Chapter I: Part 1

Transcriber's Notes

Text Emphasis ================ _Text_ - Italics =Text= - Bold

Symbolic Representations ============================== [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

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.

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.

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.

=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.

) 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).

), _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).

); _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