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=