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diff --git a/40112.txt b/40112.txt deleted file mode 100644 index e39caaa..0000000 --- a/40112.txt +++ /dev/null @@ -1,3705 +0,0 @@ -The Project Gutenberg EBook of Systematics of Megachiropteran Bats in the -Solomon Islands, by Carleton J. Phillips - -This eBook is for the use of anyone anywhere at no cost and with -almost no restrictions whatsoever. You may copy it, give it away or -re-use it under the terms of the Project Gutenberg License included -with this eBook or online at www.gutenberg.org - - -Title: Systematics of Megachiropteran Bats in the Solomon Islands - -Author: Carleton J. Phillips - -Release Date: July 1, 2012 [EBook #40112] - -Language: English - -Character set encoding: ASCII - -*** START OF THIS PROJECT GUTENBERG EBOOK SYSTEMATICS OF MEGACHIROPTERAN BATS *** - - - - -Produced by Chris Curnow, Joseph Cooper, Tom Cosmas and -the Online Distributed Proofreading Team at -http://www.pgdp.net - - - - - - - -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 - -[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 deg. and -83 deg. F. and at sea level ranges from about 70 deg. to 93 deg. 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 deg. 07' S, 155 deg. 54' E. - -BANIKA.--9 deg. 05' S, 155 deg. 13' E. - -BARA (Gera).--9 deg. 31' S, 160 deg. 31' E. - -BELLONA (Bello).--11 deg. 18' S, 159 deg. 48' E. - -BOUGAINVILLE (Mamamolimo).--6 deg. 12' S, 155 deg. 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 deg. 15' S, 154 deg. 38' E. - -CHOISEUL.--7 deg. 04' S, 157 deg. 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 deg. 55' S, 156 deg. 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 deg. 05' S, 160 deg. 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 deg. 03' S, 156 deg. 35' E. - -GATUKAI.--8 deg. 47' S, 158 deg. 12' E. - -GHIZO (Gizo, Keso).--8 deg. 05' S, 156 deg. 59' E. - -GOWER (N'dai).--7 deg. 54' S, 160 deg. 34' E. - -GUADALCANAL (Guadalcanar).--9 deg. 15' S, 159 deg. 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 deg. 44' S, 155 deg. 28' E. - -KOLOMBANGARA (Duki, Kulambangara).--8 deg. 00' S, 157 deg. 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 deg. 00' S, 161 deg. 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 deg. 49' S, 160 deg. 53' E. - -MONO (Treasury).--7 deg. 22' S, 155 deg. 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 deg. 20' S, 157 deg. 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 deg. 31' S, 154 deg. 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 deg. 31' S, 154 deg. 11' E. - -NUKUMANU (Le Maira, Tasman).--4 deg. 32' S, 159 deg. 25' E. - -ONTONG JAVA (Lord Howe Atoll, Liuniuwu).--5 deg. 25' S, 159 deg. 30' E. - -PAVUVO.--9 deg. 04' S, 159 deg. 08' E. - -RAMOS.--8 deg. 16' S, 160 deg. 11' E. - -RENNELL.--11 deg. 38' S, 160 deg. 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 deg. 21' S, 157 deg. 20' E. - -RUSSELL.--9 deg. 04' S, 159 deg. 12' E. - -SAN CRISTOBAL (San Christoval, Bauro, Makira, Arussi).--11 deg. 33' S, 161 deg. -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 deg. 00' S, -159 deg. 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 deg. 08' S, 159 deg. 49' E. - -SHORTLAND.--7 deg. 03' S, 155 deg. 47' E. - -SIKAIANA (Stewart).--8 deg. 22' S, 162 deg. 44' E. - -SIMBO (Narovo, Naorovo, Naravo, Navoro, Sembo).--8 deg. 16' S, 156 deg. 31' E. - -STIRLING.--7 deg. 25' S, 155 deg. 35' E. - -TANABULI (Tanambuli, Tunnibili, Tunnibilis, Tunnibul, Tunnivula).--8 deg. -24' S, 159 deg. 35' E. - -TAUU (Marqueen, Mortlock).--4 deg. 48' S, 157 deg. 32' E. - -TELIPARI.--8 deg. 15' S, 157 deg. 32' E. - -UGI.--10 deg. 14' S, 161 deg. 44' E. - -VANGUNO (Vangunu).--8 deg. 39' S, 158 deg. 00' E. - -VELLA LAVELLA.--7 deg. 43' S, 156 deg. 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 deg. 20' S, 161 deg. 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. Saeugetierk., - 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, Mem. 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. Saeugetierk., 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 deg. 34' E, was incorrectly listed in Laurie -and Hill (1954:152) as 159 deg. 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. - Saeugetierk., 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) E. 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. Saeugetierk., - 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. Saeugetierk., 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. Saeugetierk., 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_ E. 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. Saeugetierk., - 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. - - - - -LITERATURE CITED - - -ANONYMOUS. - - 1944. Gazetteer of Solomon Islands, Bismarck Archipelago, and - Islands of the south-eastern end of New Guinea. Hydrographic - office of the United States Navy Department, No. 881. - - -ANDERSEN, K. - - 1908. Twenty new forms of _Pteropus_. Ann. Mag. Nat. Hist., ser. 8, - 2:361-370, October. - - 1909a. On the characters and affinities of "Desmalopex" and - Pteralopex. Ann. Mag. Nat. Hist., ser. 8, 3:213-222, February. - - 1909b. Two new bats from the Solomon Islands. Ann. Mag. Nat. Hist., - ser. 8, 3:266-270, March. - - 1909c. On the fruit-bats of the genus _Dobsonia_. Ann. Mag. Nat. - Hist., ser. 8, 4:528-533, December. - - 1911. Six new fruit-bats of the genera _Macroglossus_ and - _Syconycteris_. Ann. Mag. Nat. Hist., ser. 8, 7:641-643, June. - - 1912. Catalogue of the Chiroptera in the collection of the British - Museum, British Mus. (Nat. Hist.), London, 1:ci + 1-854, 79 figs. - - -BADER, R. S., and HALL, J. S. - - 1960. Osteometric variation and function in bats. Evol., 14:8-17, 3 - figs., March 21. - - -BAKER, R. H. - - 1951. The avifauna of Micronesia, its origin, evolution, and - distribution. Univ. Kansas Publ., Mus. Nat. Hist., 3:1-359, 16 - figs., June 12. - - -BEAUFORT, L. E. DE. - - 1951. Zoogeography of the land and inland waters. Sidgwick and - Jackson, London, viii + 208 pp., maps. - - -BELKIN, J. N. - - 1962. The mosquitoes of the South Pacific (Diptera, Culicidae). - Univ. California Press, Berkeley, 1:xii + 1-608, July 18. - - -BENEDICT, F. - - 1957. Hair structure as a generic character in bats. Univ. - California Publ. Zool., 59:285-548, 9 pls., 4 figs., October 10. - - -BRIGHAM, W. T. - - 1900. An index to the islands of the Pacific Ocean. Mem. Bernice P. - Bishop Mus., Honolulu, 2:1-170, maps. - - -CRANSTONE, B. A. L. - - 1961. Melanesia: a short ethnography. British Mus., London, 115 - pp., 26 pls., 43 figs., map. - - -DARLINGTON, P. J. - - 1938. The origin of the fauna of the Greater Antilles, with - discussion of dispersal of animals over water and through the - air. Quart. Rev. Biol., 13:274-300, 5 figs. - - -DEIGNAN, H. G. - - 1963. Birds in the tropical Pacific, pp. 263-269, _in_ Pacific - Basin biogeography (J. L. Gressitt, ed.). Bernice P. Bishop Mus. - Press, Honolulu, xi + 563 pp. - - -DOBSON, G. E. - - 1878. Catalogue of the Chiroptera in the ... British Museum. - British Mus., London, xlii + 567 pp., 30 pls. - - -DOBZHANSKY, T. G. - - 1941. Genetics and the origin of species. Columbia Univ. Press, New - York, 2nd. ed., rev., ix + 466 pp., 24 figs. - - -DURHAM, J. W. - - 1963. Paleogeographic conclusions in light of biological data, pp. - 355-365, 4 figs., _in_ Pacific Basin biogeography (J. L. - Gressitt, ed.). Bernice P. Bishop Mus. Press, Honolulu, - xi + 563 pp. - - -ELLERMAN, J. R., and MORRISON-SCOTT, T. C. S. - - 1966. Checklist of Palaearctic and Indian mammals, 1758-1946. - British Mus. (Nat. Hist.), London, 2nd. ed., 810 pp., July. - - -FELTEN, H. - - 1964a. Flughunde der Gattung _Pteropus_ von den Neuen Hebriden - (Mammalia, Chiroptera). Senck. biol., 45:87-92, 6 figs., May 15. - - 1964b. Flughunde der Gattung _Pteropus_ von Neukaledonien und den - Loyalty-Inseln (Mammalia, Chiroptera). Senck. biol., 45:671-683, - December 21. - - -HALL, E. R. - - 1946. Mammals of Nevada. Univ. California Press, Berkeley, xi + 710 - pp., frontispiece, 11 pls., 485 figs., July 1. - - -HILL, J. E. - - 1956. The mammals of Rennell Island. The natural history of Rennell - Island, British Solomon Islands, Copenhagen, 1:73-84, - November 28. - - 1962. A little-known fruit-bat from Rennell Island. The natural - history of Rennell Island, British Solomon Islands, Copenhagen, - 4:7-9, February 15. - - 1963. A revision of the genus _Hipposideros_. Bull. British Mus. - (Nat. Hist.), 2:1-129, October. - - -KOOPMAN, K. F. - - 1957. Evolution in the genus _Myzomela_ (Aves: Meliphagidae). The - Auk, 74:49-72, 5 figs., January. - - 1958. Land bridges and ecology in bat distribution on islands off - the northern coast of South America. Evol., 12:429-439, 2 figs., - December. - - -LACK, D. L. - - 1947. Darwin's finches. Cambridge Univ. Press, Cambridge, x + 209 - pp., 8 pls., 27 figs. - - -LAURIE, E. M. O., and HILL, J. E. - - 1954. List of land mammals of New Guinea, Celebes, and adjacent - islands. British Mus. (Nat. Hist.), London, 175 pp., June 30. - - -LAWRENCE, B. - - 1945. Three new Pteropus from New Caledonia and the Solomons. Proc. - New England Zool. Club, 23:59-69, March 26. - - -LEVER, R. J. A. W. - - 1934. Notes on mosquitoes of the British Solomon Islands. British - Solomon Islands Agr. Gaz., 2:16. - - -LEWIS, A. B. - - 1951. The Melanesians: People of the South Pacific. Chicago Nat. - Hist. Mus. Press, Chicago, 259 pp., 56 figs., maps, September. - - -MATSCHIE, P. - - 1899. Die Megachiroptera des Berliner Museums fuer Naturkunde. Druck - und verlag von George Reimer, Berlin, viii + 102 pp., 14 figs. - - -MAYR, E. - - 1931. A systematic list of the birds of Rennell Island with - descriptions of new species and subspecies. Amer. Mus. Novit., - 486:1-29, August 29. - - 1940. The origin and the history of the bird fauna of Polynesia. - Proc. 6th Pacific Sci. Cong., 1939, 4:197-216. - - 1942. Systematics and the origin of species from the viewpoint of a - zoologist. Columbia Univ. Press, New York, xiv + 334 pp., - 29 figs. - - -MILLER, A. H. - - 1966. Animal evolution on islands, pp. 10-16, _in_ The Galapagos - (R. I. Bowman, ed.), Univ. California Press, Berkeley. - - -MILLER, G. S. - - 1907. The families and genera of bats. Bull. U. S. Nat. Mus., - 57:xvii + 282, 14 pls., 49 figs., June 29. - - -PHILLIPS, C. J. - - 1966. A new species of bat of the genus _Melonycteris_ from the - Solomon Islands. Jour. Mamm., 47:23-27, 1 fig., March 12. - - 1967. A new subspecies of horseshoe bat (_Hipposideros diadema_) - from the Solomon Islands. Proc. Biol. Soc. Washington, 80:35-40, - 2 figs., March 24. - - -POHLE, H. - - 1953. Ueber die Fledertiere von Bougainville. Z. Saeugetierk., - 17:127-137, October 27. - - -RIDGWAY, R. - - 1912. Color standards and color nomenclature. Washington, D. C., - iv + 44 pp., 53 pls. - - -SANBORN, C. C. - - 1931. Bats from Polynesia, Melanesia, and Malaysia. Publ. Field - Mus. Nat. Hist., Zool. Ser., 18:7-29, February 12. - - -SANBORN, C. C., and BEECHER, W. J. - - 1947. Bats from the Solomon Islands. Jour. Mamm., 28:387-391, - November 19. - - -SANBORN, C. C., and NICHOLSON, A. J. - - 1950. Bats from New Caledonia, the Solomon Islands, and New - Hebrides. Fieldiana (Zool.), 31:313-338, 6 figs., August 31. - - -SIMPSON, G. G. - - 1945. The principles of classification and a classification of - mammals. Bull. Amer. Mus. Nat. Hist., 85:xvi + 350, October 5. - - -TATE, G. H. H. - - 1934. An apparently new fruit bat of the _Pteropus hypomelanus_ - group from Gower Island, Solomon Islands. Amer. Mus. Novit., - 718:1-2, May 4. - - 1942. Pteropodidae (Chiroptera) of the Archbold Collections. Bull. - Amer. Mus. Nat. Hist., 80:331-347, December 31. - - -THOMAS, O. - - 1887a. Diagnoses of two new fruit-eating bats from the Solomon - Islands. Ann. Mag. Nat. Hist., ser. 5, 60:147, February. - - 1887b. On the bats collected by Mr. C. M. Woodford in the Solomon - Islands. Proc. Zool. Soc. London, 23:320-328, 3 figs., 2 pls., - March 15. - - 1888a. Diagnoses of six new mammals from the Solomon Islands. Ann. - Mag. Nat. Hist., ser. 6, 2:155-158, February. - - 1888b. The mammals of the Solomon Islands, based on the collections - made by Mr. C. M. Woodford during his second expedition to the - Archipelago. Proc. Zool. Soc. London, 33:470-484, 3 pls., - December 4. - - -TROUESSART, E. L. - - 1904. Catalogus Mammalium tam viventium quam fossilium. - Quinquennale Supplementum, Berlin, vii + 929 pp. - - -TROUGHTON, E. LEG. - - 1931. Three new bats of the genera _Pteropus Nyctimene_, and - _Chaerephon_ from Melanesia. Proc. Linn. Soc. New South Wales, - 56:204-209, June 15. - - 1936. The mammalian fauna of Bougainville Island, Solomons Group. - Rec. Australian Mus., 14:341-353, April 7. - - -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. 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