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diff --git a/40112-0.txt b/40112-0.txt new file mode 100644 index 0000000..06a1e23 --- /dev/null +++ b/40112-0.txt @@ -0,0 +1,3317 @@ +*** START OF THE PROJECT GUTENBERG EBOOK 40112 *** + +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° and +83° F. and at sea level ranges from about 70° to 93° F. yearly (Belkin, +1962:42). + +Southeast tradewinds are relatively constant from May to October and +this period, in general, is a dry season except at higher elevations on +windward coasts. From December to March prevailing winds are from the +north and precipitation throughout the island group is especially heavy. +Rainfall on the island of Tulagi averages about 120 inches per year +(Bryan, Edwin H., 1941; MS, p. 2, at Pacific Sci. Information Center, +Bishop Museum) and up to 300 inches have been recorded on the north +coast of Guadalcanal (Belkin, 1962:42-43). Occasional dry periods occur +even in the period of December to March. + +Most islands of the Solomon Group support dense tropical rain forest. +Much of it has been modified by man. Some clearings and scattered +coconut plantations are found along coasts. On some of the larger +islands (for example, Guadalcanal) coastal scrub (especially on leeward +coasts) and extensive grassy areas are to be found. Additional notes on +vegetation are in the gazetteer. + +The 165,000 persons living on the Solomon Islands are mostly +Melanesians but some are mixed Papuan, Malay, and Polynesian. These +native peoples are notorious for their cannibalistic tendencies; the +eating of human flesh usually was related to warfare, although +malefactors and human sacrifices accounted for some of the cannibalism +(Cranstone, 1961:29). Prior to the Second World War few Europeans +visited the Solomons and several islands still remain beyond reach of +modern-day technology. For example, Rennell and Bellona islands, south +of the main part of the archipelago, are visited only rarely, and then +only by a medical officer or the Resident Commissioner. According to +Troughton (1936:341), the islanders in the interior of Bougainville as +late as 1935, were prone to kill and feast upon strangers. In 1932, +Lewis (1951:37) felt that the natives of Malaita Island were especially +resistant to outside interference by Caucasians and reported that no +"white man or foreigner" was safe on Malaita. + +Troughton (1936), who listed Melanesian names for mammals, indicated +that the native peoples distinguished between kinds of bats that closely +resembled one another. Of these, the only bats that seem to be used as +food belong to the genus _Pteropus_. + + + + +GAZETTEER + + +In the following list, currently-used names of islands are given; when +available, older names and variant spellings are indicated in +parentheses. For certain islands, especially those visited by field +parties from the Bishop Museum or those frequently mentioned in previous +literature on bats, some descriptive and ecological information also is +provided. + +Latitude and longitude of islands are from publication no. 881 of the +Hydrographic Office of the United States Navy Department (Anonymous, +1944); names of islands were checked against a list by Brigham (1900); +descriptive information mostly is from reports by Temple and Straatmann +(1964, field notes, at the Department of Entomology, Bishop Museum). + +ALU.--7° 07' S, 155° 54' E. + +BANIKA.--9° 05' S, 155° 13' E. + +BARA (Gera).--9° 31' S, 160° 31' E. + +BELLONA (Bello).--11° 18' S, 159° 48' E. + +BOUGAINVILLE (Mamamolimo).--6° 12' S, 155° 15' E. This is the largest +island in the Solomon Group, being 127 miles long (northwest to +southeast) and about 59 miles across at the widest place. The highest +elevations are 9850 and 10171 feet, at the tops of active volcanoes. +Ecologically, Bougainville is mostly dense rain forest, which is less +dense on the summits of higher mountains. + +BUKA.--5° 15' S, 154° 38' E. + +CHOISEUL.--7° 04' S, 157° 01' E. This island, formed along a +northwest-southeast line of low mountains (maximum elevation of 3500 +feet), is about 90 miles long and 20 miles wide. Most collecting was at +Malangona (Sasamunga on some maps) on the southwestern coast. + +FAURO.--6° 55' S, 156° 07' E. This small island, about 14 miles long +(north-south) and six miles wide (east-west), lies about 10 miles south +and east of Bougainville. Fauro is formed around a volcanic cone having +a maximum elevation of 1925 feet; it has considerable dense mangrove +swamp along the west coast, and mature rain forest with little +understory growth. Most collecting was at Toumoa, on one of two southern +peninsulas. + +FLORIDA (Nggela).--9° 05' S, 160° 16' E. Florida, the main island in the +Nggela Island Subgroup, is mountainous and except for some small grassy +areas, supports dense rain forest. It is nearly 25 miles long +(east-west) and nine miles wide (north-south), with a maximum +elevation, at Mount Barnett, of about 1366 feet. Most collecting was at +Haleta, on the southwestern coast. At this locality there were scattered +mangrove swamps, rain forest, and gardens inland. + +GANONGGA (Ronogo, Ronongo).--8° 03' S, 156° 35' E. + +GATUKAI.--8° 47' S, 158° 12' E. + +GHIZO (Gizo, Keso).--8° 05' S, 156° 59' E. + +GOWER (N'dai).--7° 54' S, 160° 34' E. + +GUADALCANAL (Guadalcanar).--9° 15' S, 159° 35' E. Guadalcanal is mostly +of volcanic origin and has an irregular chain of mountains along the +southern coast. The highest elevation is 8005 feet at Mount Popomanasiu. +This large island is nearly 80 miles long (east-west) and 25 miles wide +(north-south). Most of the northwestern part of Guadalcanal supports +_alang-alang_ grass. The remainder of the island is heavily wooded. + +KILINAILAU (Cartaret).--4° 44' S, 155° 28' E. + +KOLOMBANGARA (Duki, Kulambangara).--8° 00' S, 157° 05' E. Kolombangara, +formed from an extinct volcano, is about 18 miles in diameter and nearly +circular. The highest peaks, rising as precipitous cliffs in some +places, reach a maximum elevation of about 5000 feet. The vegetation is +mostly virgin rain forest. Mangrove swamp and small coconut groves occur +along the coast. Field parties from the Bishop Museum were able to reach +the highest elevations, and concentrated their work along the +southwestern side of the island. + + [Illustration: FIG. 2. Solomon Islands. Principal islands + are named.] + +MALAITA (Mala, Malanta, Malayta).--9° 00' S, 161° 00' E. This long (104 +miles northwest to southeast), narrow (about 23 miles at its widest +spot) island, between Santa Ysabel and San Cristobal islands, is +basically of volcanic origin with some limestone (coral) deposits along +the coast. Mount Kolovrat, having an elevation of 4275 feet, is the +highest point. The Bishop Museum field party lived at Dala, in dense +rain forest about 12 miles north of Auki on the northwestern coast of +Malaita. + +MALAPA.--9° 49' S, 160° 53' E. + +MONO (Treasury).--7° 22' S, 155° 35' E. This is a small island (maximum +elevation 1150 feet) in the Treasury Island Subgroup just south of +Bougainville. Mono is about nine miles long (east-west) and five and one +half miles wide (north-south). The basic volcanic core is described in +field notes as topped with coral limestone. + +NEW GEORGIA (Kausagi).--8° 20' S, 157° 30' E. The New Georgia Subgroup +is composed of 11 moderate-sized islands and islets. New Georgia Island, +the main member of the subgroup, is 50 miles long (northwest to +southeast) and from five to 30 miles wide. On the northern side several +volcanic peaks attain an elevation of about 3000 feet. The entire island +is forested. + +NGGELA (Florida Islands).--4° 31' S, 154° 11' E. This subgroup consists +of several small to medium-sized islands between Guadalcanal and +Malaita. Florida is the main island. + +NISSAN (Green, Sir Charles Hardy's).--4° 31' S, 154° 11' E. + +NUKUMANU (Le Maira, Tasman).--4° 32' S, 159° 25' E. + +ONTONG JAVA (Lord Howe Atoll, Liuniuwu).--5° 25' S, 159° 30' E. + +PAVUVO.--9° 04' S, 159° 08' E. + +RAMOS.--8° 16' S, 160° 11' E. + +RENNELL.--11° 38' S, 160° 14' E. This island, of limestone (coral) +origin, along with Bellona, is nearly 100 miles southwest of any other +member of the Solomons and has been regarded, because of this distance, +as an oceanic island instead of a continental island. It is about 50 +miles long (east-west) and nine miles wide (north-south); its highest +elevation is 500 feet. + +ROVIANA (Rendova, Rovianna, Rubiana).--8° 21' S, 157° 20' E. + +RUSSELL.--9° 04' S, 159° 12' E. + +SAN CRISTOBAL (San Christoval, Bauro, Makira, Arussi).--11° 33' S, 161° +43' E. This island is composed mostly of ancient volcanic rock, has a +maximum elevation of 4100 feet, is nearly 70 miles long (northwest to +southeast) and 24 miles wide, and supports a dense rain forest. + +SANTA YSABEL (George, Ysabel, San Isabel, Isbel, Mahaga).--8° 00' S, +159° 07' E. Santa Ysabel is a long (90 miles from northwest to +southeast), narrow (19 miles at the widest spot), forested island, +consisting of a single chain of volcanic mountains. The numerous bays +and mouths of rivers provide excellent anchorages. Collecting was at +Tatamba approximately two miles south of Tanambuli where the +considerable area of forest was dense and bamboo thickets were abundant. + +SAVO (Savu).--9° 08' S, 159° 49' E. + +SHORTLAND.--7° 03' S, 155° 47' E. + +SIKAIANA (Stewart).--8° 22' S, 162° 44' E. + +SIMBO (Narovo, Naorovo, Naravo, Navoro, Sembo).--8° 16' S, 156° 31' E. + +STIRLING.--7° 25' S, 155° 35' E. + +TANABULI (Tanambuli, Tunnibili, Tunnibilis, Tunnibul, Tunnivula).--8° +24' S, 159° 35' E. + +TAUU (Marqueen, Mortlock).--4° 48' S, 157° 32' E. + +TELIPARI.--8° 15' S, 157° 32' E. + +UGI.--10° 14' S, 161° 44' E. + +VANGUNO (Vangunu).--8° 39' S, 158° 00' E. + +VELLA LAVELLA.--7° 43' S, 156° 40' E. The coastline is rugged and +indented by numerous small bays. Some peaks are 3000 feet high. The +southeastern half of Vella Lavella is said to consist of uplifted coral, +and to be thickly planted to coconut palms. The native population is +concentrated here. The northwestern half of the island is rain forest +and is nearly uninhabited. Most of the collecting was at Pusisama, on +the southern beach and on Ulo Crater, an extinct volcano at the middle +of the island. + +YANNTA.--10° 20' S, 161° 20' E. + + + + +METHODS AND MATERIALS + + +The phylogenetic arrangement and nomenclature in the text beyond are +mainly that of Laurie and Hill (1954). The synonymies for accounts of +genera are as follows: (1) first use of the generic name employed along +with the original description, and (2) original proposals, in +chronological order, of other generic names subsequently applied to the +bat in the Solomons. The synonymies in accounts of species and +subspecies are as follows: (1) first use of the accepted name, followed +by its type locality, followed, in chronological order, by other +references to the first name-combination, (2) first use of the +name-combination employed herein (if different from the original +combination), followed, in chronological order, by other references to +the present name-combination, and (3) other name-combinations, in +chronological order, employed for the bat in the Solomons. The word +"part" is used in parentheses after a name if some specimens listed +under that name are from the Solomon Islands and are referable to the +species or subspecies being written about. + +Unless noted otherwise, specimens listed as examined were prepared +originally as museum skins with skulls. Approximately 70 per cent of +bats collected in the Solomons were preserved in formalin and now are +stored in alcohol. Because it was necessary to obtain dimensions and +examine various morphological characteristics of skulls, many crania +were extracted from bats preserved in alcohol. + +Although all specimens in the Bishop Museum from the Solomon Islands +have been catalogued with the prefix BBM-BSIP, catalogue numbers without +prefixes in the lists of specimens examined refer to this museum. +Catalogue numbers with the prefix USNM refer to specimens in the U. S. +National Museum and those with the prefix AM-M refer to specimens in the +Australian Museum. + +Unless indicated otherwise, all measurements in this paper are in +millimeters and are of adults. Cranial measurements, and external +measurements of specimens stored in alcohol, were taken by me. The +cranial measurements were taken with dial calipers using techniques +described by Hall (1946:672-685). External measurements (except length +of forearm) of specimens originally prepared as dried study skins, were +transcribed from specimen labels. + +Capitalized color nomenclature is from Ridgway (1912). Noncapitalized +color terms are from published reports that did not use Ridgway's +terminology. + + + + +ACKNOWLEDGMENTS + + +Financial support for this investigation was from (1) a United +States Army Medical Research and Development Command grant +(DA-MD-49-193-62-G65) to the Entomology Department of the Bernice P. +Bishop Museum, and (2) a National Science Foundation grant (2185-4703) +to the author, through the Committee on Systematics and Evolutionary +Biology of The University of Kansas. I am grateful to many individuals +who have helped me in various ways throughout the course of this study. +Dr. J. Linsley Gressitt, Chairman of the Entomology Department, Bernice +Bishop Museum, allowed me to study specimens collected by his +expeditions; Professors E. Raymond Hall and J. Knox Jones, Jr., of the +Museum of Natural History and the Department of Zoology, The University +of Kansas, offered advice and guidance and constructively reviewed the +manuscript. Other persons who have given me assistance and, in some +cases, arranged for loans of comparative materials, are: Dr. David H. +Johnson, Division of Mammals, United States National Museum; Mr. Hobart +M. Van Deusen and Dr. Richard G. Van Gelder, Archbold Expeditions and +Department of Mammalogy, American Museum of Natural History; Messrs. +Ellis LeG. Troughton and Basil Marlow, Mammal Department, The Australian +Museum; Dr. Joseph Curtis Moore, Department of Mammalogy, Field Museum +of Natural History; Mr. John Edwards Hill, Mammal Room, British Museum +(Natural History); Prof. William B. Davis, Department of Zoology, Texas +A & M University; Miss Barbara Lawrence, Museum of Comparative Zoology, +Harvard University. Messrs. Jerry R. Choate and H. H. Genoways, two +colleagues in zoology at The University of Kansas, have assisted me in +many ways, for which I am grateful. Linda Anne Phillips, my wife, +prepared many of the figures and tables used herein. I thank also +Setsuko Nakata, Edwin H. Bryan, Robert Bowan, and Ilse Koehler, who, as +staff members of the Bishop Museum, were especially helpful to me. Most +of the specimens reported herein were collected by Philip Temple and +Peter Shanahan. + + + + +Key to Genera + + 1. Uropatagium lacking, or, if present, deeply indented in + center; tail vertebrae absent, or if present, free 2 + + 1'. Uropatagium present, not indented; tail vertebrae present, + free or in uropatagium MICROCHIROPTERA 1 + + 2(1). External tail-vertebrae lacking, or, if present, less + than 3 mm long 3 + + 2'. External tail-vertebrae more than 3 mm long 6 + + 3(2). Small or medium-sized (forearm less than 50); + tongue long, extensile 4 + + 3'. Large (forearm more than 80); tongue not long and + extensile 5 + + 4(3). Uropatagium present; small claw present on second phalanx + of second digit; tail short (about 3 mm) + =Macroglossus=, p. 812 + + 4'. Uropatagium absent; no claw on second phalanx of second + digit; no tail =Melonycteris=, p. 814 + + 5(3'). Entire back set with hair; wing membranes not meeting + at middle of back =Pteropus=, p. 793 + + 5'. Back naked; wing membranes meeting at middle of back, + =Pteralopex=, p. 790 + + 6(2'). Nostrils having definite tubelike extensions + =Nyctimene=, p. 817 + + 6'. Nostrils lacking tubelike extensions 7 + + 7(6'). Forearm less than 80; large, sharp claw on second phalanx + of second digit; four upper incisors =Rousettus=, p. 787 + + 7'. Forearm more than 90; small, blunt claw on second phalanx + of second digit; two upper incisors =Dobsonia=, p. 807 + + +Family PTEROPODIDAE + + +Subfamily Pteropodinae + + +Rousettus Gray + + 1821. _Rousettus_ Gray, London Medical Repository, 15:299, April 1. + + 1843. _Xantharpyia_ Gray, List of species ... British Museum, p. 37. + + 1852. _Cynonycteris_ Peters, Reise nach Mossambique, p. 25. + +The genus _Rousettus_ occurs throughout the tropical regions of the Old +World, and in the Solomons is readily distinguished from all other +megachiropteran genera by having both a small claw on the second digit +and free caudal vertebrae. The oriental species have been divided into +two groups on the basis of size (Tate, 1942:344). The subspecies +_Rousettus amplexicaudatus hedigeri_ appears to be the sole +representative of this genus in the Solomon Islands. Prior to 1953, +several workers (Thomas, 1887b:323, 1888b:475; Matschie, 1899:68; +Sanborn, 1931:11) used the name _Rousettus amplexicaudatus brachyotis_ +for it, but Pohle (1953) suggested that the specimens from the Solomons +recorded by earlier workers were _R. a. hedigeri_ named by him on the +basis of the specimen that he saw from Bougainville. + + +=Rousettus amplexicaudatus= + +_Rousettus amplexicaudatus_ has at least three subspecies, one of which +is endemic to the Solomon Islands. The species is wide-ranging, being +known from as far west as Thailand (Ellerman and Morrison-Scott, +1966:93) and as far east as the Solomons. + + [Illustration: FIG. 3. Distribution of _Rousettus amplexicaudatus + hedigeri_. For names of islands see Fig. 2.] + + +=Rousettus amplexicaudatus hedigeri= Pohle + + 1953. _Rousettus amplexicaudatus hedigeri_ Pohle, Z. Säugetierk., + 17:127, October 27, type from Bougainville. + + 1887. _Cynonycteris brachyotis_, Thomas, Proc. Zool. Soc. London, + p. 323, March 15; 1888, Thomas, Proc. Zool. Soc. London, p. 475, + December 4, from Fauro. + + 1889. _Xantharpyia brachyotis_, Matschie, Die Megachiroptera ... + naturkunde, p. 68, from Guadalcanal. + + 1912. _Rousettus brachyotis_, Andersen, Catalogue of the Chiroptera + ... British Museum, 1:809; 1931, Sanborn, Publ. Field Mus. Nat. + Hist., Zool. Ser., 18:11, February 12, from Santa Ysabel. + +_Specimens examined_ (20 males and 21 females; all in alcohol; ten +crania extracted and cleaned).--Guadalcanal in May, 23863, 23915; Fauro +in April, 23804-5; Malaita in June, 24079; Choiseul in March, 23563-4, +23616, 23627, 23630, 23632-3, 23642, 23658, 23663-4, 23680, 23692-3, +23713, 23722; Kolombangara in January and February, 23343, 23366, +23382-4, 23389-90, 23408-9, 23424, 23455, 23471-4, 23501. + +_Measurements._--Average and extreme external measurements of 13 males +and 18 females are, respectively, as follows: Length of head and body, +104.4 (99-118), 108.6 (104-117); tail vertebrae, 16.8 (13-19), 17.6 +(15-24); hind foot, 18.0 (16-19), 16.2 (12-18); ear, 15.9 (15-17), 15.0 +(14-16); length of forearm, 70.1 (66.0-74.1), 68.1 (65.0-69.1). Average +and extreme measurements of skulls of five males and five females are, +respectively, as follows: Greatest length of skull, 33.2 (33.0-33.7), +31.5 (30.9-32.1); condylobasal length, 31.3 (30.9-31.9), 30.1 +(29.3-30.8); palatal length, 14.0 (13.3-14.8), 13.3 (13.0-13.7); +zygomatic breadth, 20.8 (19.8-21.8), 19.4 (18.7-20.8); length of +maxillary tooth-row, 11.0 (10.9-11.3), 10.3 (10.1-10.6); length of +mandibular tooth-row, 12.6 (12.4-12.9), 11.8 (11.7-12.2). + +_Remarks._--The specimens from Choiseul, Kolombangara, and Malaita +islands provide new records of distribution for _Rousettus +amplexicaudatus hedigeri_ (Fig. 3). It was described as smaller than _R. +a. brachyotis_ Dobson, which is known from New Guinea, Amboina, and the +Bismarck Archipelago (Pohle, 1953:127-128). Andersen (1912:809) gave the +range of length of forearm in _R. a. brachyotis_ as 73-81, whereas Pohle +(1953:127) gave the length of forearm of the type specimen of _R. a. +hedigeri_ (adult male) as 67. Measurements of specimens examined by me +indicate that _hedigeri_ occurs throughout the Solomon Islands. Cranial +measurements of my specimens and Pohle's type are less than those of _R. +a. brachyotis_ (see Andersen, 1912:48). + +Sanborn (1931:11) noted that the forearms of three males examined by him +were longer than that of a female. Mean and range for length of forearm +of males and females listed herein, respectively, are 70.1 (66.0-74.1) +and 68.1 (65.0-69.1). Also, each of seven cranial measurements taken by +me averaged more in males than in females. Sagittal and lambdoidal +crests are more prominent in males than in females. + + TABLE 1. A Summary of Breeding Data for Females of _Rousettus + amplexicaudatus hedigeri_ Collected December to June. + + ===========+===========+==============+===========+============= + | Total | Number | | Number of + MONTH | number | adult [F][F] | Number | immature + | collected | collected | lactating | individuals + -----------+-----------+--------------+-----------+------------- + December | 3 | 3 | 3 | 0 + January | 11 | 11 | 8 | 0 + February | 6 | 0 | -- | 1 + March | 16 | 1 | 0 | 9 + April | 2 | 2 | 0 | 0 + June | 1 | 1 | 0 | 0 + -----------+-----------+--------------+-----------+------------- + +As shown in Table 1, adult females obtained in December and January were +lactating when captured whereas those obtained in March, April, and +June were not. More than half of the individuals collected in March were +immature (judging from small size, unfused epiphyses, and lack of wear +on teeth). The immature individuals probably had been nursing in +December and January. + + + +=Pteralopex= Thomas + + 1888. _Pteralopex_ Thomas, Ann. Mag. Nat. Hist., ser. 6, 1:155, + February 1. + + 1762. _Pteropus_ Brisson, Regnum animale ..., ed. 2, p. 153. + +_Pteralopex_, with one species and two subspecies, is the only +megachiropteran genus endemic to the Solomons. Thomas (1888b:475) +considered this unusual bat a relic, isolated from the time when +pteropodids had cuspidate cheek-teeth. Although two workers (Matschie, +1899:11; Simpson, 1945:54) have synonymized _Pteralopex_ with +_Pteropus_, I regard _Pteralopex_ as a morphologically distinct genus. + +Individuals of _Pteralopex_ can be distinguished from all species of +_Pteropus_ in the Solomon Islands by the following features: wing +membranes originate along dorsal midline; braincase diminutive relative +to rest of skull; sagittal crest pronounced; cheek-teeth cuspidate, +broad and massive; i2 about 10 times larger than i1; upper canines with +well-developed secondary cusp; postorbital process fused with zygomatic +arch, forming complete bony ring around orbit. + +Andersen (1909a:216; 1912:436) considered the relationships of +_Pteralopex_ and _Pteropus_ and concluded that _Pteropus pselaphon_ Lay, +1829, from the Sulphur Islands east of Taiwan, and _Pteropus samoensis_ +Peale, 1848, from the Samoan Islands, were the "closest" living +relatives of _Pteralopex_. He stated further that _Pteralopex_ "presents +in fact scarcely a single character which is not either developed to a +certain extent or at least distinctly foreshadowed in _Pteropus +pselaphon_, _pilosus_, _tuberculatus_, or _leucopterus_." In summary, +Andersen thought several species of _Pteropus_ had undergone +evolutionary development resembling that in _Pteralopex_, and that the +latter, with its massive, cuspidate cheek-teeth, could be considered a +highly modified _Pteropus_. For this hypothesis to be plausible, one +must assume that the originally complex cheek-teeth of pteropodids +became simple and, at least in the case of _Pteralopex_, secondarily +became complex once again. According to present-day theory of +evolutionary development, his hypothesis is improbable. Thomas +(1888b:475) probably was correct when he considered _Pteralopex_ an +isolated relic. + +Although _Pteralopex_ usually is listed after _Pteropus_ in phylogenetic +arrangements (see, for example, Sanborn, 1931:21; Pohle, 1953:129; +Laurie and Hill, 1954:40), I have placed _Pteralopex_ before _Pteropus_. + + +=Pteralopex atrata= + +Two subspecies of _Pteralopex atrata_ (_P. a. atrata_ and _P. a. +anceps_) have been named; specimens of both are rare in museum +collections. Thomas (1888_a_:155) described adults of _atrata_. Sanborn +(1931:21) examined the one additional specimen known to me and reported +that it agreed with Thomas' description. + +Andersen (1909_b_:266) used a subadult female ("nearly fully grown") as +the holotype of _anceps_. At least five additional specimens, all +adults, of _anceps_ now are housed in various collections. Judging from +these individuals, the holotype of _anceps_ was only four-fifths grown +and because he used an immature individual, Andersen's (1912:437) +criteria for distinguishing the two subspecies mostly are invalid. + + [Illustration: FIG. 4. Distribution of _Pteralopex atrata_; + _P. atrata atrata_ ([RW]) and _P. atrata anceps_ ([BC]). + For names of islands see Fig. 2.] + + +Key to Subspecies of _Pteralopex atrata_ + + 1. Length of forearm 139-144 mm.; dorsal surface of distal + one-fourth of tibia and entire metatarsus naked; known only + from Guadalcanal and Santa Ysabel islands + _Pteralopex atrata atrata_ + + 1'. Length of forearm 162-166 mm.; dorsal surface of distal + one-fourth of tibia and entire metatarsus furred; known only + from Bougainville and Choiseul islands + _Pteralopex atrata anceps_ + + +=Pteralopex atrata atrata= Thomas + + 1888. _Pteralopex atrata_ Thomas, Ann. Mag. Nat. Hist., ser. 6, + 1:155, February, type from Guadalcanal; 1888, Thomas, Proc. Zool. + Soc. London, p. 475, December 4; 1896, Heude, Mém. Hist. Nat. + Emp. China, 3:179; 1897, Trouessart, Catalogus Mammalium ..., + 1:83; 1907, Miller, Bull. U. S. Nat. Mus., 57:60, June 29; 1912, + Andersen, Catalogue of the Chiroptera ... British Museum, 1:439; + 1931, Sanborn, Publ. Field Mus. Nat. Hist., Zool. Ser., 18:21, + February 12, from Santa Ysabel. + + 1954. _Pteralopex atrata atrata_, Laurie and Hill, List of land + mammals of New Guinea, Celebes and adjacent islands, p. 40, + June 30. + + 1899. _Pteropus (Pteralopex) atrata_, Matschie, Die Megachiroptera + ... naturkunde, p. 11; 1904, Trouessart, Catalogus Mammalium ..., + Suppl., p. 49. + +_Specimens examined._--None. + +_Remarks._--_Pteralopex atrata atrata_ is known from four specimens from +Guadalcanal and one from Santa Ysabel (Sanborn, 1931:21). + +Sanborn (_loc. cit._) reported that a specimen wounded at night, while +feeding on young green coconuts, was the only fruit bat that attempted +to attack the collectors. Troughton (1936:348) has suggested, on the +basis of his experiences with _Pteropus_, that this behavior probably +was a reaction from fear rather than an indication of general +aggressiveness on the part of _Pteralopex_. + + +=Pteralopex atrata anceps= Andersen + + 1909. _Pteralopex anceps_ Andersen, Ann. Mag. Nat. Hist., ser. 8, + 3:266, March, type from Bougainville; 1912, Andersen, Catalogue + of the Chiroptera ... British Museum, 1:437; 1936, Troughton, + Rec. Australian Mus., 14:348, April 7; 1953, Pohle, + Z. Säugetierk., 17:129, October 27. + + 1954. _Pteralopex atrata anceps_, Laurie and Hill, List of land + mammals of New Guinea, Celebes and adjacent islands, p. 40, + June 30. + +_Specimens examined_ (three males, two females; one skull-only and one +in alcohol).--Choiseul in March, 23682; Bougainville in July, USNM +276973-74, USNM 276928, USNM 277112. + +_Measurements._--Measurements of three males and one female are, +respectively, as follows: Length of head and body, 280, 271, 261, 255; +hind foot, 50, 54, 52, 59; ear, 23, 23, 26, 22; length of forearm, 160, +162, 166, 171; greatest length of skull, 77.6, 77.9, 78.9, 77.0; +condylobasal length, 74.3, 74.3, 75.5, 73.8; zygomatic breadth, 42.2, +45.4, 43.1, 42.6; breadth across upper canines, 18.7, 21.1, 19.0, 19.0; +breadth across first upper molars, 22.2, 25.3, 22.9, 22.0; length of +maxillary tooth-row, 29.3, 29.8, 28.9, 28.2; length of mandibular +tooth-row, 32.8, 32.8, 32.1, 31.4. + +_Remarks._--Heretofore, _Pteralopex atrata anceps_ was not known from +Choiseul. The specimen from that island agrees well with specimens in +the U. S. National Museum from Cape Torokina, Bougainville. + +The type specimen of this subspecies is a subadult and is smaller than +the specimens examined by me; Andersen (1912:440) gave length of forearm +of the type as 137 (as opposed to 164 in adults). He (1912:438) figured +the dentition of _anceps_ and described the ways in which it differed +from the dentition of _atrata_. Although he (1912:437) concluded that +_anceps_ and _atrata_ represented "two stages of specialization of ... +dentition," there apparently are few, if any, dental differences between +the two subspecies. Teeth of adults of _anceps_ differ from teeth of the +immature type of _anceps_ as follows: in adults the anterior basal ledge +of P4 extends onto the labial surface, whereas in the type it does not; +and maxillary and mandibular teeth in adults are spaced as in the +subspecies _atrata_ (see Andersen, 1912:438, fig. 22) and not crowded as +in the type of _anceps_. Distance between individual cheek-teeth +apparently increases with growth of the cranium and mandible. + +Adults of _P. a. anceps_ that I examined are darker than the subadult +type. The mantle in these adults is black, whereas it is seal-brown in +the type (Andersen, 1912:439). + +An adult female was lactating when obtained on Bougainville in July +(USNM 276928). + + +Key to _Pteropus_ in the Solomon Islands + + 1. Premolars having distinct basal ledges; + molars 2.5-4.0 wide 2 + + 1'. Premolars lacking definite basal ledges; + molars 1.0-2.4 wide 14 + + 2(1). Rostrum unshortened (orbit to anterior tip of nasals + about one-third greatest length of skull); dorsal + surface of tibiae nearly naked 3 + + 2'. Rostrum shortened (orbit to anterior tip of nasals less + than one-third greatest length of skull); dorsal surface + of tibiae usually at least partially furred 8 + + 3(2). Forearm more than 128 4 + + 3'. Forearm less than 128 5 + + 4(3). Forearm about 155; venter and dorsum nearly black, + mantle pale yellow =P. tonganus geddiei=, p. 798 + + 4'. Forearm 128-136; venter and dorsum near Mars Brown, + mantle Ochraceous or Cream-Buff + =P. hypomelanus luteus=, p. 796 + + 5(3'). Mantle dark, russet or cinnamon, not strongly + contrasting with color of back + =P. admiralitatum solomonis=, p. 796 + + 5'. Mantle pale, Ochraceous-Buff or Cream-Buff, strongly + contrasting with color of back 6 + + 6(5'). Mantle Ochraceous-Orange to Ochraceous-Buff, hairs pale + basally; forearm 108-111 =P. admiralitatum goweri=, p. 797 + + 6'. Mantle Ochraceous to Cream-Buff, but hairs dark brown + basally; forearm 110-122 7 + + 7(6'). Length of forearm 110-112 + =P. admiralitatum colonus=, p. 796 + + 7'. Length of forearm about 122 =P. howensis=, p. 797 + + 8(2'). Forearm more than 145 9 + + 8'. Forearm less than 144 12 + + 9(8). Forearm more than 162 10 + + 9'. Forearm less than 162 11 + + 10(9). Forearm 167-173 =P. rayneri grandis=, p. 801 + + 10'. Forearm about 164 =P. rayneri rubianus=, p. 802 + + 11(9'). Flanks and lower belly brightly colored, Burnt Sienna + to Sanford's Brown; forearm less than 150 + =P. rayneri monoensis=, p. 803 + + 11'. Flanks and lower belly darker, near tawny; forearm + more than 150, =P. rayneri lavellanus=, p. 802 + + 12(8'). Pelage of dorsum tricolored; rump brightly colored; + forearm 139-141, =P. rayneri rayneri=, p. 800 + + 12'. Pelage of dorsum bicolored; rump dark; forearm + less than 135 13 + + 13(12'). Mantle tawny with some Ochraceous-Buff; forearm + about 130, =P. rayneri rennelli=, p. 804 + + 13'. Mantle russet, lacking Ochraceous-Buff; forearm + about 121, =P. rayneri cognatus=, p. 803 + + 14(1'). Forearm more than 131; dorsum Tawny Olive + =P. mahaganus=, p. 806 + + 14'. Forearm less than 100; dorsum dark brown + =P. woodfordi=, p. 804 + + +=Pteropus= Brisson + + 1762. _Pteropus_ Brisson, Regnum animale ..., ed. 2, p. 153. + +_Remarks._--More species (seven) and subspecies (12) of _Pteropus_ occur +in the Solomon Islands than of any other chiropteran genus. Other kinds +of _Pteropus_, as yet unknown, may live there. + +The relationships among the species of these large fruit-eating bats, +commonly termed "flying foxes," are obscure and the genus is in need of +revision. The basic, definitive work is still that of Andersen (1912). +Tate (1942) and Felten (1964_a_, 1964_b_) have offered some additional +remarks but groupings and suggested relationships of species of +_Pteropus_ almost entirely are the products of Kund Andersen. According +to present-day concepts of variation and speciation, Andersen's criteria +are artificial. + +Basically, there are three "species-groups" of _Pteropus_ in the Solomon +Islands. The first is composed of species in which the rostrum is +"unshortened" (its length about one third of greatest length of skull), +and the cheek-teeth are of moderate size (M1 is 2.8-3.2 wide). The +species are _P. hypomelanus_, _P. admiralitatum_, _P. tonganus_, and _P. +howensis_. The first and second species were placed in the _Pteropus +hypomelanus_ group by Andersen (1912:98). + +In the second group the rostrum is "shortened" (its length less than one +third of greatest length of skull) and the cheek-teeth are of moderate +to large size (M1 3.3-4.1 wide). _Pteropus rayneri_, endemic to the +Solomons and represented there by at least seven subspecies, fits into +this category. + +The third group is represented by _P. mahaganus_ and _P. woodfordi_. +Both species are endemic to the Solomon Islands. In these species the +rostrum is unshortened but the cheek-teeth are greatly reduced, +especially in width (M1 is 1.0-2.2 wide). Both _P. mahaganus_ and _P. +woodfordi_ can be included in the _Pteropus scapulatus_ group of +Andersen (1912:402). + + +=Pteropus hypomelanus= + +_Pteropus hypomelanus_ is a wide-ranging species of flying fox having at +least seven subspecies; three occur in southeastern Asia, two on and +near Celebes, and two in New Guinea and islands adjacent to the +southeastern coast of New Guinea, including one island in the Solomons +(Ellerman and Morrison-Scott, 1966:95; Laurie and Hill, 1954:32-33). + + [Illustration: FIG. 5. Distribution of _Pteropus hypomelanus luteus_ + ([TW]), _Pteropus admiralitatum solomonis_ ([RW]), _Pteropus a. + colonus_ ([BW]), _Pteropus a. goweri_ ([LW]), _Pteropus tonganus + geddiei_ ([RTW]), and _Pteropus howensis_ ([BC]). For names of + islands see Fig. 2.] + + +=Pteropus hypomelanus luteus= Andersen + + 1908. Pteropus hypomelanus luteus Andersen, Ann. Mag. Nat. Hist., + ser. 8, 2:362, October, type from Kiriwini Island, Trobriand + Islands; 1912, Andersen, Catalogue of the Chiroptera ... British + Museum, 1:128; 1947, Sanborn and Beecher, Jour. Mamm., 28:388, + November 19, from Banika Island, Russell Islands. + +_Specimens examined._--None. + +_Remarks._--Andersen (1908:362) identified specimens of _Pteropus +hypomelanus_ from eastern New Guinea and three nearby islands (Conflict +Islands, Trobriand Islands, and Woodlark Island) as _P. hypomelanus +luteus_. Sanborn and Beecher (1947:388) identified a female from Banika +Island in the Solomons as of this subspecies although this specimen was +darker and had a slightly smaller skull than typical _P. hypomelanus +luteus_. They noted that the pelage of the venter of the female was +uniformly dark rather than the typical Ochraceous-Buff to Cream-Buff; +the specimen was regarded as a dark phase of the subspecies. Although +not recorded previously for _luteus_, other subspecies of _P. +hypomelanus_ were known in dark phase as well as pale and intermediate +phases of coloration (Andersen, 1912:122). The reported occurrence of +_P. h. luteus_ on Banika Island extended the known geographic range +about 450 miles eastward from Woodlark Island. + + +=Pteropus admiralitatum= + +Three subspecies, all about the same size but differing in coloration, +have been described from the Solomon Islands. _P. a. goweri_ is known +only from Gower (Ndai) Island, notably removed from the western chain of +islands inhabited by _P. a. colonus_ and _P. a. solomonis_. Only one +other subspecies, from the Admiralty Islands, is known. + + +=Pteropus admiralitatum solomonis= Thomas + + 1904. _Pteropus solomonis_ Thomas, Novit. Zool., 11:597, type from + Ghizo Island; 1912, Andersen, Catalogue of the Chiroptera ... + British Museum, 1:149; 1931, Sanborn, Publ. Field Mus. Nat. Hist., + Zool. Ser., 18:12, February 12, from Ronongo (Ganongga), Vella + Lavella, and Narovo (Simbo) islands; 1947, Sanborn and Beecher, + Jour. Mamm., 28:389, November 19, from Banika and Guadalcanal + islands. + + 1954. _Pteropus admiralitatum solomonis_, Laurie and Hill, List of + land mammals of New Guinea, Celebes and adjacent islands, p. 33, + June 30. + +_Specimens examined._--None. + +_Remarks._--Andersen (1912:149) considered _Pteropus admiralitatum_, and +especially the subspecies _P. a. solomonis_, to be the easternmost +"representative" of _Pteropus hypomelanus_. In comparison with _P. +hypomelanus luteus_, _P. a. solomonis_ differs mostly in size, being +much smaller (length of forearm about 110 rather than 134). It is now +known that both species occur on Banika Island in the Solomons. + +The subspecies _P. a. solomonis_ has been recorded from a "chain" of +islands that included Vella Lavella, Simbo, Ghizo, Ganongga, Banika, and +Guadalcanal (see Fig. 5). + + +=Pteropus admiralitatum colonus= Andersen + + 1908. _Pteropus colonus_ Andersen, Ann. Mag. Nat. Hist., ser. 8, + 2:363, October, type from Shortland Island; 1912, Andersen, + Catalogue of the Chiroptera ... British Museum, 1:150; 1931, + Sanborn, Publ. Field Mus. Nat. Hist., Zool. Ser., 18:12, + February 12, from Mono Island. + + 1954. _Pteropus admiralitatum colonus_, Laurie and Hill, List of + land mammals of New Guinea, Celebes and adjacent islands, p. 33, + June 30. + + 1887. _Pteropus hypomelanus_ (part), Thomas, Proc. Zool. Soc. + London, p. 471, December 4; 1898, Trouessart, Catalogus Mammalium + ..., 1:82, from "I. Salomonis." + + 1899. _Pteropus (Spectrum) hypomelanus_ (part), Matschie, Die + Megachiroptera ... naturkunde, p. 24. + +_Specimens examined._--None. + +_Remarks._--_Pteropus admiralitatum colonus_ is the largest of the three +subspecies that occur in the Solomon Islands. It closely resembles _P. +hypomelanus luteus_, except in being smaller throughout (see Andersen, +1912:151-152, for measurements) and darker on the underparts. + +This bat has been found in a group of small islands (Alu, Mono, and +Shortland) about 30 miles south of Bougainville. Because of this +proximity and because yet another subspecies of this species occurs +northward of Bougainville, it is interesting that neither Troughton +(1936) nor Pohle (1953) included the species in their faunal lists for +Bougainville. + +Andersen (1912:152) indicated that the M1 in _P. admiralitatum colonus_ +is smaller than in _P. a. solomonis_, the subspecies found in islands to +the southeast (4.4-4.5 and 5.2, respectively), but Sanborn (1931:13) +studied specimens of these two subspecies that overlapped in size of M1. + + +=Pteropus admiralitatum goweri= Tate + + 1934. _Pteropus goweri_ Tate, Amer. Mus. Novit., 718:1, May 4, type + from Gower (Ndai) Island. + + 1954. _Pteropus admiralitatum goweri_, Laurie and Hill, List of + land mammals of New Guinea, Celebes and adjacent islands, p. 33, + June 30. + +_Specimens examined._--None. + +_Remarks._--_Pteropus admiralitatum goweri_ was described from six +specimens collected in 1930 by the Whitney South Sea Expedition (Tate, +1934:1). This subspecies closely resembles the other two subspecies of +_P. admiralitatum_ (_colonus_ and _solomonis_) found in the Solomon +Islands. Color and length of forearm (see key on p. 793) seem to be the +only reliable criteria for distinguishing between these subspecies. The +longitude of Gower Island, 160° 34' E, was incorrectly listed in Laurie +and Hill (1954:152) as 159° 34' E. + + +=Pteropus howensis= Troughton + + 1931. _Pteropus howensis_ Troughton, Proc. Linn. Soc. New South + Wales, 56:204, June 24, type from Lord Howe Islands (Ontong Java); + 1950, Sanborn and Nicholson, Fieldiana:Zool., 31:329, August 31. + +_Specimens examined_ (one male, three females, and two sex unknown; two +embryos in alcohol).--Liuniuwu, Lord Howe Islands (Ontong Java) in +August, USNM 278703-6, USNM 279715-6. + +_Measurements._--Average and extreme measurements of one male and three +females are as follows: Length of head and body, 185.2 (176-196); hind +foot, 34.5 (33-36); ear, 21.5 (21-23); forearm not measured [broken in +all specimens examined]. Cranial measurements of a male and a female +are, respectively, as follows: Greatest length of skull, 55.3, 53.8; +condylobasal length, 54.2, 52.8; palatal length, 26.7, 26.0; zygomatic +breadth, 30.6, 29.9; breadth of braincase, 19.9, 19.2; breadth across +first upper molars, 14.3, 14.3; length of maxillary tooth-row, 20.7, +19.6; length of mandibular tooth-row, 23.1, --. + +_Remarks._--Apparently _Pteropus howensis_ is confined to Ontong Java +(Lord Howe Islands) located northeastward of the main body of islands +that constitute the Solomon Archipelago (see Fig. 5). According to A. J. +Nicholson, who collected the specimens listed above, _P. howensis_ is +not abundant in Ontong Java. He related this circumstance to the fact +that these small islands are nothing more than parts of a coral atoll +used almost entirely for the production of coconuts (see Sanborn and +Nicholson, 1950:329). + +Specimens of _Pteropus howensis_ deposited in the U. S. National Museum +agree well in most ways with the original description of the species by +Troughton (1931:204-205). Slight variation in color is evident; in two +specimens, the mantle, just posterior to the ears, is Ochraceous-Buff. + +The relationship of this species to other kinds of _Pteropus_ known from +Melanesia is not clear. Troughton (1931:204, 206) compared _P. howensis_ +with _P. hypomelanus_ and _P. admiralitatum_ and found that it resembled +each of them. Tate (1934:2) noted that the skull of _P. admiralitatum +goweri_ was similar to that of _P. howensis_ in structure. The latter +species is, however, larger (length of forearm 122 according to +Troughton, 1931:205) than any subspecies of _P. admiralitatum_ (length +of forearm 108-112). Also, the cheek-teeth of _P. howensis_ that I have +studied are relatively larger than those of either _P. hypomelanus_ or +_P. admiralitatum_. Furthermore, in _P. howensis_ there is a small but +distinct cusp located medio-posteriorly on P4 (most noticeable in young +individuals) that is more reduced or undeveloped in specimens of the +other two species. Cheek-teeth of _P. howensis_ resemble those in a +dull-colored specimen of _P. tonganus_ from Fiji Island with which I +compared the specimens listed above. + +Weights and crown-rump lengths of the two embryos (in an advanced stage +of development) examined were 20 and 29 grams and 43 and 51 mm. +(apparently these are the specimens listed by Sanborn and Nicholson, +1950:329). + + +=Pteropus tonganus= + +_Pteropus tonganus_ has at least three subspecies, one of which has been +recorded from the Solomons. The species ranges from a small island off +the eastern coast of New Guinea, where there is an endemic subspecies, +eastward to Tonga and the New Hebrides (Laurie and Hill, 1954:33-34). +Felten (1964_a_) recently has reported on the species in the New +Hebrides. + + +=Pteropus tonganus geddiei= MacGillivary + + 1860. _Pteropus geddiei_ MacGillivary, Zoologist, 18:7134, + September, type from Aneitum Island, New Hebrides; 1912, + Andersen, Catalogue of the Chiroptera ... British Museum, + 1:189; 1931, Sanborn, Publ. Field Mus. Nat. Hist., Zool. + Ser., 18:13, February 12, from Rennell Island in the Solomons. + + 1914. _Pteropus tonganus geddiei_, Revilliod, _in_ Sarasin and + Roux, Nova Caledonia (A), 1:341; 1954, Laurie and Hill, List + of land mammals of New Guinea, Celebes and adjacent islands, + p. 34, June 30. + +_Specimens examined._--None. + +_Remarks._--_Pteropus tonganus geddiei_, as far as is known, is the +widest ranging subspecies of this genus. It is the only megachiropteran +in the Solomon Islands having affinities with bats to the southeast (the +New Hebrides, Santa Cruz Islands, Samoan Islands and Fiji Islands) +rather than with those to the west (New Guinea). The subspecies _P. +tonganus geddiei_, which ranges from the Solomons to the New Hebrides +(about 500 miles straight-line distance), is said to be remarkably +uniform throughout its range. Sanborn (1931:14) compared color and size +in specimens from the Solomon Islands and the New Hebrides and found +little variation. Another subspecies, _P. t. bascilicus_ Thomas 1915, +apparently closely related to _geddiei_, is known from Dampier [= +Kar-kar] Island off the northeastern coast of New Guinea and therefore +farther westward from the New Hebrides than are the Solomon Islands. +Additional remarks on the distribution of this species are in the +section on Zoogeography and Speciation. + + +=Pteropus rayneri= + +_Pteropus rayneri_ is endemic to the Solomon Islands. It is divisible +into seven subspecies (see Fig. 6), which, excepting _P. r. rennelli_ +and _P. r. cognatus_, are strikingly colored--the mantle, back, and rump +being of different colors. Differences in color and size provide +characters differentiating the subspecies (see key, p. 793). Recorded +lengths of forearms do not overlap between any two subspecies. _P. r. +grandis_, northernmost in distribution, has the longest (about 170) +forearm and _P. r. cognatus_, known from two of the southernmost +islands, has the shortest (about 121). + + [Illustration: FIG. 6. Distribution of _Pteropus rayneri_: _P. r. + rayneri_ ([RTW]); _P. r. grandis_ ([RW]); _P. r. lavellanus_ ([BC]); + _P. r. monoensis_ ([BW]); _P. r. rubianus_ ([TW]); _P. r. cognatus_ + ([LW]); _P. r. rennelli_ ([LTW]). For names of islands see Fig. 2.] + +Adult males of _Pteropus rayneri_ have well-developed tufts of hair on +each side of the neck where a gland is located (see Andersen, +1912:259). Apparently these glands are not present in females as none +were found in specimens studied by me or those reported by Sanborn +(1931:16). Evidently, these glands are associated with sexual maturity +in males because neither Sanborn nor I found them in subadult males. + + +=Pteropus rayneri rayneri= Gray + + 1870. _Pteropus rayneri_ (part), Gray, Catalogue of monkeys, lemurs + and fruit-eating bats ... British Museum, p. 108, cotypes from + Guadalcanal; 1878, Dobson, Catalogue of the Chiroptera ... + British Museum, p. 33; 1879, Trouessart, Rev. Mag. Zool., 6:204; + 1879, Trouessart, Ann. Sci. Nat. Zool, 8:16; 1887, Thomas, Proc. + Zool. Soc. London, p. 322, March 15; 1888, Thomas, Proc. Zool. + Soc. London, p. 472, December 4; 1898, Trouessart, Catalogus + Mammalium ..., 1:78; 1912, Andersen, Catalogue of the Chiroptera + ... British Museum, p. 254; 1931, Sanborn, Publ. Field Mus. Nat. + Hist., Zool. Ser., 18:15, February 12, from Guadalcanal + and Malaita. + + 1954. _Pteropus rayneri rayneri_, Laurie and Hill, List of land + mammals of New Guinea, Celebes and adjacent islands, p. 35, + June 30. + + 1899. _Pteropus (Spectrum) rayneri_ (part), Matschie, Die + Megachiroptera ... naturkunde, p. 22; 1904, Trouessart, + Catalogus Mammalium ..., Suppl., p. 51. + +_Specimens examined_ (four males and one female; one embryo in +alcohol).--Guadalcanal in July and November, USNM 278700-02, USNM +278142, USNM 278714. + +_Measurements._--Measurements of three males and one female are, +respectively, as follows: Length of head and body, --, 210, 214, 215; +hind foot, --, 33, 39, 42; ear, --, 23, 23, 23; length of forearm, --, +138, 136, 134; greatest length of skull, 61.5, 59.2, 61.6, 61.2; +condylobasal length, 61.4, 58.2, 60.3, 60.0; zygomatic breadth, 36.6, +35.3, 35.4, 36.5; breadth of braincase, 23.7, 22.5, 22.6, 24.1; breadth +across first upper molars, --, 16.9, 16.7, 16.8; width of M1, 3.4, 3.5, +3.5, 3.5; length of maxillary tooth-row, 22.4, 22.1, 23.6, 23.2; length +of mandibular tooth-row, 26.4, 25.5, 25.9, 25.6. + +_Remarks._--_Pteropus rayneri_ was named on the basis of two specimens +(cotypes) obtained on Guadalcanal and listed as "male" and "female"; +according to Andersen (1912:254), however, both are females. + +_P. r. rayneri_ is known from Guadalcanal and Malaita (see Fig. 6), and +is of almost the same size as _P. r. cognatus_, which is known from San +Cristobal and Ugi, only about 40 miles to the southeast. In the latter +subspecies the back and rump are the same color (Prouts Brown), whereas +in _P. r. rayneri_ the rump is brightly colored and therefore contrasts +strongly with the dark brown back. A specimen of _rayneri_ from Malaita +was reported by Sanborn (1931:15) as unusually small and having a +dark-colored rump patch. In the specimens examined from Guadalcanal, +there is noticeable variation in color of the mantle that does not seem +related to age or sex. In two specimens (adult male and female) the +mantle is Cinnamon-Rufous tinged with Russet, strongly contrasting with +the crown, which is Ochraceous-Tawny and has scattered silvery hairs. +Another specimen has a darker mantle (near Chestnut-Brown) and a crown +of about the same color, but with a few scattered Ochraceous-Tawny +hairs. + +The skull of one adult male bears an extra peglike tooth posterior to M3 +on the right side. + +An embryo, in an advanced stage of development, in the collection of the +U. S. National Museum, measures: Length of head and body, 98; hind foot, +30; ear, 8.5; length of forearm, 48 (this may be the same specimen +listed by Sanborn and Nicholson, 1950:329). + + +=Pteropus rayneri grandis= Thomas + + 1887. _Pteropus grandis_ Thomas, Ann. Mag. Nat. Hist., ser. 5, + 19:147, March, type from Shortland; 1887, Thomas, Proc. Zool. + Soc. London, p. 320, March 15, from Alu and Shortland; 1897, + Trouessart, Catalogus Mammalium ..., 1:80, from "I. Salomonis"; + 1899, Matschie, Die Megachiroptera ... naturkunde, p. 15; 1904, + Trouessart, Catalogus Mammalium ..., Suppl., p. 49; 1907, Miller, + Bull. U. S. Nat. Mus., 57:58, June 29; 1912, Andersen, Catalogue + of the Chiroptera ... British Museum, 1:259, from Bougainville; + 1931, Sanborn, Publ. Field Mus. Nat. Hist., Zool. Ser., 18:16, + February 12, from Choiseul, and Santa Ysabel; 1936, Troughton, + Rec. Australian Mus., 19:348, April 7; 1953, Pohle, Z. + Säugetierk., 17:128, October 27. + + 1954. _Pteropus rayneri grandis_, Laurie and Hill, List of land + mammals of New Guinea, Celebes and adjacent islands, p. 35, + June 30. + +_Specimens examined_ (six males and 10 females; five in +alcohol).--Choiseul in March, 23580, 23644, 23593; Bougainville in July, +August, September, and October, USNM 276926-7, USNM 276968, USNM +277091-9. + +_Measurements._--Average and extreme measurements of four males and +seven females are as follows: Length of head and body, 281 (260-302); +hind foot, 52.3 (50-58); ear, 33.1 (31-37); length of forearm, 173 +(168-180). Average and extreme measurements of skulls of three males and +six females are as follows: Greatest length of skull, 73.7 (71.3-77.7); +condylobasal length, 73.1 (70.5-77.4); zygomatic breadth, 40 +(36.4-41.5); breadth across first-upper molars, 20.9 (18.3-22.1); length +of maxillary tooth-row, 28.1 (26.9-29.9); length of mandibular +tooth-row, 31.8 (29.7-32.7). + +_Remarks._--_Pteropus rayneri grandis_ is the largest subspecies of the +species. It is also the widest ranging subspecies, being found on six +islands (see Fig. 6). + +Although the specimens listed above agree well with descriptions of +color given by Thomas (1887_a_:147) and Andersen (1912:259, 263-264), +some individual variation is noticeable. In bats not yet fully grown +(judging from small size, unfused epiphyses, and lack of wear on teeth), +numerous scattered hairs on the sides of the face and crown are buffy. +In adults the face and crown are blackish. With regard to individual +variation in color of mantle and rump patch, specimens with the +following combinations were noted (1) mantle Brick Red, rump patch +bright, basal three-quarters of hairs white, tips Warm Buff (2) mantle +darker, near Hessian Brown, rump patch dark, Chestnut along edges, +center Ochraceous-Tawny (3) mantle Brick Red, rump patch intermediate +between the two other types. Size of rump patch also is variable. In +some specimens it extends onto the upper parts of the thighs whereas in +other specimens it does not. + +Sanborn (1931:16) reported an extra tooth, behind the last lower molar, +in a specimen from Choiseul. In one of three specimens in the Bishop +Museum, m3 is lacking. Judging from Troughton's (1936:346) remarks, size +of individuals varies considerably. Specimens that he examined from +Bougainville had longer forearms (up to 177) and larger hind feet +(54-57) than those examined by me from Choiseul. On the other hand, +specimens listed above from Bougainville agree well with those from +Choiseul. In many specimens in the U. S. National Museum, length of the +right- and left-forearm differ. For example, in No. 276926 the right +forearm measures 180 whereas the left is 174; in No. 277098 the right is +172 and the left is 167. Troughton (1936:346) gave standard ear +measurement in _P. r. grandis_ as ranging from 29.5 to 31.5. Ears of +specimens that I examined varied from 31.0 to 37.0. + + +=Pteropus rayneri rubianus= Andersen + + 1908. _Pteropus rubianus_ Andersen, Ann. Mag. Nat. Hist., ser. 8, + 2:366, October, type from Rubiana; 1912, Andersen, Catalogue + of the Chiroptera ... British Museum, 1:255; 1931, Sanborn, + Publ. Field Mus. Nat. Hist., Zool. Ser., 18:15, February 12, + from Narovo (Simbo). + + 1954. _Pteropus rayneri rubianus_, Laurie and Hill, List of land + mammals of New Guinea, Celebes and adjacent islands, p. 35, + June 30. + + 1888. _Pteropus grandis_ (part), Thomas, Proc. Zool. Soc. London, + p. 470, December 4, from Rubiana; 1899, Matschie, Die + Megachiroptera ... naturkunde, p. 15; 1904, Trouessart, + Catalogus Mammalium ..., Suppl., p. 49. + +_Specimens examined_ (two males and one female).--Kolombangara, in +February, 23458-60. + +_Measurements._--Measurements of two males and one female are, +respectively, as follows: Length of head and body, 253, 265, 251; hind +foot, 53, 50, 50; ear, 30, 31, 32; length of forearm, 158, 161, 160; +greatest length of skull, 70.2, 67.4, --; condylobasal length, 67.0, --, +68.4; zygomatic breadth, 40.0, 39.4, 40.7; breadth across first upper +molars, 19.4, 20.4, 19.9; length of mandible, 53.9, 49.4, 51.3. + +_Remarks._--Kolombangara Island is a new locality for _Pteropus rayneri +rubianus_; heretofore this subspecies was known only from Rubiana and +Narovo islands (Andersen, 1908:366; Sanborn, 1931:15). The coloration of +a specimen from Narovo Island was described as between that of _P. r. +rubianus_ and _P. r. lavellanus_. Sanborn (1931:16) allocated it to the +subspecies _rubianus_ on the basis of length of forearm. + +Andersen's descriptions (1908:366; 1912:256) of _rubianus_ were of a +specimen stored in alcohol. Coloration of the museum skins examined by +me is as follows: Dorsum from shoulders to rump near Vandyke Brown; +crown and mantle Brick Red; face close to Mummy Brown; rump patch and +thighs close to Warm Buff, strongly contrasting with back and mantle; +base of hairs dark, Seal Brown; venter dark; chest about same as back +but paler laterally (to Ochraceous Tawny); throat Brick Red. + + +=Pteropus rayneri lavellanus= Andersen + + 1908. _Pteropus lavellanus_ Andersen, Ann. Mag. Nat. Hist., ser. 8, + 2:366, October, type from Vella Lavella; 1912, Andersen, + Catalogue of the Chiroptera ... British Museum, 1:259; 1931, + Sanborn, Publ. Field Mus. Nat. Hist., Zool. Ser., 18:16, + February 12, from Ghizo and Ronongo. + + 1954. _Pteropus rayneri lavellanus_, Laurie and Hill, List of land + mammals of New Guinea, Celebes and adjacent islands, p. 36, + June 30. + +_Specimens examined_ (one male and one female).--Vella Lavella in +November, 23192, 23142. + +_Measurements._--Measurements of a male and a female are, respectively, +as follows: Length of head and body, 286, 282; hind foot, 55, 56; ear, +30, 30; length of forearm, 156, 155; greatest length of skull, 72.9, +67.6; condylobasal length, 71.8, 64.2; zygomatic breadth, 38.4, 37.9; +breadth across first upper molars, 19.9, 19.8; length of mandible, 54.6, +50.8. + +_Remarks._--_Pteropus rayneri lavellanus_ inhabits islands +geographically near those from which _P. r. rubianus_ is known (see Fig. +6) and in most respects the two subspecies closely resemble each other. +_P. r. lavellanus_ is slightly the smaller (average length of forearm +about 156 instead of 160) and darker. A bat from Narovo [Simbo] Island, +only a few miles from Vella Lavella, identified by Sanborn (1931:16) on +basis of its size as _P. r. rubianus_, resembled the subspecies +_lavellanus_ in color and probably represents an intergrade between the +two populations. + +The color of _P. r. lavellanus_ is close to that of _P. r. rubianus_ +except that the crown, mantle, and foreneck are near Chestnut-Brown, the +basal portions of hair black, and the fur of the venter, from sternum to +pectoral region, is dark, almost black (compare with description of _P. +r. rubianus_ under account of that subspecies). + +Measurements of the male examined are greater than those of the female +studied. Andersen (1912:259) noted that the canine teeth are heavier in +males than in females. + + +=Pteropus rayneri monoensis= Lawrence + + 1945. _Pteropus rayneri monoensis_ Lawrence, Proc. New England + Zool. Club, 23:63, March 26, type from Mono (Treasury); 1954, + Laurie and Hill, List of land mammals of New Guinea, Celebes + and adjacent islands, p. 36, June 30. + +_Specimens examined._--None. + +_Remarks._--_Pteropus rayneri monoensis_ is the most recently described +subspecies of _P. rayneri_. Lawrence (1945:63) judged that in most ways +this bat is intermediate between _P. r. grandis_ and _P. r. lavellanus_. +Coloration of _monoensis_ indicates affinity with the former, whereas +length of forearm (145-148) approaches that in the latter. The small +skull, narrow palate, and whitish rump patch of _monoensis_ are +differences that distinguish it from _grandis_ and _lavellanus_. The +relatively isolated position of Mono Island may have been important in +establishment of the distinctive features of this bat. + +Lawrence (1945:65) quoted a collector as stating: "They [individuals of +_P. r. monoensis_] rest quietly during the day in the tops of +heavy-leaved, tall jungle trees, and start flying about dusk, looking +for feeding spots. There is usually quite a flight for fifteen to twenty +minutes at twilight...." + +No additional specimens of this subspecies have been collected on small +adjacent islands and _monoensis_ may therefore be confined to Mono +Island. + + +=Pteropus rayneri cognatus= Andersen + + 1908. _Pteropus cognatus_ Andersen, Ann. Mag. Nat. Hist., ser. 8, + 2:365, October 1, type from San Cristobal; 1912, Andersen, + Catalogue of the Chiroptera ... British Museum, 1:251; 1931, + Sanborn, Publ. Field Mus. Nat. Hist., Zool. Ser., 18:15, + February 12, from San Cristobal and Ugi; 1954, Laurie and Hill, + List of land mammals of New Guinea, Celebes and adjacent islands, + p. 35, June 30. + + 1962. _Pteropus rayneri cognatus_, Hill, The natural history of + Rennell Island, British Solomon Islands, 4:9, February. + + 1870. _Pteropus rayneri_ (part), Gray, Catalogue of monkeys, lemurs + and fruit-eating bats ... British Museum, p. 108, from San + Cristobal; 1878, Dobson, Catalogue of the Chiroptera ... British + Museum, p. 33. + + 1904. _Pteropus_ (_Spectrum_) _rayneri_ (part), Trouessart, + Catalogus Mammalium ..., Suppl., p. 51. + +_Specimens examined._--None. + +_Remarks._--Specimens of _Pteropus rayneri cognatus_ first were reported +under the name _Pteropus rayneri_ based on three specimens (one from San +Cristobal and two from Guadalcanal). Because the description was based +mostly on the two specimens from Guadalcanal, the name _rayneri_ is +applicable to the bats from that island. Andersen (1908:365) thought +that specimens that he studied, from San Cristobal, were specifically +distinct from _P. rayneri_ and he proposed the name _Pteropus cognatus_ +for them. Later, Hill (1962:9) reduced _cognatus_ to subspecific status +under _P. rayneri_. + +Presently _P. r. cognatus_ is known only from San Cristobal and the +small adjacent island of Ugi (see Fig. 6). + + +=Pteropus rayneri rennelli= Troughton + + 1929. _Pteropus rennelli_ Troughton, Rec. Australian Mus., 17:193, + September 4, type from Rennell Island; 1954, Laurie and Hill, + List of land mammals of New Guinea, Celebes and adjacent islands, + p. 35, June 30. + + 1962. _Pteropus rayneri rennelli_, Hill, The natural history of + Rennell Island, British Solomon Islands, 4:7, February. + +_Specimens examined._--None. + +_Remarks._--Until recently, _Pteropus rayneri rennelli_ was known from +but a single specimen. Hill (1962:7) reported two additional specimens +and pointed out that _P. r. cognatus_ and _P. r. rennelli_ probably +represent the extremes of an east-west cline in size. _P. r. rennelli_ +and _P. r. cognatus_ differ from other subspecies of the species in +lacking tricolored pelage on the dorsum, but their short rostrum clearly +indicates affinity with other members of this complex group in the +Solomon Islands (Hill, 1962:8). + +The relationship of the subspecies _rennelli_ and _cognatus_ is close, +both geographically and genetically. Longer forearm, longer metacarpals, +and longer mandibular tooth-row serve to differentiate _rennelli_ from +_cognatus_. + + +=Pteropus woodfordi= Thomas + + 1888. _Pteropus woodfordi_ Thomas, Ann. Mag. Nat. Hist., ser. 6, + 1:156, February, type from Guadalcanal; 1888, Thomas, Proc. Zool. + Soc. London, p. 472, December 4; 1898, Trouessart, Catalogus + Mammalium ..., 1:78; 1907, Elliot, Field Columbian Mus., Zool. + Ser., 8:491; 1912, Andersen, Catalogue of the Chiroptera ... + British Museum, 1:410, from New Georgia and Guadalcanal; 1931, + Sanborn, Publ. Field Mus. Nat. Hist., Zool. Ser., 18:19, February + 12, from Kolombangara; 1947, Sanborn and Beecher, Jour. Mamm., + 28:389, November 19, from Banika and Guadalcanal; 1954, Laurie + and Hill, List of land mammals of New Guinea, Celebes and + adjacent islands, p. 39, June 30. + + 1899. _Pteropus (Sericonycteris) woodfordi_, Matschie, Die + Megachiroptera ... naturkunde, p. 83; 1904, Trouessart, Catalogus + Mammalium ..., Suppl., p. 54. + + 1945. _Pteropus austini_ Lawrence, Proc. New England Zool. Club, + 23:59, March 26, from Florida. + +_Specimens examined_ (four males and three females; five in alcohol and +two skin-onlys).--Fauro, in April, 23727, 23790; Guadalcanal in May and +June, 23823, 23931; Pavuvo (Russell Islands) in August and October, USNM +277887, USNM 283872-3. + +_Measurements._--External measurements of two males and two females are, +respectively, as follows: Length of head and body, 152, 128, 132, 155; +hind foot, 29, 26, 31, 28; ear, 16, 14, 14, 17; length of forearm, 79, +76, 86, 90. + +_Remarks._--Heretofore, _Pteropus woodfordi_ was known from New Georgia, +Guadalcanal, Kolombangara, and Banika (see Fig. 7); specimens from Fauro +and Pavuvo islands, listed above, provide new northern localities of +record for this species. + +Judging by small size and unfused epiphyses, a bat obtained in April and +another obtained in June are subadults. Specimens of adults, examined by +me, agree well with the descriptions of _P. woodfordi_ by Thomas +(1888_a_:156) and Andersen (1912:407-409), but are slightly smaller than +specimens listed by Sanborn and Beecher (1947:389). Color of pelage in +this species seems to vary. Adults seen have a pale head and mantle, +contrasting strongly with the dark back. Andersen (1912:409) and +Lawrence (1945:61) discussed individuals that had scattered silvery +hairs mixed with dark fur dorsally and darker mantles that did not +contrast noticeably with the rest of the dorsum. + +Lawrence (1945:389) named _Pteropus austini_ as a new species closely +related to _P. woodfordi_ and other species of the _P. scapulatus_ group +of Andersen (1912:402) and Tate (1942:336). Sanborn and Beecher +(1947:389), studied a series of _P. woodfordi_ from Banika and +Guadalcanal and found that skulls of two subadults agreed well with +cranial characteristics ascribed to _P. austini_, which was based on two +subadults. Lawrence (1945:61) stated also that "the interfemoral +membrane is entirely absent medially in _austini_, while in _woodfordi_ +it is present as a barely discernible ridge 8 mm. wide." Andersen +(1912:408) had earlier reported that in the type of _woodfordi_ the +interfemoral membrane was "undeveloped in [the] centre." In 13 adults +(in alcohol) studied by Sanborn and Beecher (1947:389), as well as in +adults examined by me, the uropatagium is not present. In size, however, +these specimens agree with dimensions given for _woodfordi_ by Thomas +(1888_a_:156) and Andersen (1912:410); for example, length of forearm is +93-99. According to Lawrence (1945:59) _austini_, in which the +interfemoral membrane is lacking, is smaller than _woodfordi_ and has a +forearm of about 84. In two juveniles of _P. woodfordi_ in the U. S. +National Museum, the medially-developed interfemoral membrane is about 7 +wide. One specimen has small but distinct calcars whereas the other +(slightly larger) apparently lacks calcars. This suggests individual +variation in the presence or absence, as well as in the size, of the +uropatagium in _Pteropus woodfordi_. + +Sanborn and Beecher (1947:389) decided that "until fully adult specimens +showing the characters of _austini_ are available, it best be considered +a synonym of _woodfordi_." For the following reasons I agree with these +authors: (1) _austini_ is known from only two specimens, both of which +are apparently subadults; (2) _austini_ is reported to have a forearm 84 +long and no interfemoral membrane, whereas _woodfordi_ has a forearm +about 96 long and an interfemoral membrane that is only slightly +developed; (3) specimens that agree in size and cranial characters with +the type of _woodfordi_ but that lack an interfemoral membrane have been +obtained; and (4) skulls of subadults of _woodfordi_ agree with the +description of skulls of _austini_. + +Sanborn (1931:19) reported that specimens of _Pteropus woodfordi_ were +obtained at night, while feeding on young green coconuts. Lawrence +(1945:62) reported that in the late afternoon a collector found +individuals of _austini_ [= _woodfordi_] in the fronds of a coconut +tree, apparently feeding on pollen shoots. Sanborn and Beecher +(1947:388) have reported malaria (_Plasmodium_) in _P. woodfordi_ +obtained on Guadalcanal. They suggested that malaria might have rendered +one individual helpless because when it was found, on the ground, no +wounds were evident and parasites were present in the blood. + + [Illustration: FIG. 7. Distribution of _Pteropus woodfordi_ ([BW]) + and _P. mahaganus_ ([BC]). For names of islands see Fig. 2.] + + +=Pteropus mahaganus= Sanborn + + 1931. _Pteropus mahaganus_ Sanborn, Publ. Field Mus. Nat. Hist., + Zool. Ser., 18:19, February 12, type from Santa Ysabel, also + reported from Bougainville; 1954, Laurie and Hill, List of + land mammals of New Guinea, Celebes and adjacent islands, + p. 39, June 30. + +_Specimens examined_ (one male and two females; one in +alcohol).--Bougainville, in August and October, USNM 276972, USNM +277104-5. + +_Measurements._--Measurements of one male and two females are, +respectively, as follows: Length of head and body, 180, 204, 198; hind +foot, 42, 38, 44; ear, 25, 23, 22; length of forearm, 134, 138, 140. +Measurements of the skull of the male and one female are, respectively, +as follows: Greatest length of skull, 52.5, 55.8; condylobasal length, +50.9, 54.3; palatal length, 24.1, 26.0; zygomatic breadth, 28.9, 32.5; +breadth across first upper molars, 14.4, 15.0; width of M1, 2.2, 2.2; +length of maxillary tooth-row, 17.4, 18.4; length of mandibular +tooth-row, 20.1, 21.4. + +_Remarks._--Sanborn (1931:19-21) described _Pteropus mahaganus_ on basis +of six specimens, five from Santa Ysabel and one from Bougainville. The +latter was in poor condition and only provisionally allocated to this +species. The specimens examined by me (listed above) confirm the +occurrence of _P. mahaganus_ on Bougainville. + +Sanborn (1931:20) described _mahaganus_ as "similar to and about the +size of [_Pteropus scapulatus_] from Australia, but lighter in color," +and considered it, along with _P. woodfordi_, a member of the _Pteropus +scapulatus_ group of Andersen (1912:402) and Tate (1942:336). I would +judge, however, that _P. mahaganus_ and _P. woodfordi_ are much more +closely related to one another than to _P. scapulatus_ of Australia. The +only significant characteristic that the latter has in common with the +two species from the Solomons is small cheek-teeth. In fact, teeth of +_scapulatus_ are relatively smaller than teeth of either _mahaganus_ or +_woodfordi_. Also, in _scapulatus_ the upper canines are widely +separated due to lateral expansion of the palate at that point, whereas +in _mahaganus_ and _woodfordi_ the width across the upper canines is +relatively much less. + + +=Dobsonia= Palmer + + 1898. _Dobsonia_ Palmer, Proc. Biol. Soc. Washington, 12:114, + April 30. + + 1810. _Cephalotes_ (part) É. Geoffroy, Ann. du Mus. d'Hist. Nat., + 15:104. + +_Dobsonia_, a genus of large to medium-sized fruit bats, occurring from +Celebes to the Solomon Islands, contains at least nine species. One +species and its two subspecies are endemic to the Solomons. + +_Dobsonia_ differs from all other genera of megachiropteran bats in the +Solomons by combining absence of a small claw on the second digit and +presence of external tail vertebrae. + +The cranium of _Dobsonia_ resembles, in some ways, the cranium of +_Rousettus_ as well as that of _Pteropus_. Even so, in _Dobsonia_ the +rostrum is shorter and the cheek-teeth, especially in the upper jaw, are +more crowded. The anterior part of the mandible is narrow and the lower +incisors are diminutive and often concealed by the flesh of the gum. + + +=Dobsonia inermis= + +In a review of the genus _Dobsonia_, Andersen (1909_c_:532) named and +described _D. inermis_ and _D. nesea_ from the Solomons. Specimens of +_Dobsonia inermis_ from San Cristobal and Ugi were said to differ from +specimens of _D. nesea_ from Alu, Shortland, and Rubiana in having +perpendicular as opposed to anteriorly slanted upper canines. Andersen +(1909_c_:532) reported that the two species were of "... the same +general size." Troughton (1936:348-349) studied specimens of _Dobsonia_ +from Bougainville and Santa Ysabel and, because of individual variation +in proclivity of the upper canines, concluded that _D. nesea_ was +conspecific with _D. inermis_. He (p. 349) noted that the ears were +shorter in _inermis_ than in _nesea_, but the size of teeth showed +insular variation and a "... confusing amount of intergradation ... +[that obscures] ... diagnostic importance." + +Specimens of _Dobsonia_ from Choiseul are smaller (externally and +cranially) than those from Alu, Shortland, Rubiana, Bougainville, Fauro, +Vella Lavella, Guadalcanal, Florida, Ugi, San Cristobal, and Rennell. +Specimens from Santa Ysabel (see Fig. 8) are intermediate in size +between those from Choiseul and the other islands listed. Judging from +available specimens, two subspecies of _Dobsonia inermis_ occur in the +Solomons. Specimens from Choiseul (see A, Fig. 8), which are smaller +than those from other islands, represent one subspecies (heretofore +unrecognized), whereas specimens from other islands (except Santa +Ysabel) represent a second subspecies. Specimens from Santa Ysabel are +slightly larger than those on Choiseul and are regarded as intergrades +between the two subspecies. Specimens from Rennell, Ugi, San Cristobal, +Florida, Fauro, and Guadalcanal are slightly smaller than those from +Bougainville, Vella Lavella, Shortland, and Rubiana, but the differences +are not great enough to warrant recognition of two subspecies. +Therefore, the subspecific name _nesea_ is arranged as a synonym of +_inermis_, which has priority, and the latter name is used for specimens +of _Dobsonia inermis_ from the Solomon islands other than Choiseul and +Santa Ysabel. Additional remarks on the distribution of this species are +in the section on Zoogeography and Speciation. + +Pohle (1953:130) suggested that _Dobsonia inermis_ (as well as _D. +crenulata_ and _D. praedatrix_) is conspecific with _D. viridis_, but +Laurie and Hill (1954:41) did not adopt his suggestion. I have not seen +adequate series of _crenulata_, _praedatrix_, and _viridis_ (none of +which occurs in the Solomons) to judge systematic relationships of these +kinds; therefore I follow Laurie and Hill. + + +=Dobsonia inermis inermis= Andersen + + 1909. _Dobsonia inermis_ Andersen, Ann. Mag. Nat. Hist., ser. 8, + 4:532, December, type from San Cristobal; 1912, Andersen, + Catalogue of the Chiroptera ... British Museum, 1:475. + + 1936. _Dobsonia inermis inermis_, Troughton, Rec. Australian Mus., + 14:349, April 7, from Santa Ysabel; 1954, Laurie and Hill, List + of land mammals of New Guinea, Celebes and adjacent islands, + p. 41, June 30; 1956, Hill, The natural history of Rennell + Island, British Solomon Islands, 1:74, November 28, from + Rennell Island. + + 1878. _Cephalotes peroni_ (part), Dobson, Catalogue of the + Chiroptera ... British Museum, p. 91; 1879, Trouessart, Rev. + Mag. Zool., 3:208; 1887, Thomas, Proc. Zool. Soc. London, + p. 323, March 15, from Ugi and San Cristobal; 1888, Thomas, + Proc. Zool. Soc. London, p. 476, December 4; 1897, Trouessart, + Catalogus Mammalium ..., 1:87. + + 1899. _Dobsonia peroni_ (part), Trouessart, Catalogus Mammalium + ..., 2:1278. + + 1909. _Dobsonia nesea_ Andersen, Ann. Mag. Nat. Hist., ser. 8, + 4:532, December 1, type from Shortland Island; 1912, Andersen, + Catalogue of the Chiroptera ... British Museum, 1:476, from + Shortland and Rubiana; 1931, Sanborn, Publ. Field Mus. Nat. + Hist., Zool. Ser., 18:22, February 12, from San Cristobal. + + 1936. _Dobsonia inermis nesea_, Troughton, Rec. Australian Mus., + 14:348, April 7, from Bougainville; 1953, Pohle, Z. Säugetierk., + 17:130, October 27; 1954, Laurie and Hill, List of land mammals + of New Guinea, Celebes and adjacent islands, p. 41, June 30, + from New Georgia. + +_Specimens examined_ (13 males and three females; three in alcohol, +crania extracted and cleaned).--Fauro in April, 23728, 23740, 23751; +Vella Lavella in November, 23134-36, 23141, 23145, 23147, 23149, 23151, +23153; Guadalcanal in May and June, 23865, 23914, 24008; Florida in +October, 24416. + +_Measurements._--See tables 2 and 3. + +_Remarks._--Heretofore, _Dobsonia inermis inermis_ was unreported from +Fauro, Vella Lavella, Guadalcanal, and Florida. Apparently the +subspecies occurs on most islands of the archipelago (see Fig. 9). + +In coloration and most cranial dimensions the specimens listed above +agree with specimens of _D. i. inermis_ from Alu, Shortland, and Rubiana +(Andersen, 1909_c_:532; 1912:475, 476), Bougainville (Troughton, +1936:348, 349), and Rennell (Hill, 1963:74). The forearm in the adult +male holotype of "_nesea_," from Shortland, is 109.5 as opposed to 109.0 +in an adult female topotype of _inermis_ from Ugi (Andersen, 1912:478) +in the southeastern part of the archipelago (see Fig. 9). Forearms of +specimens of _D. i. inermis_ from Vella Lavella are 107 to 112 +(measurements from labels because forearms of these specimens were +broken and therefore could not be remeasured). Forearms of specimens +from Fauro, Florida, Guadalcanal, and Rennell are 103.6 to 110.0 (see +Hill, 1956:74). Variation in length of forearm probably is not +significant because no cline is evident (see Fig. 9 and Table 2). + +In 1964, 11 specimens of _Dobsonia inermis_ were collected on Choiseul. +They are smaller, externally and cranially, than specimens of _D. +inermis_ from Sun Cristobal, Ugi, Rennell, Guadalcanal, Florida, +Rubiana, Vella Lavella, Shortland, Alu, Bougainville, and Fauro, and may +be named and described as follows: + + +=Dobsonia inermis minimus=, new subspecies + +_Type._--Adult male skin and skull, in good condition (originally stored +in 70 per cent alcohol for about one year), no. BBM-BSIP 23716, Bernice +P. Bishop Museum; from Choiseul Island, British Solomon Islands +Protectorate; obtained on 20 March 1964 by Philip Temple, original +number 1524. + +_Distribution._--Choiseul Island (type locality); intergrades from Santa +Ysabel also assigned to this subspecies. + +_Diagnosis._--Size small for species; wing membranes, feet, and ears +black; dorsal surface of interfemoral membrane sparsely set with silvery +hairs, other membranes naked; hair soft, medium length (10 on mantle, 5 +on crown), black hairs and scattered white hairs on face and crown; fur +of dorsal surface of mantle composed of whitish hairs having faint olive +cast imparting general color of Buffy-Citrine; hair of venter short +(about 5), soft, and fine; general coloration Buffy-Citrine; cranium +delicate; rostrum narrow in dorsal aspect (nasals not expanded +laterally); forehead (junction of nasals and frontals) pronounced in +lateral aspect; teeth resembling those of other subspecies of _D. +inermis_ but slightly smaller. + +_Comparisons._--From adults of _Dobsonia inermis inermis_, which occurs +on Rennell, San Cristobal, Ugi, Malaita, Florida, Guadalcanal, Rubiana, +Vella Lavella, Shortland, Alu, Bougainville, and Fauro, _minimus_ +differs in being smaller. Average length of mandible 31.2 and 33.4. For +other measurements see Table 2. + +From _Dobsonia praedatrix_, which occurs on New Britain, New Ireland, +and Duke of York (northward of the Solomons), _minimus_ differs in being +smaller in all dimensions; length of forearm averaging 100.5 as opposed +to 116.0, and greatest length of skull 42.4 as opposed to 50.0. + + [Illustration: FIG. 8. Greatest length of skull plotted against + zygomatic breadth for two subspecies of _Dobsonia inermis_. Symbols + represent _D. i. inermis_ ([BW]), _D. i. minimus_ ([TW]), and + intergrades assigned to _minimus_ ([BC]). Capital letters are used + to relate groups of specimens to the island or islands from which + they were collected; spatial distribution of specimens indicated in + the scatter diagram thus is shown in the inset map. Specimens from + Santa Ysabel and Bougainville are deposited in the Australian + Museum. The type specimen of _D. i. inermis_ is labeled "E." For + names of islands see Fig. 2.] + + [Illustration: FIG. 9. Distribution of _Dobsonia inermis inermis_ + ([BC]) and _D. inermis minimus_ ([RW]). For names of islands see + Fig. 2.] + + TABLE 2. Average and Extreme Measurements of Two Subspecies of + _Dobsonia inermis_. + + ================+====================================+=================== + | _D. i. minimus_ | _D. i. inermis_ + +------------------+-----------------+------------------- + | | | Guadalcanal, + MEASUREMENT | Choiseul | Santa Ysabel | Fauro, Vella + | 4 [M], 2 [F] | 1 [M], 2 [F] |Lavella, Florida + | | | 9 [M], 2 [F] + ----------------+------------------+-----------------+------------------- + Length of head | | | + and body |174.5 (170 -180 )| |174.5 (160 -190 ) + Tail vertebrae | 28.5 (24 - 33 )| 23.0 (21.5-24.0)| 30.4 ( 25 - 35 ) + Hind foot | 25.3 (25.1- 25.9)| 23.5 (23.5-24.0)| 29.3 ( 26.0- 31.6) + Ear | 21.5 (21.0- 22.9)| 21.1 (21.0-21.5)| 23.1 ( 19 - 25 ) + Length of | | | + forearm |100.5 (98.1-104.0)|105.3 (104 -107 )|108.4 (105 -112 ) + 2nd metacarpal | 43.6 (42.6- 45.2)| | 48.1 ( 45.9- 50.9) + 3rd metacarpal | 61.5 (59.8- 62.9)| | 67.6 ( 65.2- 68.5) + 4th metacarpal | 57.5 (56.5- 58.5)| | 62.5 ( 58.7- 65.5) + 5th metacarpal | 59.0 (57.0- 60.5)| | 64.4 ( 61.8- 66.0) + | | | + Greatest length | | | + of skull | 42.4 (42.1- 43.5)| 44.0 (43.0-45.6)| 45.9 ( 45.2- 47.4) + Condylobasal | | | + length | 40.4 (39.5- 41.3)| 41.7 (41.1-42.6)| 43.6 ( 43.1- 45.0) + Zygomatic | | | + breadth | 25.6 (24.9- 26.8)| 26.5 (25.7-27.6)| 27.9 ( 27.2- 28.5) + Breadth of | | | + braincase | 16.8 (16.5- 17.4)| 18.0 (17.1-19.9)| 17.9 ( 16.7- 19.0) + Breadth across | | | + upper canines | 8.4 ( 8.1- 8.7)| | 9.2 ( 9.2- 9.5) + Breadth across | | | + first upper | | | + molars | 12.1 (11.8- 12.6)| | 13.1 ( 12.6- 13.3) + Length of | | | + maxillary | | | + tooth-row | 15.6 (15.5- 15.8)| 16.4 (16.0-17.2)| 16.4 ( 15.9- 17.0) + Length of | | | + mandibular | | | + tooth-row | 17.1 (16.8- 17.6)| 17.8 (17.3-18.4)| 18.2 ( 17.8- 19.4) + ----------------+------------------+-----------------+------------------- + +_Measurements._--Comparative measurements of the subspecies _inermis_ +and _minimus_ are given in Table 2. Some measurements of the type are as +follows: Length of head and body, 147; tail vertebrae, 31; hind foot, +25; ear, 21; length of forearm, 99.5; 2nd metacarpal, 42.8; 3rd +metacarpal, 62.7; 4th metacarpal, 58.5; 5th metacarpal, 59.1; greatest +length of skull, 42.2; condylobasal length, 40.6; zygomatic breadth, +25.8; breadth of braincase, 16.8; length of maxillary tooth-row, 15.8; +length of mandible, 31.2. + +_Remarks._--_Dobsonia inermis minimus_ is the smallest subspecies of +_Dobsonia inermis_. Specimens from Santa Ysabel, southeastward of +Choiseul, are slightly larger than the type and paratypes of _minimus_. +As can be seen in the scatter diagram (Fig. 8), a male from Santa Ysabel +is as large as one male and most females of _D. i. inermis_. The other +three specimens from Santa Ysabel also are slightly larger than +specimens of _minimus_ from Choiseul, but are much smaller than +specimens of _D. i. inermis_, and, therefore, are referred to _D. i. +minimus_. + +Although there is a cline in size of _Dobsonia inermis_ from Choiseul to +Florida (generally southward; Fig. 9), no cline in size is apparent +between Choiseul and Fauro (generally westward). Specimens of _D. +inermis_ from Fauro are average for the subspecies _inermis_; there is +no evidence, in the small series available, of intergradation between +_minimus_ on Choiseul and _inermis_ on Fauro. + +_Specimens examined_ (eight males and three females, all originally in +alcohol; seven crania, all adults, extracted and cleaned).--Choiseul in +March, 23565, 23628, 23637, 23665-67, 23640, 23714, 23716 (holotype), +23717, 23720. Ellis LeG. Troughton kindly examined and measured nos. +AM-M. 3693[M], AM-M. 3694[M], AM-M. 3937[F], and AM-M. 3940[F], from +Santa Ysabel in the Australian Museum. + + +Subfamily Macroglossinae + + +=Macroglossus= F. Cuvier + + 1824. _Macroglossus_ F. Cuvier, Des dents des mammiferes ... + zoologiques, p. 248. + + 1840. _Kiodotus_ Blyth, _in_ Cuvier's animal kingdom ..., p. 69. + + 1891. _Carponycteris_ Lydekker, _in_ Flower and Lydekker, mammals + living and extinct, p. 654. + + 1902. _Odontonycteris_ Jentink, Notes Leyden Mus., 23:140, July 15. + +_Macroglossus_, the widest-ranging genus of macroglossine bats, occurs +from southeastern Asia to the southern islands of the Solomon +Archipelago (see Ellerman and Morrison-Scott, 1966:101; Laurie and Hill, +1954:44). One species, known also from Celebes and New Guinea, occurs in +the Solomons and is represented there by an endemic subspecies. + +Numerous generic names have been applied, at one time or another, to +bats now considered as _Macroglossus_. Trouessart (1904:65) and Miller +(1907:70) listed the one bat of this genus occurring in the Solomons +under _Carponycteris_ and _Kiodotus_, respectively. Andersen (1911:642; +1912:767) and, later, Sanborn (1931:22) identified this bat as +_Macroglossus lagochilus microtus_. Troughton (1936:350), reporting an +extension of range of this species in the Solomons, used the generic +name _Odontonycteris_ without explanation. Andersen (1912:754) pointed +out that Jentink originally established the name _Odontonycteris_ on the +basis of an extra premolar in each upper jaw as opposed to the usual two +in _Macroglossus_, and arranged _Odontonycteris_ as a synonym of +_Macroglossus_ because "in no genus of Megachiroptera are dental +anomalies of so frequent occurrence as in _Macroglossus_, and on no +point of the jaws are these anomalies ... so often met with as on that +occupied by the molar series." Sanborn (1931:22) and Phillips (1966:27) +noted variation in number of incisors in _Macroglossus_ as well as in +_Melonycteris_, another macroglossine genus. All of the more recent +workers (Ellerman and Morrison-Scott, 1966; Pohle, 1953; Laurie and +Hill, 1954) use the name _Macroglossus_. + + +=Macroglossus lagochilus= + +_Macroglossus lagochilus_ has at least three subspecies, one of which is +endemic to the Solomons. The species ranges from Celebes on the west to +the Solomon Islands on the east, occurring not only in New Guinea but +also on many of the small adjacent islands (see Laurie and Hill, +1954:44). + + [Illustration: FIG. 10. Distribution of _Macroglossus lagochilus + microtus_. For names of islands see Fig. 2.] + + +=Macroglossus lagochilus microtus= Andersen + + 1911. _Macroglossus lagochilus microtus_ Andersen, Ann. Mag. Nat. + Hist., Ser. 8, 7:642, June, type from Guadalcanal, additional + specimens from Florida; 1912, Andersen, Catalogue of the + Chiroptera ... British Museum, 1:767; 1931, Sanborn, Publ. + Field Mus. Nat. Hist., Zool. Ser., 18:22, February 12, from San + Cristobal; 1953, Pohle, Z. Säugetierk., 17:130, October 27, + from Bougainville; 1954, Laurie and Hill, List of land mammals + of New Guinea, Celebes and adjacent islands, p. 44, June 30. + + 1888. _Macroglossus australis_ (part). Thomas, Proc. Zool. Soc. + London, p. 476, December 4, from Guadalcanal. + + 1904. _Carponycteris nana_ (part), Trouessart, Catalogus Mammalium + ..., Suppl., p. 65. + + 1907. _Kiodotus_ sp., Miller, Bull. U. S. Nat. Mus., 57:70, + June 29. + + 1936. _Odontonycteris lagochilus microtus_, Troughton, Rec. + Australian Mus., 14:350, April 7, from Bougainville. + +_Specimens examined_ (14 males and 16 females; in alcohol).--Choiseul in +March, 23654-57, 23614, 23629, 23643, 23645, 23647, 23677-79, 23684; +Vella Lavella in December, 23277-79, 23283-84; Fauro in April, 23765; +Guadalcanal in May and June, 23830, 23864, 23935; Kolombangara in +January, 23385, 23399, 23397, 23407, 23420-21; Santa Ysabel in June, +24067; Malaita in June, 24067. + +_Measurements._--Average and extreme external measurements of 14 males +and 15 females are as follows: Length of head and body, 68.3 (63-72); +tail vertebrae present but scarcely perceptible and therefore not +measured; hind foot, 11.4 (9.0-12.9); ear, 12.0 (10.0-12.9); length of +forearm, 37.6 (36.2-39.9). + +_Remarks._--The distribution of _Macroglossus lagochilus microtus_ has +not been well known. Specimens herein reported from Choiseul, Fauro, and +Vella Lavella provide new records of distribution. As shown on Figure +10, the subspecies occurs throughout the Solomon Islands. + +_Macroglossus lagochilus microtus_ differs slightly from _M. l. nanus_ +Matschie, the subspecies of the Bismarck Archipelago and Admiralty +Islands to the north of the Solomons. _M. l. nanus_ averages slightly +larger than _microtus_ (see Andersen, 1912:768-769, for comparative +measurements) but otherwise closely resembles it. + +Individual variation is evident in several measurements of the specimens +at hand (in length of forearm, for example) but no clines are apparent. +Four females obtained in March were lactating, as was one taken in +December and one taken in January. + + +=Melonycteris= Dobson + + 1877. _Melonycteris_ Dobson, Proc. Zool. Soc. London, p. 119, + June 1. + + 1877. _Cheiropteruges_ Ramsay, Proc. Linn. Soc. New South Wales, + 2:19, July. + + 1887. _Nesonycteris_ Thomas, Ann. Mag. Nat. Hist., ser. 5, 14:147, + February. + +The genus _Melonycteris_ is known from three species, two apparently +endemic to the Solomon Islands and the third occurring in eastern New +Guinea and the Bismarck Archipelago (Laurie and Hill, 1954:45). + +Heretofore, the generic name _Nesonycteris_ has been applied to the +species in the Solomons, whereas _Melonycteris_ has been restricted to +the one species in the Bismarck Archipelago and New Guinea. Andersen +(1912:792) judged that _Nesonycteris_ was clearly distinct from +_Melonycteris_ on the basis of two characters (loss of a claw on the +second digit and loss of the inner, lower incisors). On the other hand, +he noted striking similarities in general cranial features, dentition, +palatal ridges, tongue, and external appearance of the two genera. Pohle +(1953:131) synonymized the two but Laurie and Hill (1954:45) considered +them distinct. I have suggested previously (Phillips, 1966:26, 27) that +characteristics used to distinguish between _Melonycteris_ and +_Nesonycteris_ are of less than generic value. Variability of number of +incisors in the upper jaw of specimens of _Melonycteris_ (and in other +macroglossine genera, as well) indicates a lack of selective pressure +for either increase or decrease in number of incisors. Furthermore, the +loss of the small claw on the second digit might not be important +because, as Bader and Hall (1960:15) have pointed out, limbs of bats +vary more in phenotypic expression than do other parts of the skeletal +structure. + +The discovery of a new species (_Melonycteris aurantius_) in the Solomon +Islands sheds additional light on the problem. Although _M. aurantius_ +possesses the distinguishing characteristics of the genus +"Nesonycteris," the species closely resembles _Melonycteris_ in other +features. Similarity in structure of hair of _Melonycteris_ and +_Nesonycteris_, as first reported by Benedict (1957:293), also supports +the argument for synonymy (see Phillips, 1966:26). + +_Melonycteris aurantius_ lacks a small claw on the second digit and has +only two lower incisors. In these ways this species is like _woodfordi_, +which also is restricted to the Solomons. On the other hand, the +structure of the skull of _M. aurantius_ is like that of _M. melanops_, +which is the species found in the Bismarck Archipelago. + +Although _melanops_ is not yet known from the Solomon Islands, I have +included it in the following key. + +Key to Known Species of _Melonycteris_ + + 1. Ventral surface darker than dorsum, but not strongly + contrasting with it; lacking a small claw on the + second digit, 2 + + 1´. Ventral surface nearly black, strongly contrasting with + dorsum; small claw on second digit, + =Melonycteris melanops= + + 2(1´). Pelage bright, Cinnamon-Rufous; postorbital region of + skull expanded (about 8.3 wide), + =Melonycteris aurantius=, p. 816 + + 2´. Pelage dark, near Wood-Brown or Cinnamon; postorbital + region of skull constricted (about 7.5), + =Melonycteris woodfordi=, p. 816 + + [Illustration: FIG. 11. Distribution of _Melonycteris aurantius_ + [BC] and _M. woodfordi_ [LW]. For names of islands see Fig. 2.] + + +=Melonycteris aurantius= Phillips + + 1966. _Melonycteris aurantius_ Phillips, Jour. Mamm., 47:23-27, + March 12, type from Florida Island, additional specimens from + Choiseul Island. + +_Specimens examined_ (six females; three in alcohol).--Florida in +October, 24440; Choiseul in March, 23615, 23617, 23558, 23694, 23681. + +_Measurements._--Average and extreme measurements of six females are as +follows: Length of head and body, 80.8 (77-106); hind foot, 17.2 +(16.0-18.7); ear, 12.7 (11.5-14.0); length of forearm, 49.3 (42.9-53.8). +Average and extreme measurements of skulls of five females are as +follows: Greatest length of skull, 31.8 (30.8-33.3); condylobasal +length, 29.7 (28.6-32.4); zygomatic breadth, 18 (17.2-20.0); breadth of +braincase, 12.6 (12.4-13.2); postorbital breadth, 8.3 (8.0-8.9); length +of maxillary tooth-row, 10.1 (9.4-10.4); length of mandibular tooth-row, +11.7 (10.8-12.2). + +_Remarks._--On Choiseul Island _Melonycteris aurantius_ was taken at the +same locality as its congener, _Melonycteris woodfordi_. + +Externally, _M. aurantius_ resembles _M. woodfordi_. These species are +the same size, but the former is brighter in color (nearly orange in +adults) than the latter, which is Wood-Brown dorsally. Internally, +differences between _M. aurantius_ and _M. woodfordi_ are more obvious. +In the skull of _M. aurantius_, the postorbital region is expanded +(measuring about 8.3), whereas in _M. woodfordi_ the postorbital region +is constricted. Furthermore, in lateral aspect the posterior portion of +the skull of _M. aurantius_ is down-turned and the angle of the facial +axis with the basicranial axis is much more acute than in _M. +woodfordi_. + +The number of upper incisors is highly variable in the six specimens of +_M. aurantius_ that I have examined. In two specimens an extra tooth has +erupted just anterior to I2 and there is a total of six upper incisors. +In two other specimens an extra tooth has erupted in front of I2 on one +side but not the other. I could find no trace of an extra tooth in the +remaining two specimens. + +Practically nothing is known about the natural history of _M. +aurantius_, or, indeed, that of either of the other two species of this +genus. One field collector (Temple, _in litt._) for the Bishop Museum +reported that he obtained both _M. aurantius_ and _M. woodfordi_ in the +same mist net in one night. The holotype, an adult female, was lactating +when obtained in October. + + +=Melonycteris woodfordi= (Thomas) + + 1887. _Nesonycteris woodfordi_ Thomas, Ann. Mag. Nat. Hist., ser. + 5, 14:147, February, type from Shortland Island; 1887, Thomas, + Proc. Zool. Soc. London, p. 324, March 15; 1888, Thomas, Proc. + Zool. Soc. London, p. 476, December 4; 1898, Trouessart, + Catalogus Mammalium ..., 1:90; 1899, Matschie, Die Megachiroptera + ... naturkunde, p. 91; 1904, Trouessart, Catalogus Mammalium .., + Suppl., p. 66; 1907, Miller, Bull. U. S. Nat. Mus., 57:74, + June 29; 1912, Andersen, Catalogue of the Chiroptera ... British + Museum, 1:792, from Alu, Shortland, Fauro, and Guadalcanal; 1931, + Sanborn, Publ. Field Mus. Nat. Hist., Zool. Ser., 18:23, February + 12, from Russell Island (Pavuvo); 1954, Laurie and Hill, List of + land mammals of New Guinea, Celebes and adjacent islands, p. 45, + June 30. + + 1953. _Melonycteris woodfordi_, Pohle, Z. Säugetierk., 17:130, + October 27, from Bougainville Island; 1966, Phillips, Jour. Mamm., + 47:23, March 12, from Choiseul. + +_Specimens examined_ (three males and one female; in +alcohol).--Choiseul, in April, 23413-14, 23434, 23275. + +_Measurements._--Average and extreme measurements of three males and one +female are as follows: Length of head and body, 86.1 (83.1-91.0); hind +foot, 19.6 (17.2-22.2); ear, 11.3 (10.8-11.7); length of forearm, 54.4 +(52.1-57.7). + +_Remarks._--Specimens of _Melonycteris woodfordi_ from Choiseul +constitute a new locality of occurrence for the species. Apparently _M. +woodfordi_ occurs throughout the Solomons (see Fig. 11). + +Thomas (1887_a_:147) named _Nesonycteris woodfordi_ in a preliminary +report that appeared before the publication of the more detailed +description of the genus and species (1887_b_:323-324). In the second +paper he stated that the anterior projections of the premaxillary bones +are separated distinctly in both _Nesonycteris_ and _Melonycteris_. +According to Thomas (1887_b_:323), it was by some "accident" that Dobson +(1878:4) reported the anterior projections of the premaxillary bones in +_Melonycteris melanops_ to be united. Writing at a later date, Andersen +(1912:785) reported that in _Melonycteris melanops_ the premaxillary +bones have "simple contact with each other." Furthermore, in Andersen's +(1912:791) illustration of _M. woodfordi_ the premaxillary bones are in +contact anteriorly. In specimens of _woodfordi_ and _melanops_ examined +by me, the premaxillary bones are in contact. In _M. aurantius_ the +premaxillary bones are not in contact, and it differs from _woodfordi_ +in several other respects. + +In _M. woodfordi_, as in other macroglossine bats, there is variability +in dentition. One specimen examined has a total of three upper incisors, +and another had an extra peglike tooth just anterior to I1. + + +Subfamily Nyctimeninae + + +=Nyctimene= Borkhausen + + 1797. _Nyctimene_ Borkhausen, Deutsche fauna ..., 1:86. + + 1810. _Cephalotes_ É. Geoffroy, Ann. du Mus. d'Hist. Nat., 15:104. + + 1811. _Harpyia_ Illiger, Prodr. Syst. Mamm. et Avium, p. 118. + + 1837. _Gelasinus_ Temminck, Monographe de Mammalia ..., 2:100. + +Tube-nosed bats of the genus _Nyctimene_ occur from Celebes on the west +to the Santa Cruz Islands on the east. Heretofore, two species (_N. +albiventer_ and _N. major_), each with an endemic subspecies, were known +from the Solomon Islands. Both species occur also in New Guinea and on +many adjacent islands. A new species of _Nyctimene_, apparently endemic +to the Solomons, and a new subspecies of _N. albiventer_ are named +beyond. + +_Nyctimene_ is related closely to _Cynopterus_ and the "Cynopterus +group" of Andersen (1912:691). Because _Nyctimene_ is a highly +specialized bat, Miller (1907:75) placed it in a subfamily separate from +that of _Cynopterus_ and its allies. + +Andersen (1912:696, 697) placed the species of _Nyctimene_ previously +known from the Solomons in two groups, the "papuanus" group and the +"cephalotes" group, on the basis of difference in length of forearm and +length of maxillary tooth-row. Because of its short forearm (about 58), +_N. albiventer_ is in the _papuanus_ group; and _N. major_, because of +its long forearm (about 74), is in the _cephalotes_ group. + +Key to Species of Nyctimene in the Solomons + + 1. Forearm longer than 70; males grayish-brown, females + pale gray, =N. major scitulus=, p. 825 + + 1'. Forearm shorter than 70; males dark brown, females + pale brown, 2 + + 2(1'). Forearm about 65, =N. malaitensis=, p. 822 + + 2'. Forearm less than 61, =N. albiventer=, p. 818 + + +=Nyctimene albiventer= + +This species occurs throughout New Guinea and on many adjacent islands, +including the Bismarck Archipelago and the Admiralty and Solomon +islands. The species varies geographically and five subspecies are +recognized. The two subspecies in the Solomons resemble _N. albiventer +papuanus_, the subspecies that ranges from eastern New Guinea to New +Britain. _N. albiventer bougainville_ occurs in the western chain of +islands of the Solomons, whereas another subspecies, named as new +beyond, occurs in the eastern chain of islands (see Fig. 12). + +Sexual dichromatism is striking. As Andersen (1912:690) previously +reported, females generally are paler, more brownish than males, which +are dark and have a better defined black dorsal stripe. + + +=Nyctimene albiventer bougainville= Troughton + + 1936. _Nyctimene bougainville_ Troughton, Rec. Australian Mus., + 19:349, April 7, type from Bougainville. + + 1954. _Nyctimene albiventer bougainville_, Laurie and Hill, List of + land mammals of New Guinea, Celebes and adjacent islands, p. 46, + June 30. + + 1953. _Nyctimene papuanus bougainville_, Pohle, Z. Säugetierk., + 17:130, October 27. + +_Specimens examined_ (nine males, one female; nine in alcohol, nine +crania extracted and cleaned).--Bougainville in December, AM-M. 5786 +(paratype); Guadalcanal in May, 23812, 23815, 23827; Kolombangara in +January and February, 23369, 23381, 23388, 23406, 23444, 23456. + +_Measurements._--See Table 3. + +_Remarks._--Heretofore, _Nyctimene albiventer bougainville_ was not +known from Kolombangara and Guadalcanal. The subspecies apparently +ranges throughout the western chain of the Solomons. + +Troughton (1936:350) considered _Nyctimene bougainville_ specifically +distinct from its nearest ally, _N. papuanus_. Pohle (1953:130) did not +examine specimens of either kind, but on the basis of Troughton's +description decided that _N. bougainville_ differed only subspecifically +from _N. papuanus_. Laurie and Hill (1954:46) synonymized _bougainville_ +and _papuanus_ with _N. albiventer_. However, Troughton (1936:350) +pointed out that in addition to size _bougainville_ differed from +_papuanus_ by having narrower and longer pm3 and pm4. Judging from +specimens examined by me, such is the case, and the difference is even +more pronounced in m1. + +Specimens of _N. a. bougainville_ from Kolombangara and Guadalcanal +agree with a paratype of this subspecies from Bougainville. Geographic +variation, if present in the population in the western chain of islands +(see Fig. 12), is slight and not notable in the series available. Some +individual variation was found, especially in the shape of the +interorbital region of the skull. An adult male from Kolombangara is +unusually dark, almost black; color of the other specimens (all in +alcohol) is consistent according to sex. + +_Nyctimene albiventer_ from Choiseul and Santa Ysabel is smaller, in all +respects, than _N. albiventer_ from Bougainville, Kolombangara, and +Guadalcanal (see Table 3), and therefore may be named and described as +follows: + + +=Nyctimene albiventer minor=, new subspecies + +_Type._--Adult male, skin and skull, in good condition (originally +stored in alcohol for about one year), no. BSIP 23636, Bernice P. Bishop +Museum; from Choiseul Island, British Solomon Islands Protectorate; +obtained on 11 March 1964, by Philip Temple, original number 1441. + +_Distribution._--Known only from Choiseul and Santa Ysabel islands (see +Fig. 12). + +_Diagnosis._--Small for _Nyctimene_; wing membranes brown with scattered +yellow spots (dried specimens); uropatagium, feet, and ears brown; +dorsum of tibia set with hair, ventral surface naked; dorsum of +uropatagium sparsely set with pale brown hairs, ventral surface almost +bare; fringe of hairs along two centimeters of dorsal and ventral +surfaces of trailing edge of wing membrane; proximal third of dorsal +surface of forearm sparsely set with hairs; pelage of back soft and +thick, of medium length (about 7); hair on crown and nape short (about +4); well-defined black dorsal stripe, extending from uropatagium to +shoulders; skull resembling that of other subspecies of _N. albiventer_ +but relatively smaller; zygomatic arch delicate, slender anteriorly; P2 +small (see Fig. 14). Sexually dichromatic as follows: male--dorsum +Hair-Brown, bases of hairs darker; hair on throat sparse, medium length +(about 6), Hair-Brown; fur along sides of abdomen Drab; female--dorsum +having Buffy-Brown cast, some individual hairs Hair-Brown; shoulders +Sayal-Brown; hair on throat sparse, Hair-Brown on throat and midline of +abdomen; sides of abdomen Sayal-Brown. + +_Comparisons._--From _Nyctimene major scitulus_, the largest member of +this genus in the Solomons, _N. a. minor_ differs in being smaller in +all measurements taken; forearm averaging 54.8 as opposed to 73.5; +greatest length of skull 28.2 as opposed to 37.0, and females pale brown +instead of pale gray. + +From nine adults of _Nyctimene albiventer bougainville_ from +Bougainville, Kolombangara, and Guadalcanal, _minor_ differs as follows: +averaging slightly smaller in all dimensions; forearm averaging 54.8 as +opposed to 57.9; second metacarpal averaging 27.4 as opposed to 28.3; +5th metacarpal averaging 38.5 as opposed to 40.0; condylobasal length +26.7 as opposed to 28.0; length of mandibular tooth-row 10.3 as opposed +to 10.9; mandible smaller (see Fig. 14); dorsal stripe fainter. + +From _Nyctimene albiventer papuanus_, known from eastern New Guinea, New +Britain, and the Admiralty Islands, _minor_ differs as follows: slightly +smaller in most dimensions; forearm averaging 54.8 as opposed to 57.0; +length of maxillary tooth-row 8.9 as opposed to 9.8; length of +mandibular tooth-row 10.3 as opposed to 11.0; breadth across upper third +premolars notably less (7.5 as opposed to 8.4). + +_N. a. minor_ differs from _N. albiventer albiventer_ Gray, which occurs +about 800 miles to the west of _minor_, in ways made apparent by the +description by Andersen (1912:700-701). _N. a. minor_ occurs about 1500 +miles eastward of the place from which _N. a. draconilla_ Thomas, a +subspecies essentially unknown to me, was named (see Laurie and Hill, +1954:46). + +From _Nyctimene sanctacrucis_, known from the Santa Cruz Islands, +_minor_ differs as follows: much smaller in all dimensions; forearm +averaging 54.8 as opposed to 75; greatest length of skull 28.2 as +opposed to 34.5; length of maxillary tooth-row 8.9 as opposed to 12.9. + + [Illustration: FIG. 12. Distribution of _Nyctimene albiventer + bougainville_ ([BC]) and _N. albiventer minor_ ([LW]). + For names of islands see Fig. 2.] + + [Illustration: FIG. 13. Scatter diagram comparing two subspecies + of _Nyctimene albiventer_. One individual of specimens thought + to be intergrades is as large as specimens of _Nyctimene a. + bougainville_, whereas the other three intergrades are about the + same size as specimens of _N. a. minor_. Symbols represent + _N. a. bougainville_ ([BW]), _N. a. minor_ ([TW]), and intergrades + assigned to _minor_ ([BC]). For names of islands see Fig. 2.] + + TABLE 3. Average and Extreme Measurements of _Nyctimene albiventer + bougainville_ and _N. a. minor_. + + ===============+==================+==================+=================== + | _N. a. minor_ | _Intergrades_ | _N. a. + | | | bougainville_ + | | | + MEASUREMENT | Choiseul, | Fauro | Kolombangara, + | Santa Ysabel | | Guadalcanal + | 4 [M], 1 [F] | 1 [M], 3 [F] | 8 [M], 1 [F] + ---------------+------------------+------------------+------------------- + Length of head | | | + and body |107.0 (105 -109 )|109.2 (105 -112 )|110.0 (106 -117) + Tail vertebrae | 20.0 (19.3- 20.5)| 21.0 ( 20 - 22 )| 19.2 ( 15.5- 23.0) + Hind foot | 14.2 (13.5- 15.0)| | 14.3 ( 13.0- 15.9) + Ear | 11.9 (11.0- 13.0)| | 12.8 ( 11.8- 14.5) + Length of | | | + forearm | 54.8 (54.0- 55.8)| 57.1 (55.9- 59.0)| 57.9 ( 55.8- 59.8) + Greatest | | | + length of | | | + skull | 28.2 (27.2- 28.9)| 28.6 (28.3- 29.7)| 29.7 ( 28.6- 30.1) + Condylobasal | | | + length | 26.7 (26.2- 27.5)| 27.4 (26.6- 28.0)| 28.0 ( 27.8- 28.9) + Palatal length | 11.2 (10.9- 11.9)| 11.6 (11.3- 11.8)| 11.7 ( 11.0- 12.5) + Breadth of | | | + braincase | 12.0 (11.5- 12.4)| 12.0 (11.7- 12.2)| 12.3 ( 12.1- 12.8) + Zygomatic | | | + breadth | 18.9 (18.4- 19.7)| 18.6 (18.4- 19.2)| 19.2 ( 18.7- 20.0) + Interorbital | | | + breadth | 5.0 ( 4.7- 5.6)| 5.3 ( 5.0- 5.6)| 5.1 ( 4.7- 5.5) + Breadth across | | | + first upper | | | + molars | 8.6 ( 8.4- 8.9)| 8.9 ( 8.7- 9.1)| 9.1 ( 8.8- 9.6) + Maxillary | | | + tooth-row | 8.9 ( 8.7- 9.3)| 9.3 ( 9.1- 9.5)| 9.5 ( 9.2- 9.8) + Mandibular | | | + tooth-row | 10.3 (10.0- 10.6)| 10.5 (10.2- 11.1)| 10.9 ( 10.7- 11.4) + ---------------+------------------+------------------+------------------- + +_Measurements._--Measurements of the two subspecies from the Solomons +are given in Table 3. Some measurements of the type are as follows: +Length of head and body, 108; tail vertebrae, 20.5; hind foot, 14.7; +ear, 11.3; length of forearm, 55.1; 2nd metacarpal, 27.4; 3rd +metacarpal, 39.0; 4th metacarpal, 37.5; 5th metacarpal, 39.1; greatest +length of skull, 28.6; condylobasal length, 27.5; zygomatic breadth, +18.4; length of maxillary tooth-row, 9.0; length of mandibular +tooth-row, 10.4. + +_Remarks._--_Nyctimene albiventer minor_ closely resembles _N. +albiventer bougainville_, differing from the latter mostly in size. +Although adults of _minor_ average only slightly smaller than adults of +_bougainville_ (see Table 3), there is only slight overlap (about 0.2 at +most) in most minimum dimensions of external and cranial features of +_bougainville_ and corresponding maximum dimensions of externals and +crania of _minor_. The difference in size is clearly shown in Figs. 13 +and 14. + +Four specimens of _Nyctimene albiventer_ from Fauro herein are +considered to be intergrades between _N. a. bougainville_ and _N. a. +minor_. As shown in Table 3, the specimens from Fauro average slightly +larger than those of _minor_ from Choiseul and Santa Ysabel and slightly +smaller than specimens of _bougainville_ from Kolombangara and +Guadalcanal. I have assigned the specimens from Fauro to _N. a. minor_ +because they generally are closer to _minor_ in size (see Fig. 13). + +_Specimens examined_ (five males and four females; seven in alcohol; +seven crania extracted and cleaned).--Choiseul in February and March, +23636 (holotype), 23631, 23540, 23646; Santa Ysabel in February, 23539; +Fauro in April, 23742, 23743, 23763, 23764. + +One specimen of _Nyctimene_ from Malaita Island is smaller than +_Nyctimene major_, which is known from Shortland, Alu, Florida, New +Georgia, Guadalcanal, Choiseul, and Malapa (see Fig. 15) and is larger +than either of the two subspecies of _Nyctimene albiventer_ known from +Bougainville, Fauro, Kolombangara, Guadalcanal, Choiseul, and Santa +Ysabel. This specimen represents a previously unknown species and may be +named and described as follows: + + +=Nyctimene malaitensis=, new species + +_Type._--Adult female, skin and skull, in good condition (originally +stored in alcohol for about one year), no. BSIP 24103, Bernice P. Bishop +Museum; from Malaita Island, British Solomon Islands Protectorate; +obtained on 1 July 1964, by Peter Shanahan, original no. unknown. + +_Distribution._--Known only from Malaita (see Fig. 16). + +_Diagnosis._--Size average for genus but larger than closest relative, +_Nyctimene albiventer_; wing membranes brown with scattered yellow spots +(dried specimen); uropatagium, ears, and feet brown; dorsal surface of +tibia set with hair, ventral surface bare; dorsal surface of uropatagium +sparsely set with hair, ventral surface having few, scattered hairs; +dorsal surface of trailing edge of wing membrane sparsely set with +hairs, ventral surface bare; proximal third of upper- and under-surface +of forearm set with hair; pelage of back luxuriant and soft (about 10 +long); hair on crown and nape shorter than on back (4 to 8); +well-defined black dorsal stripe from shoulders to rump (about 2 wide); +basal half of most hairs on dorsum Deep Mouse Gray, distal half Light +Buff, tips Ochraceous-Tawny; some hairs on back entirely Light Buff; +hairs of crown Light Ochraceous Buff tipped with Ochraceous-Tawny; hair +on throat and along sides of abdomen Light Ochraceous Buff; hairs of +ventral midline Smoke Gray; braincase narrow; zygomatic breadth +relatively narrow; well-developed lambdoidal crest in female; rostrum +short, wide; upper canines slanted posteriorly; upper incisors large; +foramen ovale large (see Fig. 14). + + [Illustration: FIG. 14. Dorsal and ventral views of skulls of + (A) _Nyctimene albiventer minor_ [specimen 23631 [M]], + (B) _N. a. bougainville_ [specimen 23381 [M]], and + (C) _N. malaitensis_ [specimen 24103 [F]].] + +_Comparisons._--From _Nyctimene major scitulus_, the largest kind of +_Nyctimene_ in the Solomons, _malaitensis_ differs as follows: smaller +in all dimensions (forearm 65 as opposed to 73.5); greatest length of +skull 32.4 as opposed to 37.0; length of maxillary tooth-row 10.5 as +opposed to 13.0; length of mandibular tooth-row 11.8 as opposed to 14.2. + +From nine adults of _Nyctimene albiventer bougainville_ from +Bougainville, Kolombangara, and Guadalcanal, _malaitensis_ differs as +follows: larger in all dimensions: forearm 65 as opposed to 57.9; +greatest length of skull 32.4 as opposed to 29.7; zygomatic breadth 20.4 +as opposed to 19.2; and length of maxillary tooth-row 10.5 as opposed to +9.5; length of mandibular tooth-row 11.8 as opposed to 11.1. + +From five adults of _Nyctimene albiventer minor_, from Choiseul and +Santa Ysabel, _malaitensis_ differs in the same ways it differs from _N. +a. bougainville_, but the contrast is even greater when _malaitensis_ +and _minor_ are compared. + +From _Nyctimene sanctacrucis_, known only from the Santa Cruz Islands, +_malaitensis_ differs in being smaller in all dimensions: forearm 65 as +opposed to 75; greatest length of skull 32.4 as opposed to 34.5; and +length of maxillary tooth-row 10.5 as opposed to 12.9. + +_Measurements of the holotype._--Length of head and body, 118; tail +vertebrae, 23.0; hind foot, 16.0; ear, 14.0; length of forearm, 65.0; +2nd metacarpal, 33.2; 3rd metacarpal, 46.4; 4th metacarpal, 44.3; 5th +metacarpal, 46.0; greatest length of skull, 32.4; condylobasal length, +30.6; palatal length, 13.0; breadth of braincase, 12.5; zygomatic +breadth, 20.4; interorbital breadth, 5.5; breadth across first upper +molars, 9.5; length of maxillary tooth-row, 10.5; length of mandibular +tooth-row, 11.8. + + [Illustration: FIG. 15. Distribution of _Nyctimene malaitensis_ [BC] + and _N. major scitulus_ [RW]. For names of islands see Fig. 2.] + +_Remarks._--In size, _Nyctimene malaitensis_ is intermediate between _N. +albiventer_ and _N. major_. Because the type of _malaitensis_ is brown +and not pale gray, as are females of _major_, _N. malaitensis_ most +likely is more closely related to _N. albiventer_, in which the females +are brown. The teeth of the holotype and only known specimen of +_malaitensis_ are too worn to be useful in determining the relationships +between these species. + +When more specimens are available, _N. malaitensis_ may prove to be a +subspecies of _N. albiventer_. At present, _malaitensis_ is accorded +specific rank in order not to obscure the apparent relationships of _N. +albiventer bougainville_ and _N. a. minor_. Additionally, _N. +malaitensis_ is given specific rank because (1) it is larger (especially +in external dimensions) than the largest subspecies of _N. albiventer_ +(compare above measurements with those in Table 3), and (2) +_malaitensis_ does not form a cline with either of the two subspecies of +_N. albiventer_. + +_Specimen examined_ (one female).--Malaita in July, 24103 (holotype). + + +=Nyctimene major= + +This large species of tube-nosed bat has at least four subspecies, one +of which (_N. major scitulus_) is endemic to the Solomons. The species +occurs throughout eastern New Guinea and on many of the islands adjacent +to the eastern coast of New Guinea, including the Trobriand Islands, +the Bismarck Archipelago, and the Solomons (see Laurie and Hill, +1954:47). The geographic distribution of the species generally is the +same as that of _N. albiventer_. + +In _Nyctimene major_, as in _N. albiventer_, most males are +grayish-brown, whereas most females are pale gray. + + +=Nyctimene major scitulus= Andersen + + 1910. _Nyctimene scitulus_ Andersen, Ann. Mag. Nat. Hist., ser. 8, + 6:623, December 1, type from Shortland; 1912, Andersen, Catalogue + of the Chiroptera ... British Museum, 1:711, from Shortland, New + Georgia, Florida, Guadalcanal; 1931, Troughton, Proc. Linnean Soc. + New South Wales, 56:206, July 15; 1931, Sanborn, Publ. Field Mus. + Nat. Hist., 18:22, February 12, from Choiseul and Malapa; 1942, + Tate, Bull. Amer. Mus. Nat. Hist., 80:342, December 31. + + 1954. _Nyctimene major scitulus_, Laurie and Hill, List of land + mammals of New Guinea, Celebes and adjacent islands, p. 47, + June 30. + + 1862. _Harpyia pallasi_, Gerrard, Catalogue of the bones ... + British Museum, p. 58. + + 1870. _Harpyia cephalotes_, Gray, Catalogue of monkeys, lemurs and + fruit-eating bats in the British Museum, p. 121. + + 1878. _Harpyia major_, Dobson, Catalogue of the Chiroptera ... + British Museum, p. 90; 1879, Trouessart, Rev. Mag. Zool., 3:207; + 1887, Thomas, Proc. Zool. Soc. London, p. 323; 1888, Thomas, Proc. + Zool. Soc. London, p. 476; 1897, Trouessart, Catalogus Mammalium + ..., 1:87. + + 1899. _Cephalotes major_, Trouessart, Catalogus Mammalium ..., + 2:1277. + + 1899. _Gelasinus major_, Matschie, Die Megachiroptera ... + naturkunde, p. 84; 1904, Trouessart, Catalogus Mammalium ..., + Suppl., p. 64. + +_Specimens examined_ (four males and one female; dried skins with skulls +inside).--Florida in October, 24397, 24413, 24418, 24419. + +_Measurements._--External measurements of four males and one female are, +respectively, as follows: Length of head and body, 134, 128, 134, 134, +136; tail vertebrae, 28, 23, 27, 26, 21; hind foot, 20, 16, 19, 16, 21; +ear, 17, 17, 17, 17, 18; length of forearm, 73.8, 68.0, 74.0, 73.6, +78.0. + +_Remarks._--_Nyctimene major scitulus_ has been recorded only from the +western chain of islands in the Solomons (see Fig. 15). Specimens +examined by me agree well in external dimensions and color with +specimens described by Andersen (1912:712) and Troughton (1931:206-207). + + + + +ZOOGEOGRAPHY AND SPECIATION + + +De Beaufort (1951:113) considered bats of "less zoogeographical +importance" than other mammals because the ocean is not an "absolute +barrier to their dispersal." Volant animals are ecologically terrestrial +and therefore are more nearly earthbound than De Beaufort's remarks +would suggest (see Miller, 1966:10). Indeed, many kinds of volant +animals are endemic to the Solomons. Birds, for example, are well +adapted for flight but pose some of the most complex zoogeographic +problems in the area of New Guinea and the Solomon Islands (Mayr, +1940:198; 1942:81-83; Koopman, 1957). Rapid speciation can take place +in any situation where there is a high degree of isolation (Wright, +1931; Lack, 1947). In fact, isolation is a most important factor in +speciation of insular populations (Baker, 1951:55). The one genus, nine +species, and 19 subspecies of megachiropterans that are endemic to the +Solomons (Table 4) obviously indicate that bats, although volant, can be +restricted to one or more islands long enough for new taxa to evolve. + + TABLE 4. A Summary of the Kinds of Megachiropteran Bats in the + Solomon Islands and Their Affinities with Faunas of Adjacent + Islands. + + ===========+========+==========+===========+============+=============== + | | | Common | Common to | Common to + | | Endemic | only to | Solomons, | Solomons, + | Totals | to | Solomons | Bismarcks, | New Hebrides, + | | Solomons | and | and | and + | | | Bismarcks | New Guinea | New Caledonia + -----------+--------+----------+-----------+------------+--------------- + Genera | 7 | 1 | 0 | 6 | 0 + Species | 16 | 9 | 1 | 6 | 1 + Subspecies | 20 | 19 | 0 | 0 | 1 + -----------+--------+----------+-----------+------------+--------------- + +The megachiropteran bats of the Solomons have their affinities with the +fauna of New Guinea (Table 4); the Solomons and New Guinea have six +genera and six species in common. Because the two areas never have been +connected (_via_ the Bismarck Archipelago) by dry land, bats probably +have reached the Solomons by flying from island to island (see Durham, +1963:357, 359, 361, 363). Deignan (1963:266) has dismissed voluntary or +involuntary flight as possible explanations for distributions of bats +and birds on islands of the Pacific. + +The taxonomic level of endemism can be used as an indicator of antiquity +(Dobzhansky, 1941; Koopman, 1958:429-430). The one megachiropteran genus +(_Pteralopex_) endemic to the Solomons apparently is an ancient relic. +Bats of this monotypic genus occur on Bougainville, Choiseul, Santa +Ysabel, and Guadalcanal (see Fig. 4). These four islands probably were +contiguous during the maximum lowering of sea level in the Pleistocene +(see Durham, 1963:362-363). Bats of the genus _Pteralopex_ are the only +kind in the Solomons having a distribution that can be correlated with +former land connections between islands. + +The distributions of 16 species of megachiropterans known from the +Solomons are summarized in Table 5 and in Figure 16. The larger islands +(in terms of surface area and elevation) in general have the highest +number of species (Guadalcanal 10, Choiseul 9, and Bougainville 8). But +Fauro, one of the smallest islands for which data are available, has six +species of megachiropterans whereas San Cristobal and Malaita, two of +the larger islands, have only three and four species, respectively. +Possibly this difference signals the need for additional collecting. + +Bougainville and Choiseul, about 60 miles apart, have seven species of +megachiropterans in common (Table 5). Fauro, 25 miles southeast of +Bougainville and 35 miles west of Choiseul, shares five species with +each of these islands (Fig. 16). _Pteralopex atrata_ and _Pteropus +rayneri_ occur on Choiseul and on Bougainville, but not on Fauro. +Individuals of these species are the largest fruit bats in the Solomons, +and their absence on Fauro suggests, therefore, that this small island +is ecologically unsuitable, at least in some months, for the support of +populations of bats that require relatively large amounts of food. The +small size of the island is consistent with this hypothesis, but several +other islands as small as Fauro do support populations of the large +kinds of _Pteropus_, at least in some months. + + TABLE 5. A Summary of Distribution of All Species of Megachiropteran + Bats Known from the Solomons. Only Islands Well Known Faunistically + Are Listed. + + Column headings: + + A: Bougainville I: Vella Lavella + B: Choiseul J: Kolombangara + C: Santa Ysabel K: Russell + D: Ndai L: Guadalcanal + E: Malaita M: San Cristobal + F: Florida N: Ugi + G: Fauro O: Rennell + H: Shortland P: Ontong Java + + ===================+==+==+==+==+==+==+==+==+==+==+==+==+==+==+==+== + SPECIES | A| B| C| D| E| F| G| H| I| J| K| L| M| N| O| P + -------------------+--+--+--+--+--+--+--+--+--+--+--+--+--+--+--+-- + R. amplexicaudatus | X| X| X| | X| | X| | | X| | X| | | | + P. atrata | X| X| X| | | | | | | | | X| | | | + Pt. hypomelanus | | | | | | | | | | | X| | | | | + Pt. admiralitatum | | | | X| | | | X| X| | X| X| | | | + Pt. tonganus | | | | | | | | | | | | | | | X| + Pt. howensis | | | | | | | | | | | | | | | | X + Pt. rayneri | X| X| X| | X| | | | X| X| | X| X| X| X| + Pt. woodfordi | | | | | | | X| | | X| X| X| | | | + Pt. mahaganus | X| | X| | | | | | | | | | | | | + D. inermis | X| X| X| | X| X| X| X| X| | | X| X| X| X| + M. lagochilus | X| X| | | | X| X| | | X| | X| X| | | + M. woodfordi | X| X| | | | | X| X| | | X| X| | | | + M. aurantius | | X| | | | X| | | | | | | | | | + N. albiventer | X| X| X| | | | X| | | X| | X| | | | + N. major | | X| | | | X| | X| | | | X| | | | + N. malaitensis | | | | | X| | | | | | | | | | | + +--+--+--+--+--+--+--+--+--+--+--+--+--+--+--+-- + Totals | 8| 9| 6| 1| 4| 4| 6| 4| 3| 5| 4|10| 3| 2| 3| 1 + -------------------+--+--+--+--+--+--+--+--+--+--+--+--+--+--+--+-- + +Santa Ysabel has six species of megachiropterans and 10 occur on +Guadalcanal (Table 5). These two islands, separated by about 100 miles +of water, share five species (_Rousettus amplexicaudatus_, _Pteralopex +atrata_, _Pteropus rayneri_, _Dobsonia inermis_, and _Nyctimene +albiventer_). The Nggela Group, in which Florida is the largest island +and the only one from which bats have been collected, is 50 miles +southeast of Santa Ysabel and 30 miles north of Guadalcanal (Fig. 16). +Four species of megachiropterans are known from Florida (_Dobsonia +inermis_, _Macroglossus lagochilus_, _Melonycteris aurantius_, and +_Nyctimene major_). Three of these are known from Guadalcanal and one +occurs on Santa Ysabel. This situation resembles the one involving +Fauro, Bougainville, and Choiseul because none of the large bats +(_Pteropus_ and _Pteralopex_) is known from Florida, even though two +species of large bats that occur on Santa Ysabel to the northwest occur +also on Guadalcanal to the south. Possibly Florida and the smaller +islands that comprise the Nggela Group are ecologically unsuitable for +large bats, or perhaps these small islands can support only limited +numbers of individuals during part of a year. + + [Illustration: FIG. 16. The number of megachiropteran species known + from individual islands (number within a circle) is compared with + the number of species common to two different islands (number + without a circle). For names of islands see Fig. 2.] + +Some of the small islands in the Solomons have populations of large +fruit bats. For example, _Pteropus admiralitatum_ and _P. hypomelanus_ +have been reported from the small islands in the Russell Group (Table +5). Possibly these species do not live concurrently in the Russells; +specimens of the two were obtained in different years. Two small +megachiropterans, _P. woodfordi_ and _Melonycteris woodfordi_, also +inhabit the Russells. Shortland, a small island about 15 miles south of +Bougainville, supports one large bat, _P. admiralitatum_, as well as +smaller megachiropterans. + +Kolombangara and Vella Lavella are about the same size and are separated +by about 15 miles of water. _Rousettus amplexicaudatus_, _Pteropus +rayneri_, _P. woodfordi_, _Macroglossus lagochilus_, and _Nyctimene +albiventer_ have been collected on Kolombangara but only _P. +admiralitatum_, _P. rayneri_, and _Dobsonia inermis_ have been found on +Vella Lavella. The difference in the known megachiropteran faunas is +more striking when one compares each island with adjacent islands. Two +species on Vella Lavella occur also on Choiseul, which is about 35 miles +northeastward, and two species occur also on Shortland, which is 120 +miles northwestward (Fig. 16). Four of the five megachiropterans on +Kolombangara also have been found on Choiseul, about 50 miles northward +(Table 5). _Pteropus rayneri_ is the only megachiropteran known from +both Kolombangara and Vella Lavella, even though the islands are +separated by only a few miles of water. Inadequate data possibly account +for the differences in the megachiropteran fauna, but I suspect that +some other factors are involved. Although Vella Lavella and Kolombangara +do have one species (_P. rayneri_) in common, a different subspecies +occurs on each island--_rubianus_ on Kolombangara and _lavellanus_ on +Vella Lavella (Fig. 17 and Table 6). This indicates that some factor or +factors are operating to keep megachiropterans from moving frequently or +easily from one island to the other. + +Each of several subspecies of species in the genus _Pteropus_ are known +from one or two small islands separated by only a few miles from other +islands on which different subspecies occur (see Fig. 6). Judging from +this kind of distribution, these bats do not move frequently from island +to island. Possibly this is because they cannot easily cross water +barriers, or are not inclined to do so because food is abundantly +available throughout the year on their home island. Because "flying +foxes" frequently are seen in flight over water several hundred yards +from shore, the first factor probably is unimportant--at least where +short distances are involved. It seems most likely that when abundant +food is available these bats have no reason to move even moderate +distances. + + [Illustration: FIG. 17. The number of subspecies of megachiropterans + known from individual islands (number within a circle) is compared + with the number of subspecies common to different islands (number + without a circle). For names of islands see Fig. 2.] + +Distributions of subspecies of polytypic species are summarized in Table +6 and Figure 17. Generally, more subspecies are known from the larger +islands than from the smaller islands (Guadalcanal with 5, Bougainville, +Choiseul, and Santa Ysabel with 4, Fauro with 2.) The distributions of +some subspecies can be used to judge the differential effectiveness of +water gaps between islands. The distribution of _Pteropus rayneri +lavellanus_ and _P. rayneri rubianus_ is an example. + +Choiseul and Santa Ysabel are separated by about 50 miles of water (see +Fig. 17) but have three subspecies in common (_Pteropus rayneri +grandis_, _Dobsonia inermis minimus_, and _Nyctimene albiventer +minor_.) Choiseul is about 50 miles from Kolombangara and about 35 miles +from Vella Lavella, but shares no subspecies with these smaller islands +although some species are shared (Tables 5 and 6). From these data one +can conclude that exchange of genes between populations on Choiseul and +populations on Santa Ysabel is frequent but for some reason exchange of +genes between populations on Vella Lavella and Choiseul and Kolombangara +and Choiseul is infrequent. A series of small islands (Rob Roy, Wagina, +and the Arnavon Islands, not named on the maps) connect Choiseul and +Santa Ysabel in stepping-stone fashion (see Fig. 17). Possibly these +small islands enhance movement of megachiropterans between Choiseul and +Santa Ysabel. + + TABLE 6. A Summary of Distribution of Polytypic Species of + Megachiropteran Bats in the Solomon Islands. Only Islands + Well Known Faunistically Are Listed. + + Column headings: + + A: Bougainville I: Vella Lavella + B: Choiseul J: Kolombangara + C: Santa Ysabel K: Russell + D: Ndai L: Guadalcanal + E: Malaita M: San Cristobal + F: Florida N: Ugi + G: Fauro O: Rennell + H: Shortland P: Ontong Java + + ===================+==+==+==+==+==+==+==+==+==+==+==+==+==+==+==+== + SUBSPECIES | A| B| C| D| E| F| G| H| I| J| K| L| M| N| O| P + -------------------+--+--+--+--+--+--+--+--+--+--+--+--+--+--+--+-- + P. atrata atrata | X| X| | | | | | | | | | | | | | + P. atrata anceps | | | X| | | | | | | | | X| | | | + Pt. a. solomonis | | | | | | | | | X| | X| X| | | | + Pt. a. colonus | | | | | | | | X| | | | | | | | + Pt. a. grandis | | | | X| | | | | | | | | | | | + Pt. r. rayneri | | | | | X| | | | | | | X| | | | + Pt. r. grandis | X| X| X| | | | | | | | | | | | | + Pt. r. rubianus | | | | | | | | | | X| | | | | | + Pt. r. lavellanus | | | | | | | | | X| | | | | | | + Pt. r. monoensis | | | | | | | | | | | | | | | | X + Pt. r. cognatus | | | | | | | | | | | | | X| X| | + Pt. r. rennelli | | | | | | | | | | | | | | | X| + D. i. inermis | X| | | | X| X| X| X| X| | | X| X| X| X| + D. i. minimus | | X| X| | | | | | | | | | | | | + N. a. bougainville | X| | | | | | | | | X| | X| | | | + N. a. minor | | X| X| | | | X| | | | | | | | | + +--+--+--+--+--+--+--+--+--+--+--+--+--+--+--+-- + Totals | 4| 4| 4| 1| 2| 1| 2| 2| 3| 2| 1| 5| 2| 2| 2| 1 + -------------------+--+--+--+--+--+--+--+--+--+--+--+--+--+--+--+-- + +Florida, of the Nggela Group, is approximately halfway between Santa +Ysabel and Guadalcanal. _Pteralopex atrata anceps_ occurs on Santa +Ysabel and on Guadalcanal but is unknown from Florida. Fauro lies +between Bougainville and Choiseul. _Pteralopex atrata atrata_ and +_Pteropus rayneri grandis_ occur on Choiseul and on Bougainville but are +unknown from Fauro. As suggested earlier, small islands like Fauro and +Florida possibly cannot support large fruit bats, although they probably +would utilize these small islands when in transit between larger +islands. + +Fauro apparently is important to the distribution of the two subspecies +of _Dobsonia inermis_ and _Nyctimene albiventer_ in the Solomons (see +Figs. 9 and 13). In both species, one subspecies is found in the eastern +chain of islands and one subspecies is found in the western chain. +Specimens of _Dobsonia inermis_ from Fauro and Bougainville can be +identified as the subspecies _inermis_ whereas those from Choiseul are +assignable to the subspecies _minimus_. _Nyctimene albiventer +bougainville_ occurs on Bougainville but specimens of _N. albiventer_ +from Fauro and Choiseul can be identified as the subspecies _minor_. +Although interchange of genes occurs between populations on Bougainville +and Fauro in the case of _D. inermis_, the population of _N. albiventer_ +on Fauro is at least partially isolated from the population on +Bougainville. + +Rennell and Ontong Java are relatively isolated from other islands in +the Solomons (see Fig. 17). Only one kind of bat (_Pteropus howensis_) +is known from Ontong Java and apparently is endemic to that atoll. +_Pteropus tonganus geddiei_, one of the megachiropterans that occurs on +Rennell (Table 5), also is found in the New Hebrides and on New +Caledonia (Table 4). This makes _P. t. geddiei_ the only megachiropteran +bat in the Solomons that is more closely related to bats on islands to +the southeast of the Solomons than to bats on other islands of the +Solomons, the Bismarcks, or New Guinea, to the north and west. The other +species of megachiropterans (_Dobsonia inermis_ and _Pteropus rayneri_) +on Rennell are found also on other islands in the Solomons. It is to be +noted that Mayr (1931) regarded the avifauna of Rennell as most nearly +like that of the New Hebrides and New Caledonia. He suggested that the +prevailing winds from the southeast have been important for birds that +have reached Rennell. The New Hebrides and New Caledonia are four and a +half times farther from Rennell than are San Cristobal and Guadalcanal. +On first consideration a person might doubt that the winds would be +favorable enough to compensate for the great distance between Rennell +and the New Hebrides and New Caledonia. Darlington (1938) has used the +formula X n/m to obtain a comparison of barriers of different widths. +[X = the probability of an individual crossing a barrier of width m; +the probability of an individual crossing a similar barrier of width n +is the ratio n/m.] If this formula is applied here, one finds that winds +from the southeast (that is, from the New Hebrides and New Caledonia) +would have to be more than 100 times more favorable than winds from the +northeast (from Guadalcanal and San Cristobal) in order to compensate +for the distance of Rennell from the New Hebrides and New Caledonia. +Even so, tropical storms with unusually strong winds, frequent during +some parts of the year, possibly account for the present distributional +pattern of bats and birds that live on Rennell. + +Whatever the means by which bats of the species _P. tonganus_ reached +Rennell, the fact remains that specimens from Rennell cannot be +distinguished from specimens of _P. tonganus geddiei_ from the New +Hebrides and New Caledonia, more than 500 miles to the southeast. + +NOTE: An important and interesting paper on zoogeography of bats, which +was published too late to be included here, is: Krzanowski, A., 1967, +The magnitude of islands and the size of bats (Chiroptera), Acta Zool. +Cracoviensia, 12:281-348. + + + + +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 für 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. Über die Fledertiere von Bougainville. Z. Säugetierk., + 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. Phillips + +*** END OF THE PROJECT GUTENBERG EBOOK 40112 *** |