonycteris_.
According to Thomas (1887_b_:323), it was by some "accident" that Dobson
(1878:4) reported the anterior projections of the premaxillary bones in
_Melonycteris melanops_ to be united. Writing at a later date, Andersen
(1912:785) reported that in _Melonycteris melanops_ the premaxillary
bones have "simple contact with each other." Furthermore, in Andersen's
(1912:791) illustration of _M. woodfordi_ the premaxillary bones are in
contact anteriorly. In specimens of _woodfordi_ and _melanops_ examined
by me, the premaxillary bones are in contact. In _M. aurantius_ the
premaxillary bones are not in contact, and it differs from _woodfordi_
in several other respects.

In _M. woodfordi_, as in other macroglossine bats, there is variability
in dentition. One specimen examined has a total of three upper incisors,
and another had an extra peglike tooth just anterior to I1.


Subfamily Nyctimeninae


=Nyctimene= Borkhausen

   1797. _Nyctimene_ Borkhausen, Deutsche fauna ..., 1:86.

   1810. _Cephalotes_ É. Geoffroy, Ann. du Mus. d'Hist. Nat., 15:104.

   1811. _Harpyia_ Illiger, Prodr. Syst. Mamm. et Avium, p. 118.

   1837. _Gelasinus_ Temminck, Monographe de Mammalia ..., 2:100.

Tube-nosed bats of the genus _Nyctimene_ occur from Celebes on the west
to the Santa Cruz Islands on the east. Heretofore, two species (_N.
albiventer_ and _N. major_), each with an endemic subspecies, were known
from the Solomon Islands. Both species occur also in New Guinea and on
many adjacent islands. A new species of _Nyctimene_, apparently endemic
to the Solomons, and a new subspecies of _N. albiventer_ are named
beyond.

_Nyctimene_ is related closely to _Cynopterus_ and the "Cynopterus
group" of Andersen (1912:691). Because _Nyctimene_ is a highly
specialized bat, Miller (1907:75) placed it in a subfamily separate from
that of _Cynopterus_ and its allies.

Andersen (1912:696, 697) placed the species of _Nyctimene_ previously
known from the Solomons in two groups, the "papuanus" group and the
"cephalotes" group, on the basis of difference in length of forearm and
length of maxillary tooth-row. Because of its short forearm (about 58),
_N. albiventer_ is in the _papuanus_ group; and _N. major_, because of
its long forearm (about 74), is in the _cephalotes_ group.

Key to Species of Nyctimene in the Solomons

  1.     Forearm longer than 70; males grayish-brown, females
         pale gray,                        =N. major scitulus=, p. 825

  1'.    Forearm shorter than 70; males dark brown, females
         pale brown,                                                 2

  2(1'). Forearm about 65,                    =N. malaitensis=, p. 822

  2'.    Forearm less than 61,                 =N. albiventer=, p. 818


=Nyctimene albiventer=

This species occurs throughout New Guinea and on many adjacent islands,
including the Bismarck Archipelago and the Admiralty and Solomon
islands. The species varies geographically and five subspecies are
recognized. The two subspecies in the Solomons resemble _N. albiventer
papuanus_, the subspecies that ranges from eastern New Guinea to New
Britain. _N. albiventer bougainville_ occurs in the western chain of
islands of the Solomons, whereas another subspecies, named as new
beyond, occurs in the eastern chain of islands (see Fig. 12).

Sexual dichromatism is striking. As Andersen (1912:690) previously
reported, females generally are paler, more brownish than males, which
are dark and have a better defined black dorsal stripe.


=Nyctimene albiventer bougainville= Troughton

   1936. _Nyctimene bougainville_ Troughton, Rec. Australian Mus.,
         19:349, April 7, type from Bougainville.

   1954. _Nyctimene albiventer bougainville_, Laurie and Hill, List of
         land mammals of New Guinea, Celebes and adjacent islands, p. 46,
         June 30.

   1953. _Nyctimene papuanus bougainville_, Pohle, Z. Säugetierk.,
         17:130, October 27.

_Specimens examined_ (nine males, one female; nine in alcohol, nine
crania extracted and cleaned).--Bougainville in December, AM-M. 5786
(paratype); Guadalcanal in May, 23812, 23815, 23827; Kolombangara in
January and February, 23369, 23381, 23388, 23406, 23444, 23456.

_Measurements._--See Table 3.

_Remarks._--Heretofore, _Nyctimene albiventer bougainville_ was not
known from Kolombangara and Guadalcanal. The subspecies apparently
ranges throughout the western chain of the Solomons.

Troughton (1936:350) considered _Nyctimene bougainville_ specifically
distinct from its nearest ally, _N. papuanus_. Pohle (1953:130) did not
examine specimens of either kind, but on the basis of Troughton's
description decided that _N. bougainville_ differed only subspecifically
from _N. papuanus_. Laurie and Hill (1954:46) synonymized _bougainville_
and _papuanus_ with _N. albiventer_. However, Troughton (1936:350)
pointed out that in addition to size _bougainville_ differed from
_papuanus_ by having narrower and longer pm3 and pm4. Judging from
specimens examined by me, such is the case, and the difference is even
more pronounced in m1.

Specimens of _N. a. bougainville_ from Kolombangara and Guadalcanal
agree with a paratype of this subspecies from Bougainville. Geographic
variation, if present in the population in the western chain of islands
(see Fig. 12), is slight and not notable in the series available. Some
individual variation was found, especially in the shape of the
interorbital region of the skull. An adult male from Kolombangara is
unusually dark, almost black; color of the other specimens (all in
alcohol) is consistent according to sex.

_Nyctimene albiventer_ from Choiseul and Santa Ysabel is smaller, in all
respects, than _N. albiventer_ from Bougainville, Kolombangara, and
Guadalcanal (see Table 3), and therefore may be named and described as
follows:


=Nyctimene albiventer minor=, new subspecies

_Type._--Adult male, skin and skull, in good condition (originally
stored in alcohol for about one year), no. BSIP 23636, Bernice P. Bishop
Museum; from Choiseul Island, British Solomon Islands Protectorate;
obtained on 11 March 1964, by Philip Temple, original number 1441.

_Distribution._--Known only from Choiseul and Santa Ysabel islands (see
Fig. 12).

_Diagnosis._--Small for _Nyctimene_; wing membranes brown with scattered
yellow spots (dried specimens); uropatagium, feet, and ears brown;
dorsum of tibia set with hair, ventral surface naked; dorsum of
uropatagium sparsely set with pale brown hairs, ventral surface almost
bare; fringe of hairs along two centimeters of dorsal and ventral
surfaces of trailing edge of wing membrane; proximal third of dorsal
surface of forearm sparsely set with hairs; pelage of back soft and
thick, of medium length (about 7); hair on crown and nape short (about
4); well-defined black dorsal stripe, extending from uropatagium to
shoulders; skull resembling that of other subspecies of _N. albiventer_
but relatively smaller; zygomatic arch delicate, slender anteriorly; P2
small (see Fig. 14). Sexually dichromatic as follows: male--dorsum
Hair-Brown, bases of hairs darker; hair on throat sparse, medium length
(about 6), Hair-Brown; fur along sides of abdomen Drab; female--dorsum
having Buffy-Brown cast, some individual hairs Hair-Brown; shoulders
Sayal-Brown; hair on throat sparse, Hair-Brown on throat and midline of
abdomen; sides of abdomen Sayal-Brown.

_Comparisons._--From _Nyctimene major scitulus_, the largest member of
this genus in the Solomons, _N. a. minor_ differs in being smaller in
all measurements taken; forearm averaging 54.8 as opposed to 73.5;
greatest length of skull 28.2 as opposed to 37.0, and females pale brown
instead of pale gray.

From nine adults of _Nyctimene albiventer bougainville_ from
Bougainville, Kolombangara, and Guadalcanal, _minor_ differs as follows:
averaging slightly smaller in all dimensions; forearm averaging 54.8 as
opposed to 57.9; second metacarpal averaging 27.4 as opposed to 28.3;
5th metacarpal averaging 38.5 as opposed to 40.0; condylobasal length
26.7 as opposed to 28.0; length of mandibular tooth-row 10.3 as opposed
to 10.9; mandible smaller (see Fig. 14); dorsal stripe fainter.

From _Nyctimene albiventer papuanus_, known from eastern New Guinea, New
Britain, and the Admiralty Islands, _minor_ differs as follows: slightly
smaller in most dimensions; forearm averaging 54.8 as opposed to 57.0;
length of maxillary tooth-row 8.9 as opposed to 9.8; length of
mandibular tooth-row 10.3 as opposed to 11.0; breadth across upper third
premolars notably less (7.5 as opposed to 8.4).

_N. a. minor_ differs from _N. albiventer albiventer_ Gray, which occurs
about 800 miles to the west of _minor_, in ways made apparent by the
description by Andersen (1912:700-701). _N. a. minor_ occurs about 1500
miles eastward of the place from which _N. a. draconilla_ Thomas, a
subspecies essentially unknown to me, was named (see Laurie and Hill,
1954:46).

From _Nyctimene sanctacrucis_, known from the Santa Cruz Islands,
_minor_ differs as follows: much smaller in all dimensions; forearm
averaging 54.8 as opposed to 75; greatest length of skull 28.2 as
opposed to 34.5; length of maxillary tooth-row 8.9 as opposed to 12.9.

    [Illustration: FIG. 12. Distribution of _Nyctimene albiventer
       bougainville_ ([BC]) and _N. albiventer minor_ ([LW]).
       For names of islands see Fig. 2.]

    [Illustration: FIG. 13. Scatter diagram comparing two subspecies
       of _Nyctimene albiventer_. One individual of specimens thought
       to be intergrades is as large as specimens of _Nyctimene a.
       bougainville_, whereas the other three intergrades are about the
       same size as specimens of _N. a. minor_. Symbols represent
       _N. a. bougainville_ ([BW]), _N. a. minor_ ([TW]), and intergrades
       assigned to _minor_ ([BC]). For names of islands see Fig. 2.]

    TABLE 3. Average and Extreme Measurements of _Nyctimene albiventer
       bougainville_ and _N. a. minor_.

  ===============+==================+==================+===================
                 |  _N. a. minor_   |  _Intergrades_   | _N. a.
                 |                  |                  |    bougainville_
                 |                  |                  |
    MEASUREMENT  |    Choiseul,     |      Fauro       |   Kolombangara,
                 |   Santa Ysabel   |                  |    Guadalcanal
                 |   4 [M], 1 [F]   |   1 [M], 3 [F]   |    8 [M], 1 [F]
  ---------------+------------------+------------------+-------------------
  Length of head |                  |                  |
    and body     |107.0 (105 -109  )|109.2 (105 -112  )|110.0 (106  -117)
  Tail vertebrae | 20.0 (19.3- 20.5)| 21.0 ( 20 - 22  )| 19.2 ( 15.5- 23.0)
  Hind foot      | 14.2 (13.5- 15.0)|                  | 14.3 ( 13.0- 15.9)
  Ear            | 11.9 (11.0- 13.0)|                  | 12.8 ( 11.8- 14.5)
  Length of      |                  |                  |
    forearm      | 54.8 (54.0- 55.8)| 57.1 (55.9- 59.0)| 57.9 ( 55.8- 59.8)
  Greatest       |                  |                  |
    length of    |                  |                  |
    skull        | 28.2 (27.2- 28.9)| 28.6 (28.3- 29.7)| 29.7 ( 28.6- 30.1)
  Condylobasal   |                  |                  |
    length       | 26.7 (26.2- 27.5)| 27.4 (26.6- 28.0)| 28.0 ( 27.8- 28.9)
  Palatal length | 11.2 (10.9- 11.9)| 11.6 (11.3- 11.8)| 11.7 ( 11.0- 12.5)
  Breadth of     |                  |                  |
    braincase    | 12.0 (11.5- 12.4)| 12.0 (11.7- 12.2)| 12.3 ( 12.1- 12.8)
  Zygomatic      |                  |                  |
    breadth      | 18.9 (18.4- 19.7)| 18.6 (18.4- 19.2)| 19.2 ( 18.7- 20.0)
  Interorbital   |                  |                  |
    breadth      |  5.0 ( 4.7-  5.6)|  5.3 ( 5.0-  5.6)|  5.1 (  4.7-  5.5)
  Breadth across |                  |                  |
    first upper  |                  |                  |
    molars       |  8.6 ( 8.4-  8.9)|  8.9 ( 8.7-  9.1)|  9.1 (  8.8-  9.6)
  Maxillary      |                  |                  |
    tooth-row    |  8.9 ( 8.7-  9.3)|  9.3 ( 9.1-  9.5)|  9.5 (  9.2-  9.8)
  Mandibular     |                  |                  |
    tooth-row    | 10.3 (10.0- 10.6)| 10.5 (10.2- 11.1)| 10.9 ( 10.7- 11.4)
  ---------------+------------------+------------------+-------------------

_Measurements._--Measurements of the two subspecies from the Solomons
are given in Table 3. Some measurements of the type are as follows:
Length of head and body, 108; tail vertebrae, 20.5; hind foot, 14.7;
ear, 11.3; length of forearm, 55.1; 2nd metacarpal, 27.4; 3rd
metacarpal, 39.0; 4th metacarpal, 37.5; 5th metacarpal, 39.1; greatest
length of skull, 28.6; condylobasal length, 27.5; zygomatic breadth,
18.4; length of maxillary tooth-row, 9.0; length of mandibular
tooth-row, 10.4.

_Remarks._--_Nyctimene albiventer minor_ closely resembles _N.
albiventer bougainville_, differing from the latter mostly in size.
Although adults of _minor_ average only slightly smaller than adults of
_bougainville_ (see Table 3), there is only slight overlap (about 0.2 at
most) in most minimum dimensions of external and cranial features of
_bougainville_ and corresponding maximum dimensions of externals and
crania of _minor_. The difference in size is clearly shown in Figs. 13
and 14.

Four specimens of _Nyctimene albiventer_ from Fauro herein are
considered to be intergrades between _N. a. bougainville_ and _N. a.
minor_. As shown in Table 3, the specimens from Fauro average slightly
larger than those of _minor_ from Choiseul and Santa Ysabel and slightly
smaller than specimens of _bougainville_ from Kolombangara and
Guadalcanal. I have assigned the specimens from Fauro to _N. a. minor_
because they generally are closer to _minor_ in size (see Fig. 13).

_Specimens examined_ (five males and four females; seven in alcohol;
seven crania extracted and cleaned).--Choiseul in February and March,
23636 (holotype), 23631, 23540, 23646; Santa Ysabel in February, 23539;
Fauro in April, 23742, 23743, 23763, 23764.

One specimen of _Nyctimene_ from Malaita Island is smaller than
_Nyctimene major_, which is known from Shortland, Alu, Florida, New
Georgia, Guadalcanal, Choiseul, and Malapa (see Fig. 15) and is larger
than either of the two subspecies of _Nyctimene albiventer_ known from
Bougainville, Fauro, Kolombangara, Guadalcanal, Choiseul, and Santa
Ysabel. This specimen represents a previously unknown species and may be
named and described as follows:


=Nyctimene malaitensis=, new species

_Type._--Adult female, skin and skull, in good condition (originally
stored in alcohol for about one year), no. BSIP 24103, Bernice P. Bishop
Museum; from Malaita Island, British Solomon Islands Protectorate;
obtained on 1 July 1964, by Peter Shanahan, original no. unknown.

_Distribution._--Known only from Malaita (see Fig. 16).

_Diagnosis._--Size average for genus but larger than closest relative,
_Nyctimene albiventer_; wing membranes brown with scattered yellow spots
(dried specimen); uropatagium, ears, and feet brown; dorsal surface of
tibia set with hair, ventral surface bare; dorsal surface of uropatagium
sparsely set with hair, ventral surface having few, scattered hairs;
dorsal surface of trailing edge of wing membrane sparsely set with
hairs, ventral surface bare; proximal third of upper- and under-surface
of forearm set with hair; pelage of back luxuriant and soft (about 10
long); hair on crown and nape shorter than on back (4 to 8);
well-defined black dorsal stripe from shoulders to rump (about 2 wide);
basal half of most hairs on dorsum Deep Mouse Gray, distal half Light
Buff, tips Ochraceous-Tawny; some hairs on back entirely Light Buff;
hairs of crown Light Ochraceous Buff tipped with Ochraceous-Tawny; hair
on throat and along sides of abdomen Light Ochraceous Buff; hairs of
ventral midline Smoke Gray; braincase narrow; zygomatic breadth
relatively narrow; well-developed lambdoidal crest in female; rostrum
short, wide; upper canines slanted posteriorly; upper incisors large;
foramen ovale large (see Fig. 14).

    [Illustration: FIG. 14. Dorsal and ventral views of skulls of
       (A) _Nyctimene albiventer minor_ [specimen 23631 [M]],
       (B) _N. a. bougainville_ [specimen 23381 [M]], and
       (C) _N. malaitensis_ [specimen 24103 [F]].]

_Comparisons._--From _Nyctimene major scitulus_, the largest kind of
_Nyctimene_ in the Solomons, _malaitensis_ differs as follows: smaller
in all dimensions (forearm 65 as opposed to 73.5); greatest length of
skull 32.4 as opposed to 37.0; length of maxillary tooth-row 10.5 as
opposed to 13.0; length of mandibular tooth-row 11.8 as opposed to 14.2.

From nine adults of _Nyctimene albiventer bougainville_ from
Bougainville, Kolombangara, and Guadalcanal, _malaitensis_ differs as
follows: larger in all dimensions: forearm 65 as opposed to 57.9;
greatest length of skull 32.4 as opposed to 29.7; zygomatic breadth 20.4
as opposed to 19.2; and length of maxillary tooth-row 10.5 as opposed to
9.5; length of mandibular tooth-row 11.8 as opposed to 11.1.

From five adults of _Nyctimene albiventer minor_, from Choiseul and
Santa Ysabel, _malaitensis_ differs in the same ways it differs from _N.
a. bougainville_, but the contrast is even greater when _malaitensis_
and _minor_ are compared.

From _Nyctimene sanctacrucis_, known only from the Santa Cruz Islands,
_malaitensis_ differs in being smaller in all dimensions: forearm 65 as
opposed to 75; greatest length of skull 32.4 as opposed to 34.5; and
length of maxillary tooth-row 10.5 as opposed to 12.9.

_Measurements of the holotype._--Length of head and body, 118; tail
vertebrae, 23.0; hind foot, 16.0; ear, 14.0; length of forearm, 65.0;
2nd metacarpal, 33.2; 3rd metacarpal, 46.4; 4th metacarpal, 44.3; 5th
metacarpal, 46.0; greatest length of skull, 32.4; condylobasal length,
30.6; palatal length, 13.0; breadth of braincase, 12.5; zygomatic
breadth, 20.4; interorbital breadth, 5.5; breadth across first upper
molars, 9.5; length of maxillary tooth-row, 10.5; length of mandibular
tooth-row, 11.8.

    [Illustration: FIG. 15. Distribution of _Nyctimene malaitensis_ [BC]
       and _N. major scitulus_ [RW]. For names of islands see Fig. 2.]

_Remarks._--In size, _Nyctimene malaitensis_ is intermediate between _N.
albiventer_ and _N. major_. Because the type of _malaitensis_ is brown
and not pale gray, as are females of _major_, _N. malaitensis_ most
likely is more closely related to _N. albiventer_, in which the females
are brown. The teeth of the holotype and only known specimen of
_malaitensis_ are too worn to be useful in determining the relationships
between these species.

When more specimens are available, _N. malaitensis_ may prove to be a
subspecies of _N. albiventer_. At present, _malaitensis_ is accorded
specific rank in order not to obscure the apparent relationships of _N.
albiventer bougainville_ and _N. a. minor_. Additionally, _N.
malaitensis_ is given specific rank because (1) it is larger (especially
in external dimensions) than the largest subspecies of _N. albiventer_
(compare above measurements with those in Table 3), and (2)
_malaitensis_ does not form a cline with either of the two subspecies of
_N. albiventer_.

_Specimen examined_ (one female).--Malaita in July, 24103 (holotype).


=Nyctimene major=

This large species of tube-nosed bat has at least four subspecies, one
of which (_N. major scitulus_) is endemic to the Solomons. The species
occurs throughout eastern New Guinea and on many of the islands adjacent
to the eastern coast of New Guinea, including the Trobriand Islands,
the Bismarck Archipelago, and the Solomons (see Laurie and Hill,
1954:47). The geographic distribution of the species generally is the
same as that of _N. albiventer_.

In _Nyctimene major_, as in _N. albiventer_, most males are
grayish-brown, whereas most females are pale gray.


=Nyctimene major scitulus= Andersen

   1910. _Nyctimene scitulus_ Andersen, Ann. Mag. Nat. Hist., ser. 8,
         6:623, December 1, type from Shortland; 1912, Andersen, Catalogue
         of the Chiroptera ... British Museum, 1:711, from Shortland, New
         Georgia, Florida, Guadalcanal; 1931, Troughton, Proc. Linnean Soc.
         New South Wales, 56:206, July 15; 1931, Sanborn, Publ. Field Mus.
         Nat. Hist., 18:22, February 12, from Choiseul and Malapa; 1942,
         Tate, Bull. Amer. Mus. Nat. Hist., 80:342, December 31.

   1954. _Nyctimene major scitulus_, Laurie and Hill, List of land
         mammals of New Guinea, Celebes and adjacent islands, p. 47,
         June 30.

   1862. _Harpyia pallasi_, Gerrard, Catalogue of the bones ...
         British Museum, p. 58.

   1870. _Harpyia cephalotes_, Gray, Catalogue of monkeys, lemurs and
         fruit-eating bats in the British Museum, p. 121.

   1878. _Harpyia major_, Dobson, Catalogue of the Chiroptera ...
         British Museum, p. 90; 1879, Trouessart, Rev. Mag. Zool., 3:207;
         1887, Thomas, Proc. Zool. Soc. London, p. 323; 1888, Thomas, Proc.
         Zool. Soc. London, p. 476; 1897, Trouessart, Catalogus Mammalium
         ..., 1:87.

   1899. _Cephalotes major_, Trouessart, Catalogus Mammalium ...,
         2:1277.

   1899. _Gelasinus major_, Matschie, Die Megachiroptera ...
         naturkunde, p. 84; 1904, Trouessart, Catalogus Mammalium ...,
         Suppl., p. 64.

_Specimens examined_ (four males and one female; dried skins with skulls
inside).--Florida in October, 24397, 24413, 24418, 24419.

_Measurements._--External measurements of four males and one female are,
respectively, as follows: Length of head and body, 134, 128, 134, 134,
136; tail vertebrae, 28, 23, 27, 26, 21; hind foot, 20, 16, 19, 16, 21;
ear, 17, 17, 17, 17, 18; length of forearm, 73.8, 68.0, 74.0, 73.6,
78.0.

_Remarks._--_Nyctimene major scitulus_ has been recorded only from the
western chain of islands in the Solomons (see Fig. 15). Specimens
examined by me agree well in external dimensions and color with
specimens described by Andersen (1912:712) and Troughton (1931:206-207).




ZOOGEOGRAPHY AND SPECIATION


De Beaufort (1951:113) considered bats of "less zoogeographical
importance" than other mammals because the ocean is not an "absolute
barrier to their dispersal." Volant animals are ecologically terrestrial
and therefore are more nearly earthbound than De Beaufort's remarks
would suggest (see Miller, 1966:10). Indeed, many kinds of volant
animals are endemic to the Solomons. Birds, for example, are well
adapted for flight but pose some of the most complex zoogeographic
problems in the area of New Guinea and the Solomon Islands (Mayr,
1940:198; 1942:81-83; Koopman, 1957). Rapid speciation can take place
in any situation where there is a high degree of isolation (Wright,
1931; Lack, 1947). In fact, isolation is a most important factor in
speciation of insular populations (Baker, 1951:55). The one genus, nine
species, and 19 subspecies of megachiropterans that are endemic to the
Solomons (Table 4) obviously indicate that bats, although volant, can be
restricted to one or more islands long enough for new taxa to evolve.

    TABLE 4. A Summary of the Kinds of Megachiropteran Bats in the
       Solomon Islands and Their Affinities with Faunas of Adjacent
       Islands.

  ===========+========+==========+===========+============+===============
             |        |          |  Common   | Common to  |   Common to
             |        | Endemic  |  only to  | Solomons,  |   Solomons,
             | Totals |    to    | Solomons  | Bismarcks, | New Hebrides,
             |        | Solomons |    and    |    and     |      and
             |        |          | Bismarcks | New Guinea | New Caledonia
  -----------+--------+----------+-----------+------------+---------------
  Genera     |    7   |     1    |     0     |      6     |       0
  Species    |   16   |     9    |     1     |      6     |       1
  Subspecies |   20   |    19    |     0     |      0     |       1
  -----------+--------+----------+-----------+------------+---------------

The megachiropteran bats of the Solomons have their affinities with the
fauna of New Guinea (Table 4); the Solomons and New Guinea have six
genera and six species in common. Because the two areas never have been
connected (_via_ the Bismarck Archipelago) by dry land, bats probably
have reached the Solomons by flying from island to island (see Durham,
1963:357, 359, 361, 363). Deignan (1963:266) has dismissed voluntary or
involuntary flight as possible explanations for distributions of bats
and birds on islands of the Pacific.

The taxonomic level of endemism can be used as an indicator of antiquity
(Dobzhansky, 1941; Koopman, 1958:429-430). The one megachiropteran genus
(_Pteralopex_) endemic to the Solomons apparently is an ancient relic.
Bats of this monotypic genus occur on Bougainville, Choiseul, Santa
Ysabel, and Guadalcanal (see Fig. 4). These four islands probably were
contiguous during the maximum lowering of sea level in the Pleistocene
(see Durham, 1963:362-363). Bats of the genus _Pteralopex_ are the only
kind in the Solomons having a distribution that can be correlated with
former land connections between islands.

The distributions of 16 species of megachiropterans known from the
Solomons are summarized in Table 5 and in Figure 16. The larger islands
(in terms of surface area and elevation) in general have the highest
number of species (Guadalcanal 10, Choiseul 9, and Bougainville 8). But
Fauro, one of the smallest islands for which data are available, has six
species of megachiropterans whereas San Cristobal and Malaita, two of
the larger islands, have only three and four species, respectively.
Possibly this difference signals the need for additional collecting.

Bougainville and Choiseul, about 60 miles apart, have seven species of
megachiropterans in common (Table 5). Fauro, 25 miles southeast of
Bougainville and 35 miles west of Choiseul, shares five species with
each of these islands (Fig. 16). _Pteralopex atrata_ and _Pteropus
rayneri_ occur on Choiseul and on Bougainville, but not on Fauro.
Individuals of these species are the largest fruit bats in the Solomons,
and their absence on Fauro suggests, therefore, that this small island
is ecologically unsuitable, at least in some months, for the support of
populations of bats that require relatively large amounts of food. The
small size of the island is consistent with this hypothesis, but several
other islands as small as Fauro do support populations of the large
kinds of _Pteropus_, at least in some months.

    TABLE 5. A Summary of Distribution of All Species of Megachiropteran
       Bats Known from the Solomons. Only Islands Well Known Faunistically
       Are Listed.

  Column headings:

  A: Bougainville     I: Vella Lavella
  B: Choiseul         J: Kolombangara
  C: Santa Ysabel     K: Russell
  D: Ndai             L: Guadalcanal
  E: Malaita          M: San Cristobal
  F: Florida          N: Ugi
  G: Fauro            O: Rennell
  H: Shortland        P: Ontong Java

  ===================+==+==+==+==+==+==+==+==+==+==+==+==+==+==+==+==
        SPECIES      | A| B| C| D| E| F| G| H| I| J| K| L| M| N| O| P
  -------------------+--+--+--+--+--+--+--+--+--+--+--+--+--+--+--+--
  R. amplexicaudatus | X| X| X|  | X|  | X|  |  | X|  | X|  |  |  |
  P. atrata          | X| X| X|  |  |  |  |  |  |  |  | X|  |  |  |
  Pt. hypomelanus    |  |  |  |  |  |  |  |  |  |  | X|  |  |  |  |
  Pt. admiralitatum  |  |  |  | X|  |  |  | X| X|  | X| X|  |  |  |
  Pt. tonganus       |  |  |  |  |  |  |  |  |  |  |  |  |  |  | X|
  Pt. howensis       |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  | X
  Pt. rayneri        | X| X| X|  | X|  |  |  | X| X|  | X| X| X| X|
  Pt. woodfordi      |  |  |  |  |  |  | X|  |  | X| X| X|  |  |  |
  Pt. mahaganus      | X|  | X|  |  |  |  |  |  |  |  |  |  |  |  |
  D. inermis         | X| X| X|  | X| X| X| X| X|  |  | X| X| X| X|
  M. lagochilus      | X| X|  |  |  | X| X|  |  | X|  | X| X|  |  |
  M. woodfordi       | X| X|  |  |  |  | X| X|  |  | X| X|  |  |  |
  M. aurantius       |  | X|  |  |  | X|  |  |  |  |  |  |  |  |  |
  N. albiventer      | X| X| X|  |  |  | X|  |  | X|  | X|  |  |  |
  N. major           |  | X|  |  |  | X|  | X|  |  |  | X|  |  |  |
  N. malaitensis     |  |  |  |  | X|  |  |  |  |  |  |  |  |  |  |
                     +--+--+--+--+--+--+--+--+--+--+--+--+--+--+--+--
      Totals         | 8| 9| 6| 1| 4| 4| 6| 4| 3| 5| 4|10| 3| 2| 3| 1
  -------------------+--+--+--+--+--+--+--+--+--+--+--+--+--+--+--+--

Santa Ysabel has six species of megachiropterans and 10 occur on
Guadalcanal (Table 5). These two islands, separated by about 100 miles
of water, share five species (_Rousettus amplexicaudatus_, _Pteralopex
atrata_, _Pteropus rayneri_, _Dobsonia inermis_, and _Nyctimene
albiventer_). The Nggela Group, in which Florida is the largest island
and the only one from which bats have been collected, is 50 miles
southeast of Santa Ysabel and 30 miles north of Guadalcanal (Fig. 16).
Four species of megachiropterans are known from Florida (_Dobsonia
inermis_, _Macroglossus lagochilus_, _Melonycteris aurantius_, and
_Nyctimene major_). Three of these are known from Guadalcanal and one
occurs on Santa Ysabel. This situation resembles the one involving
Fauro, Bougainville, and Choiseul because none of the large bats
(_Pteropus_ and _Pteralopex_) is known from Florida, even though two
species of large bats that occur on Santa Ysabel to the northwest occur
also on Guadalcanal to the south. Possibly Florida and the smaller
islands that comprise the Nggela Group are ecologically unsuitable for
large bats, or perhaps these small islands can support only limited
numbers of individuals during part of a year.

    [Illustration: FIG. 16. The number of megachiropteran species known
       from individual islands (number within a circle) is compared with
       the number of species common to two different islands (number
       without a circle). For names of islands see Fig. 2.]

Some of the small islands in the Solomons have populations of large
fruit bats. For example, _Pteropus admiralitatum_ and _P. hypomelanus_
have been reported from the small islands in the Russell Group (Table
5). Possibly these species do not live concurrently in the Russells;
specimens of the two were obtained in different years. Two small
megachiropterans, _P. woodfordi_ and _Melonycteris woodfordi_, also
inhabit the Russells. Shortland, a small island about 15 miles south of
Bougainville, supports one large bat, _P. admiralitatum_, as well as
smaller megachiropterans.

Kolombangara and Vella Lavella are about the same size and are separated
by about 15 miles of water. _Rousettus amplexicaudatus_, _Pteropus
rayneri_, _P. woodfordi_, _Macroglossus lagochilus_, and _Nyctimene
albiventer_ have been collected on Kolombangara but only _P.
admiralitatum_, _P. rayneri_, and _Dobsonia inermis_ have been found on
Vella Lavella. The difference in the known megachiropteran faunas is
more striking when one compares each island with adjacent islands. Two
species on Vella Lavella occur also on Choiseul, which is about 35 miles
northeastward, and two species occur also on Shortland, which is 120
miles northwestward (Fig. 16). Four of the five megachiropterans on
Kolombangara also have been found on Choiseul, about 50 miles northward
(Table 5). _Pteropus rayneri_ is the only megachiropteran known from
both Kolombangara and Vella Lavella, even though the islands are
separated by only a few miles of water. Inadequate data possibly account
for the differences in the megachiropteran fauna, but I suspect that
some other factors are involved. Although Vella Lavella and Kolombangara
do have one species (_P. rayneri_) in common, a different subspecies
occurs on each island--_rubianus_ on Kolombangara and _lavellanus_ on
Vella Lavella (Fig. 17 and Table 6). This indicates that some factor or
factors are operating to keep megachiropterans from moving frequently or
easily from one island to the other.

Each of several subspecies of species in the genus _Pteropus_ are known
from one or two small islands separated by only a few miles from other
islands on which different subspecies occur (see Fig. 6). Judging from
this kind of distribution, these bats do not move frequently from island
to island. Possibly this is because they cannot easily cross water
barriers, or are not inclined to do so because food is abundantly
available throughout the year on their home island. Because "flying
foxes" frequently are seen in flight over water several hundred yards
from shore, the first factor probably is unimportant--at least where
short distances are involved. It seems most likely that when abundant
food is available these bats have no reason to move even moderate
distances.

    [Illustration: FIG. 17. The number of subspecies of megachiropterans
       known from individual islands (number within a circle) is compared
       with the number of subspecies common to different islands (number
       without a circle). For names of islands see Fig. 2.]

Distributions of subspecies of polytypic species are summarized in Table
6 and Figure 17. Generally, more subspecies are known from the larger
islands than from the smaller islands (Guadalcanal with 5, Bougainville,
Choiseul, and Santa Ysabel with 4, Fauro with 2.) The distributions of
some subspecies can be used to judge the differential effectiveness of
water gaps between islands. The distribution of _Pteropus rayneri
lavellanus_ and _P. rayneri rubianus_ is an example.

Choiseul and Santa Ysabel are separated by about 50 miles of water (see
Fig. 17) but have three subspecies in common (_Pteropus rayneri
grandis_, _Dobsonia inermis minimus_, and _Nyctimene albiventer
minor_.) Choiseul is about 50 miles from Kolombangara and about 35 miles
from Vella Lavella, but shares no subspecies with these smaller islands
although some species are shared (Tables 5 and 6). From these data one
can conclude that exchange of genes between populations on Choiseul and
populations on Santa Ysabel is frequent but for some reason exchange of
genes between populations on Vella Lavella and Choiseul and Kolombangara
and Choiseul is infrequent. A series of small islands (Rob Roy, Wagina,
and the Arnavon Islands, not named on the maps) connect Choiseul and
Santa Ysabel in stepping-stone fashion (see Fig. 17). Possibly these
small islands enhance movement of megachiropterans between Choiseul and
Santa Ysabel.

    TABLE 6. A Summary of Distribution of Polytypic Species of
       Megachiropteran Bats in the Solomon Islands. Only Islands
       Well Known Faunistically Are Listed.

  Column headings:

  A: Bougainville     I: Vella Lavella
  B: Choiseul         J: Kolombangara
  C: Santa Ysabel     K: Russell
  D: Ndai             L: Guadalcanal
  E: Malaita          M: San Cristobal
  F: Florida          N: Ugi
  G: Fauro            O: Rennell
  H: Shortland        P: Ontong Java

  ===================+==+==+==+==+==+==+==+==+==+==+==+==+==+==+==+==
      SUBSPECIES     | A| B| C| D| E| F| G| H| I| J| K| L| M| N| O| P
  -------------------+--+--+--+--+--+--+--+--+--+--+--+--+--+--+--+--
  P. atrata atrata   | X| X|  |  |  |  |  |  |  |  |  |  |  |  |  |
  P. atrata anceps   |  |  | X|  |  |  |  |  |  |  |  | X|  |  |  |
  Pt. a. solomonis   |  |  |  |  |  |  |  |  | X|  | X| X|  |  |  |
  Pt. a. colonus     |  |  |  |  |  |  |  | X|  |  |  |  |  |  |  |
  Pt. a. grandis     |  |  |  | X|  |  |  |  |  |  |  |  |  |  |  |
  Pt. r. rayneri     |  |  |  |  | X|  |  |  |  |  |  | X|  |  |  |
  Pt. r. grandis     | X| X| X|  |  |  |  |  |  |  |  |  |  |  |  |
  Pt. r. rubianus    |  |  |  |  |  |  |  |  |  | X|  |  |  |  |  |
  Pt. r. lavellanus  |  |  |  |  |  |  |  |  | X|  |  |  |  |  |  |
  Pt. r. monoensis   |  |  |  |  |  |  |  |  |  |  |  |  |  |  |  | X
  Pt. r. cognatus    |  |  |  |  |  |  |  |  |  |  |  |  | X| X|  |
  Pt. r. rennelli    |  |  |  |  |  |  |  |  |  |  |  |  |  |  | X|
  D. i. inermis      | X|  |  |  | X| X| X| X| X|  |  | X| X| X| X|
  D. i. minimus      |  | X| X|  |  |  |  |  |  |  |  |  |  |  |  |
  N. a. bougainville | X|  |  |  |  |  |  |  |  | X|  | X|  |  |  |
  N. a. minor        |  | X| X|  |  |  | X|  |  |  |  |  |  |  |  |
                     +--+--+--+--+--+--+--+--+--+--+--+--+--+--+--+--
      Totals         | 4| 4| 4| 1| 2| 1| 2| 2| 3| 2| 1| 5| 2| 2| 2| 1
  -------------------+--+--+--+--+--+--+--+--+--+--+--+--+--+--+--+--

Florida, of the Nggela Group, is approximately halfway between Santa
Ysabel and Guadalcanal. _Pteralopex atrata anceps_ occurs on Santa
Ysabel and on Guadalcanal but is unknown from Florida. Fauro lies
between Bougainville and Choiseul. _Pteralopex atrata atrata_ and
_Pteropus rayneri grandis_ occur on Choiseul and on Bougainville but are
unknown from Fauro. As suggested earlier, small islands like Fauro and
Florida possibly cannot support large fruit bats, although they probably
would utilize these small islands when in transit between larger
islands.

Fauro apparently is important to the distribution of the two subspecies
of _Dobsonia inermis_ and _Nyctimene albiventer_ in the Solomons (see
Figs. 9 and 13). In both species, one subspecies is found in the eastern
chain of islands and one subspecies is found in the western chain.
Specimens of _Dobsonia inermis_ from Fauro and Bougainville can be
identified as the subspecies _inermis_ whereas those from Choiseul are
assignable to the subspecies _minimus_. _Nyctimene albiventer
bougainville_ occurs on Bougainville but specimens of _N. albiventer_
from Fauro and Choiseul can be identified as the subspecies _minor_.
Although interchange of genes occurs between populations on Bougainville
and Fauro in the case of _D. inermis_, the population of _N. albiventer_
on Fauro is at least partially isolated from the population on
Bougainville.

Rennell and Ontong Java are relatively isolated from other islands in
the Solomons (see Fig. 17). Only one kind of bat (_Pteropus howensis_)
is known from Ontong Java and apparently is endemic to that atoll.
_Pteropus tonganus geddiei_, one of the megachiropterans that occurs on
Rennell (Table 5), also is found in the New Hebrides and on New
Caledonia (Table 4). This makes _P. t. geddiei_ the only megachiropteran
bat in the Solomons that is more closely related to bats on islands to
the southeast of the Solomons than to bats on other islands of the
Solomons, the Bismarcks, or New Guinea, to the north and west. The other
species of megachiropterans (_Dobsonia inermis_ and _Pteropus rayneri_)
on Rennell are found also on other islands in the Solomons. It is to be
noted that Mayr (1931) regarded the avifauna of Rennell as most nearly
like that of the New Hebrides and New Caledonia. He suggested that the
prevailing winds from the southeast have been important for birds that
have reached Rennell. The New Hebrides and New Caledonia are four and a
half times farther from Rennell than are San Cristobal and Guadalcanal.
On first consideration a person might doubt that the winds would be
favorable enough to compensate for the great distance between Rennell
and the New Hebrides and New Caledonia. Darlington (1938) has used the
formula X n/m to obtain a comparison of barriers of different widths.
[X = the probability of an individual crossing a barrier of width m;
the probability of an individual crossing a similar barrier of width n
is the ratio n/m.] If this formula is applied here, one finds that winds
from the southeast (that is, from the New Hebrides and New Caledonia)
would have to be more than 100 times more favorable than winds from the
northeast (from Guadalcanal and San Cristobal) in order to compensate
for the distance of Rennell from the New Hebrides and New Caledonia.
Even so, tropical storms with unusually strong winds, frequent during
some parts of the year, possibly account for the present distributional
pattern of bats and birds that live on Rennell.

Whatever the means by which bats of the species _P. tonganus_ reached
Rennell, the fact remains that specimens from Rennell cannot be
distinguished from specimens of _P. tonganus geddiei_ from the New
Hebrides and New Caledonia, more than 500 miles to the southeast.

NOTE: An important and interesting paper on zoogeography of bats, which
was published too late to be included here, is: Krzanowski, A., 1967,
The magnitude of islands and the size of bats (Chiroptera), Acta Zool.
Cracoviensia, 12:281-348.




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WRIGHT, S.

   1931.  Evolution in Mendelian populations. Genetics, 16:97-159,
          21 figs.


_Transmitted August 10, 1967._




Transcriber's Notes

Obvious typographical and punctuation errors repaired. The "Key to
_Pteropus_ in the Solomon Islands" was moved above the beginning of the
listing for =Pteropus= Brisson. Where figures or tables split
paragraphs, they were moved above or below the split.


Typographical Corrections

  Page  Correction
  ====  =======================
   797  Liuinuwu => Liuniuwu
   809  intermis => inermis
   824  adbiventer => albiventer
   832  Gaudalcanal => Guadalcanal







End of the Project Gutenberg EBook of Systematics of Megachiropteran Bats in
the Solomon Islands, by Carleton J. Phillips

*** END OF THE PROJECT GUTENBERG EBOOK 40112 ***