Transcriber's Notes
Text Emphasis
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_Text_ - Italics
=Text= - Bold
Symbolic Representations
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[F] Female symbol
[M] Male symbol
[BC] Circle Black
[TW] Circle Top White
[RW] Circle Right White
[BW] Circle Bottom White
[LW] Circle Left White
[RTW] Circle Right Third White
[LTW] Circle Left Third White
UNIVERSITY OF KANSAS PUBLICATIONS
MUSEUM OF NATURAL HISTORY
Vol. 16, No. 8, pp. 777-837, 17 figures in text
December 16, 1968
Systematics of Megachiropteran Bats in the Solomon Islands
BY
CARLETON J. PHILLIPS
UNIVERSITY OF KANSAS
LAWRENCE
1968
UNIVERSITY OF KANSAS PUBLICATIONS, MUSEUM OF NATURAL HISTORY
Editorial Committee: E. Raymond Hall, Chairman; Frank B. Cross, Editor;
Henry S. Fitch; J. Knox Jones, Jr.
Volume 16, No. 8, pp. 777-837, 17 figs.
Published December 16, 1968
UNIVERSITY OF KANSAS
Lawrence, Kansas
PRINTED BY
ROBERT R. (BOB) SANDERS, STATE PRINTER
TOPEKA, KANSAS
1968
[Illustration]
31-9490
Systematics of Megachiropteran Bats in the Solomon Islands
BY
CARLETON J. PHILLIPS
CONTENTS
PAGE
INTRODUCTION 781
GAZETTEER 783
METHODS AND MATERIALS 786
ACKNOWLEDGMENTS 786
KEY TO GENERA 787
_Rousettus amplexicaudatus hedigeri_ Pohle 788
_Pteralopex atrata atrata_ Thomas 791
_Pteralopex atrata anceps_ Andersen 792
_Pteropus hypomelanus luteus_ Andersen 796
_Pteropus admiralitatum solomonis_ Thomas 796
_Pteropus admiralitatum colonus_ Andersen 796
_Pteropus admiralitatum goweri_ Tate 797
_Pteropus howensis_ Troughton 797
_Pteropus tonganus geddiei_ MacGillivary 798
_Pteropus rayneri rayneri_ Gray 800
_Pteropus rayneri grandis_ Thomas 801
_Pteropus rayneri rubianus_ Andersen 802
_Pteropus rayneri lavellanus_ Andersen 802
_Pteropus rayneri monoensis_ Lawrence 803
_Pteropus rayneri cognatus_ Andersen 803
_Pteropus rayneri rennelli_ Troughton 804
_Pteropus woodfordi_ Thomas 804
_Pteropus mahaganus_ Sanborn 806
_Dobsonia inermis inermis_ Andersen 808
_Dobsonia inermis_ new subspecies 809
_Macroglossus lagochilus microtus_ Andersen 813
_Melonycteris aurantius_ Phillips 816
_Melonycteris woodfordi_ Thomas 816
_Nyctimene albiventer bougainville_ Troughton 818
_Nyctimene albiventer_ new subspecies 819
_Nyctimene_ new species 822
_Nyctimene major scitulus_ Andersen 825
ZOOGEOGRAPHY AND SPECIATION 825
LITERATURE CITED 834
INTRODUCTION
The Solomon Islands constitute an archipelago east of the large island
of New Guinea and more than a thousand miles off the northeastern coast
of Australia. This archipelago, which is principally of volcanic origin
although sedimentary layers of calcareous rocks occur on many islands
(Lever, 1934; Belkin, 1962), consists of a double chain of islands
having a northwest-southeast axis of more than 600 miles. The
archipelago is more or less an extension of New Guinea and in fact is
connected to it in stepping-stone fashion by New Britain, New Ireland,
and numerous smaller islands (see Fig. 1).
Australia and New Guinea have many kinds of mammals but the only
terrestrial mammals in the Solomon Islands are a species of the genus
_Phalanger_ (order Marsupialia), and several species of four genera of
rodents, one genus of which probably was introduced by man.
Additionally, several kinds of bats have reached and colonized the
Solomon Islands.
[Illustration: FIG. 1. Showing the Solomon Islands in relation to
major adjacent land masses.]
In the past 100 years at least 43 species and subspecies of Chiroptera
of 16 genera have been recorded from the Solomon Islands; of these 27
species and subspecies of seven genera are in the suborder
Megachiroptera. At least one genus of Megachiroptera is endemic as are
numerous species of other genera, and subspecies of still other species.
In 1963 and 1964, the Bernice P. Bishop Museum sent several collecting
parties to the British Solomon Islands Protectorate and the Australian
Trust Territory of New Guinea. In the Solomons, J. Linsley Gressitt,
Philip Temple, Peter Shanahan, and Ray Straatmann visited many of the
larger and more accessible islands and collected a wealth of zoological
materials. I have had the opportunity to study and report on specimens
of mammals, especially bats, collected by the persons named and
deposited in the Bishop Museum. This report is the third in a series on
bats from the Solomons (Phillips, 1966; 1967). Other specimens, mostly
obtained in 1944 by personnel of United States military units, are
stored in the United States National Museum and have been available for
study. Aims of the following report are to (1) identify the
megachiropteran bats to species and subspecies and (2) discuss
distribution of these bats in the Solomon Islands.
In all, 27 kinds (subspecies and monotypic species) of the order
Megachiroptera are known from the Solomon Islands. These pertain to
three subfamilies of the one family Pteropodidae.
The 43 Solomon Islands, having a total land area of more than 15,300
square miles (see Belkin, 1962:42-43), are listed in the gazetteer (see
also Figure 2). Politically, all of the Solomon Islands except Buka and
Bougainville, which are included in New Guinea Trust Territory under
mandate to Australia, are in the British Solomon Islands Protectorate.
The Solomons are within 300 to 700 miles of the equator and have a
fairly constant tropical climate, except at high elevations. The
temperature varies little; monthly mean temperature is between 81° and
83° F. and at sea level ranges from about 70° to 93° F. yearly (Belkin,
1962:42).
Southeast tradewinds are relatively constant from May to October and
this period, in general, is a dry season except at higher elevations on
windward coasts. From December to March prevailing winds are from the
north and precipitation throughout the island group is especially heavy.
Rainfall on the island of Tulagi averages about 120 inches per year
(Bryan, Edwin H., 1941; MS, p. 2, at Pacific Sci. Information Center,
Bishop Museum) and up to 300 inches have been recorded on the north
coast of Guadalcanal (Belkin, 1962:42-43). Occasional dry periods occur
even in the period of December to March.
Most islands of the Solomon Group support dense tropical rain forest.
Much of it has been modified by man. Some clearings and scattered
coconut plantations are found along coasts. On some of the larger
islands (for example, Guadalcanal) coastal scrub (especially on leeward
coasts) and extensive grassy areas are to be found. Additional notes on
vegetation are in the gazetteer.
The 165,000 persons living on the Solomon Islands are mostly
Melanesians but some are mixed Papuan, Malay, and Polynesian. These
native peoples are notorious for their cannibalistic tendencies; the
eating of human flesh usually was related to warfare, although
malefactors and human sacrifices accounted for some of the cannibalism
(Cranstone, 1961:29). Prior to the Second World War few Europeans
visited the Solomons and several islands still remain beyond reach of
modern-day technology. For example, Rennell and Bellona islands, south
of the main part of the archipelago, are visited only rarely, and then
only by a medical officer or the Resident Commissioner. According to
Troughton (1936:341), the islanders in the interior of Bougainville as
late as 1935, were prone to kill and feast upon strangers. In 1932,
Lewis (1951:37) felt that the natives of Malaita Island were especially
resistant to outside interference by Caucasians and reported that no
"white man or foreigner" was safe on Malaita.
Troughton (1936), who listed Melanesian names for mammals, indicated
that the native peoples distinguished between kinds of bats that closely
resembled one another. Of these, the only bats that seem to be used as
food belong to the genus _Pteropus_.
GAZETTEER
In the following list, currently-used names of islands are given; when
available, older names and variant spellings are indicated in
parentheses. For certain islands, especially those visited by field
parties from the Bishop Museum or those frequently mentioned in previous
literature on bats, some descriptive and ecological information also is
provided.
Latitude and longitude of islands are from publication no. 881 of the
Hydrographic Office of the United States Navy Department (Anonymous,
1944); names of islands were checked against a list by Brigham (1900);
descriptive information mostly is from reports by Temple and Straatmann
(1964, field notes, at the Department of Entomology, Bishop Museum).
ALU.--7° 07' S, 155° 54' E.
BANIKA.--9° 05' S, 155° 13' E.
BARA (Gera).--9° 31' S, 160° 31' E.
BELLONA (Bello).--11° 18' S, 159° 48' E.
BOUGAINVILLE (Mamamolimo).--6° 12' S, 155° 15' E. This is the largest
island in the Solomon Group, being 127 miles long (northwest to
southeast) and about 59 miles across at the widest place. The highest
elevations are 9850 and 10171 feet, at the tops of active volcanoes.
Ecologically, Bougainville is mostly dense rain forest, which is less
dense on the summits of higher mountains.
BUKA.--5° 15' S, 154° 38' E.
CHOISEUL.--7° 04' S, 157° 01' E. This island, formed along a
northwest-southeast line of low mountains (maximum elevation of 3500
feet), is about 90 miles long and 20 miles wide. Most collecting was at
Malangona (Sasamunga on some maps) on the southwestern coast.
FAURO.--6° 55' S, 156° 07' E. This small island, about 14 miles long
(north-south) and six miles wide (east-west), lies about 10 miles south
and east of Bougainville. Fauro is formed around a volcanic cone having
a maximum elevation of 1925 feet; it has considerable dense mangrove
swamp along the west coast, and mature rain forest with little
understory growth. Most collecting was at Toumoa, on one of two southern
peninsulas.
FLORIDA (Nggela).--9° 05' S, 160° 16' E. Florida, the main island in the
Nggela Island Subgroup, is mountainous and except for some small grassy
areas, supports dense rain forest. It is nearly 25 miles long
(east-west) and nine miles wide (north-south), with a maximum
elevation, at Mount Barnett, of about 1366 feet. Most collecting was at
Haleta, on the southwestern coast. At this locality there were scattered
mangrove swamps, rain forest, and gardens inland.
GANONGGA (Ronogo, Ronongo).--8° 03' S, 156° 35' E.
GATUKAI.--8° 47' S, 158° 12' E.
GHIZO (Gizo, Keso).--8° 05' S, 156° 59' E.
GOWER (N'dai).--7° 54' S, 160° 34' E.
GUADALCANAL (Guadalcanar).--9° 15' S, 159° 35' E. Guadalcanal is mostly
of volcanic origin and has an irregular chain of mountains along the
southern coast. The highest elevation is 8005 feet at Mount Popomanasiu.
This large island is nearly 80 miles long (east-west) and 25 miles wide
(north-south). Most of the northwestern part of Guadalcanal supports
_alang-alang_ grass. The remainder of the island is heavily wooded.
KILINAILAU (Cartaret).--4° 44' S, 155° 28' E.
KOLOMBANGARA (Duki, Kulambangara).--8° 00' S, 157° 05' E. Kolombangara,
formed from an extinct volcano, is about 18 miles in diameter and nearly
circular. The highest peaks, rising as precipitous cliffs in some
places, reach a maximum elevation of about 5000 feet. The vegetation is
mostly virgin rain forest. Mangrove swamp and small coconut groves occur
along the coast. Field parties from the Bishop Museum were able to reach
the highest elevations, and concentrated their work along the
southwestern side of the island.
[Illustration: FIG. 2. Solomon Islands. Principal islands
are named.]
MALAITA (Mala, Malanta, Malayta).--9° 00' S, 161° 00' E. This long (104
miles northwest to southeast), narrow (about 23 miles at its widest
spot) island, between Santa Ysabel and San Cristobal islands, is
basically of volcanic origin with some limestone (coral) deposits along
the coast. Mount Kolovrat, having an elevation of 4275 feet, is the
highest point. The Bishop Museum field party lived at Dala, in dense
rain forest about 12 miles north of Auki on the northwestern coast of
Malaita.
MALAPA.--9° 49' S, 160° 53' E.
MONO (Treasury).--7° 22' S, 155° 35' E. This is a small island (maximum
elevation 1150 feet) in the Treasury Island Subgroup just south of
Bougainville. Mono is about nine miles long (east-west) and five and one
half miles wide (north-south). The basic volcanic core is described in
field notes as topped with coral limestone.
NEW GEORGIA (Kausagi).--8° 20' S, 157° 30' E. The New Georgia Subgroup
is composed of 11 moderate-sized islands and islets. New Georgia Island,
the main member of the subgroup, is 50 miles long (northwest to
southeast) and from five to 30 miles wide. On the northern side several
volcanic peaks attain an elevation of about 3000 feet. The entire island
is forested.
NGGELA (Florida Islands).--4° 31' S, 154° 11' E. This subgroup consists
of several small to medium-sized islands between Guadalcanal and
Malaita. Florida is the main island.
NISSAN (Green, Sir Charles Hardy's).--4° 31' S, 154° 11' E.
NUKUMANU (Le Maira, Tasman).--4° 32' S, 159° 25' E.
ONTONG JAVA (Lord Howe Atoll, Liuniuwu).--5° 25' S, 159° 30' E.
PAVUVO.--9° 04' S, 159° 08' E.
RAMOS.--8° 16' S, 160° 11' E.
RENNELL.--11° 38' S, 160° 14' E. This island, of limestone (coral)
origin, along with Bellona, is nearly 100 miles southwest of any other
member of the Solomons and has been regarded, because of this distance,
as an oceanic island instead of a continental island. It is about 50
miles long (east-west) and nine miles wide (north-south); its highest
elevation is 500 feet.
ROVIANA (Rendova, Rovianna, Rubiana).--8° 21' S, 157° 20' E.
RUSSELL.--9° 04' S, 159° 12' E.
SAN CRISTOBAL (San Christoval, Bauro, Makira, Arussi).--11° 33' S, 161°
43' E. This island is composed mostly of ancient volcanic rock, has a
maximum elevation of 4100 feet, is nearly 70 miles long (northwest to
southeast) and 24 miles wide, and supports a dense rain forest.
SANTA YSABEL (George, Ysabel, San Isabel, Isbel, Mahaga).--8° 00' S,
159° 07' E. Santa Ysabel is a long (90 miles from northwest to
southeast), narrow (19 miles at the widest spot), forested island,
consisting of a single chain of volcanic mountains. The numerous bays
and mouths of rivers provide excellent anchorages. Collecting was at
Tatamba approximately two miles south of Tanambuli where the
considerable area of forest was dense and bamboo thickets were abundant.
SAVO (Savu).--9° 08' S, 159° 49' E.
SHORTLAND.--7° 03' S, 155° 47' E.
SIKAIANA (Stewart).--8° 22' S, 162° 44' E.
SIMBO (Narovo, Naorovo, Naravo, Navoro, Sembo).--8° 16' S, 156° 31' E.
STIRLING.--7° 25' S, 155° 35' E.
TANABULI (Tanambuli, Tunnibili, Tunnibilis, Tunnibul, Tunnivula).--8°
24' S, 159° 35' E.
TAUU (Marqueen, Mortlock).--4° 48' S, 157° 32' E.
TELIPARI.--8° 15' S, 157° 32' E.
UGI.--10° 14' S, 161° 44' E.
VANGUNO (Vangunu).--8° 39' S, 158° 00' E.
VELLA LAVELLA.--7° 43' S, 156° 40' E. The coastline is rugged and
indented by numerous small bays. Some peaks are 3000 feet high. The
southeastern half of Vella Lavella is said to consist of uplifted coral,
and to be thickly planted to coconut palms. The native population is
concentrated here. The northwestern half of the island is rain forest
and is nearly uninhabited. Most of the collecting was at Pusisama, on
the southern beach and on Ulo Crater, an extinct volcano at the middle
of the island.
YANNTA.--10° 20' S, 161° 20' E.
METHODS AND MATERIALS
The phylogenetic arrangement and nomenclature in the text beyond are
mainly that of Laurie and Hill (1954). The synonymies for accounts of
genera are as follows: (1) first use of the generic name employed along
with the original description, and (2) original proposals, in
chronological order, of other generic names subsequently applied to the
bat in the Solomons. The synonymies in accounts of species and
subspecies are as follows: (1) first use of the accepted name, followed
by its type locality, followed, in chronological order, by other
references to the first name-combination, (2) first use of the
name-combination employed herein (if different from the original
combination), followed, in chronological order, by other references to
the present name-combination, and (3) other name-combinations, in
chronological order, employed for the bat in the Solomons. The word
"part" is used in parentheses after a name if some specimens listed
under that name are from the Solomon Islands and are referable to the
species or subspecies being written about.
Unless noted otherwise, specimens listed as examined were prepared
originally as museum skins with skulls. Approximately 70 per cent of
bats collected in the Solomons were preserved in formalin and now are
stored in alcohol. Because it was necessary to obtain dimensions and
examine various morphological characteristics of skulls, many crania
were extracted from bats preserved in alcohol.
Although all specimens in the Bishop Museum from the Solomon Islands
have been catalogued with the prefix BBM-BSIP, catalogue numbers without
prefixes in the lists of specimens examined refer to this museum.
Catalogue numbers with the prefix USNM refer to specimens in the U. S.
National Museum and those with the prefix AM-M refer to specimens in the
Australian Museum.
Unless indicated otherwise, all measurements in this paper are in
millimeters and are of adults. Cranial measurements, and external
measurements of specimens stored in alcohol, were taken by me. The
cranial measurements were taken with dial calipers using techniques
described by Hall (1946:672-685). External measurements (except length
of forearm) of specimens originally prepared as dried study skins, were
transcribed from specimen labels.
Capitalized color nomenclature is from Ridgway (1912). Noncapitalized
color terms are from published reports that did not use Ridgway's
terminology.
ACKNOWLEDGMENTS
Financial support for this investigation was from (1) a United
States Army Medical Research and Development Command grant
(DA-MD-49-193-62-G65) to the Entomology Department of the Bernice P.
Bishop Museum, and (2) a National Science Foundation grant (2185-4703)
to the author, through the Committee on Systematics and Evolutionary
Biology of The University of Kansas. I am grateful to many individuals
who have helped me in various ways throughout the course of this study.
Dr. J. Linsley Gressitt, Chairman of the Entomology Department, Bernice
Bishop Museum, allowed me to study specimens collected by his
expeditions; Professors E. Raymond Hall and J. Knox Jones, Jr., of the
Museum of Natural History and the Department of Zoology, The University
of Kansas, offered advice and guidance and constructively reviewed the
manuscript. Other persons who have given me assistance and, in some
cases, arranged for loans of comparative materials, are: Dr. David H.
Johnson, Division of Mammals, United States National Museum; Mr. Hobart
M. Van Deusen and Dr. Richard G. Van Gelder, Archbold Expeditions and
Department of Mammalogy, American Museum of Natural History; Messrs.
Ellis LeG. Troughton and Basil Marlow, Mammal Department, The Australian
Museum; Dr. Joseph Curtis Moore, Department of Mammalogy, Field Museum
of Natural History; Mr. John Edwards Hill, Mammal Room, British Museum
(Natural History); Prof. William B. Davis, Department of Zoology, Texas
A & M University; Miss Barbara Lawrence, Museum of Comparative Zoology,
Harvard University. Messrs. Jerry R. Choate and H. H. Genoways, two
colleagues in zoology at The University of Kansas, have assisted me in
many ways, for which I am grateful. Linda Anne Phillips, my wife,
prepared many of the figures and tables used herein. I thank also
Setsuko Nakata, Edwin H. Bryan, Robert Bowan, and Ilse Koehler, who, as
staff members of the Bishop Museum, were especially helpful to me. Most
of the specimens reported herein were collected by Philip Temple and
Peter Shanahan.
Key to Genera
1. Uropatagium lacking, or, if present, deeply indented in
center; tail vertebrae absent, or if present, free 2
1'. Uropatagium present, not indented; tail vertebrae present,
free or in uropatagium MICROCHIROPTERA 1
2(1). External tail-vertebrae lacking, or, if present, less
than 3 mm long 3
2'. External tail-vertebrae more than 3 mm long 6
3(2). Small or medium-sized (forearm less than 50);
tongue long, extensile 4
3'. Large (forearm more than 80); tongue not long and
extensile 5
4(3). Uropatagium present; small claw present on second phalanx
of second digit; tail short (about 3 mm)
=Macroglossus=, p. 812
4'. Uropatagium absent; no claw on second phalanx of second
digit; no tail =Melonycteris=, p. 814
5(3'). Entire back set with hair; wing membranes not meeting
at middle of back =Pteropus=, p. 793
5'. Back naked; wing membranes meeting at middle of back,
=Pteralopex=, p. 790
6(2'). Nostrils having definite tubelike extensions
=Nyctimene=, p. 817
6'. Nostrils lacking tubelike extensions 7
7(6'). Forearm less than 80; large, sharp claw on second phalanx
of second digit; four upper incisors =Rousettus=, p. 787
7'. Forearm more than 90; small, blunt claw on second phalanx
of second digit; two upper incisors =Dobsonia=, p. 807
Family PTEROPODIDAE
Subfamily Pteropodinae
Rousettus Gray
1821. _Rousettus_ Gray, London Medical Repository, 15:299, April 1.
1843. _Xantharpyia_ Gray, List of species ... British Museum, p. 37.
1852. _Cynonycteris_ Peters, Reise nach Mossambique, p. 25.
The genus _Rousettus_ occurs throughout the tropical regions of the Old
World, and in the Solomons is readily distinguished from all other
megachiropteran genera by having both a small claw on the second digit
and free caudal vertebrae. The oriental species have been divided into
two groups on the basis of size (Tate, 1942:344). The subspecies
_Rousettus amplexicaudatus hedigeri_ appears to be the sole
representative of this genus in the Solomon Islands. Prior to 1953,
several workers (Thomas, 1887b:323, 1888b:475; Matschie, 1899:68;
Sanborn, 1931:11) used the name _Rousettus amplexicaudatus brachyotis_
for it, but Pohle (1953) suggested that the specimens from the Solomons
recorded by earlier workers were _R. a. hedigeri_ named by him on the
basis of the specimen that he saw from Bougainville.
=Rousettus amplexicaudatus=
_Rousettus amplexicaudatus_ has at least three subspecies, one of which
is endemic to the Solomon Islands. The species is wide-ranging, being
known from as far west as Thailand (Ellerman and Morrison-Scott,
1966:93) and as far east as the Solomons.
[Illustration: FIG. 3. Distribution of _Rousettus amplexicaudatus
hedigeri_. For names of islands see Fig. 2.]
=Rousettus amplexicaudatus hedigeri= Pohle
1953. _Rousettus amplexicaudatus hedigeri_ Pohle, Z. Säugetierk.,
17:127, October 27, type from Bougainville.
1887. _Cynonycteris brachyotis_, Thomas, Proc. Zool. Soc. London,
p. 323, March 15; 1888, Thomas, Proc. Zool. Soc. London, p. 475,
December 4, from Fauro.
1889. _Xantharpyia brachyotis_, Matschie, Die Megachiroptera ...
naturkunde, p. 68, from Guadalcanal.
1912. _Rousettus brachyotis_, Andersen, Catalogue of the Chiroptera
... British Museum, 1:809; 1931, Sanborn, Publ. Field Mus. Nat.
Hist., Zool. Ser., 18:11, February 12, from Santa Ysabel.
_Specimens examined_ (20 males and 21 females; all in alcohol; ten
crania extracted and cleaned).--Guadalcanal in May, 23863, 23915; Fauro
in April, 23804-5; Malaita in June, 24079; Choiseul in March, 23563-4,
23616, 23627, 23630, 23632-3, 23642, 23658, 23663-4, 23680, 23692-3,
23713, 23722; Kolombangara in January and February, 23343, 23366,
23382-4, 23389-90, 23408-9, 23424, 23455, 23471-4, 23501.
_Measurements._--Average and extreme external measurements of 13 males
and 18 females are, respectively, as follows: Length of head and body,
104.4 (99-118), 108.6 (104-117); tail vertebrae, 16.8 (13-19), 17.6
(15-24); hind foot, 18.0 (16-19), 16.2 (12-18); ear, 15.9 (15-17), 15.0
(14-16); length of forearm, 70.1 (66.0-74.1), 68.1 (65.0-69.1). Average
and extreme measurements of skulls of five males and five females are,
respectively, as follows: Greatest length of skull, 33.2 (33.0-33.7),
31.5 (30.9-32.1); condylobasal length, 31.3 (30.9-31.9), 30.1
(29.3-30.8); palatal length, 14.0 (13.3-14.8), 13.3 (13.0-13.7);
zygomatic breadth, 20.8 (19.8-21.8), 19.4 (18.7-20.8); length of
maxillary tooth-row, 11.0 (10.9-11.3), 10.3 (10.1-10.6); length of
mandibular tooth-row, 12.6 (12.4-12.9), 11.8 (11.7-12.2).
_Remarks._--The specimens from Choiseul, Kolombangara, and Malaita
islands provide new records of distribution for _Rousettus
amplexicaudatus hedigeri_ (Fig. 3). It was described as smaller than _R.
a. brachyotis_ Dobson, which is known from New Guinea, Amboina, and the
Bismarck Archipelago (Pohle, 1953:127-128). Andersen (1912:809) gave the
range of length of forearm in _R. a. brachyotis_ as 73-81, whereas Pohle
(1953:127) gave the length of forearm of the type specimen of _R. a.
hedigeri_ (adult male) as 67. Measurements of specimens examined by me
indicate that _hedigeri_ occurs throughout the Solomon Islands. Cranial
measurements of my specimens and Pohle's type are less than those of _R.
a. brachyotis_ (see Andersen, 1912:48).
Sanborn (1931:11) noted that the forearms of three males examined by him
were longer than that of a female. Mean and range for length of forearm
of males and females listed herein, respectively, are 70.1 (66.0-74.1)
and 68.1 (65.0-69.1). Also, each of seven cranial measurements taken by
me averaged more in males than in females. Sagittal and lambdoidal
crests are more prominent in males than in females.
TABLE 1. A Summary of Breeding Data for Females of _Rousettus
amplexicaudatus hedigeri_ Collected December to June.
===========+===========+==============+===========+=============
| Total | Number | | Number of
MONTH | number | adult [F][F] | Number | immature
| collected | collected | lactating | individuals
-----------+-----------+--------------+-----------+-------------
December | 3 | 3 | 3 | 0
January | 11 | 11 | 8 | 0
February | 6 | 0 | -- | 1
March | 16 | 1 | 0 | 9
April | 2 | 2 | 0 | 0
June | 1 | 1 | 0 | 0
-----------+-----------+--------------+-----------+-------------
As shown in Table 1, adult females obtained in December and January were
lactating when captured whereas those obtained in March, April, and
June were not. More than half of the individuals collected in March were
immature (judging from small size, unfused epiphyses, and lack of wear
on teeth). The immature individuals probably had been nursing in
December and January.
=Pteralopex= Thomas
1888. _Pteralopex_ Thomas, Ann. Mag. Nat. Hist., ser. 6, 1:155,
February 1.
1762. _Pteropus_ Brisson, Regnum animale ..., ed. 2, p. 153.
_Pteralopex_, with one species and two subspecies, is the only
megachiropteran genus endemic to the Solomons. Thomas (1888b:475)
considered this unusual bat a relic, isolated from the time when
pteropodids had cuspidate cheek-teeth. Although two workers (Matschie,
1899:11; Simpson, 1945:54) have synonymized _Pteralopex_ with
_Pteropus_, I regard _Pteralopex_ as a morphologically distinct genus.
Individuals of _Pteralopex_ can be distinguished from all species of
_Pteropus_ in the Solomon Islands by the following features: wing
membranes originate along dorsal midline; braincase diminutive relative
to rest of skull; sagittal crest pronounced; cheek-teeth cuspidate,
broad and massive; i2 about 10 times larger than i1; upper canines with
well-developed secondary cusp; postorbital process fused with zygomatic
arch, forming complete bony ring around orbit.
Andersen (1909a:216; 1912:436) considered the relationships of
_Pteralopex_ and _Pteropus_ and concluded that _Pteropus pselaphon_ Lay,
1829, from the Sulphur Islands east of Taiwan, and _Pteropus samoensis_
Peale, 1848, from the Samoan Islands, were the "closest" living
relatives of _Pteralopex_. He stated further that _Pteralopex_ "presents
in fact scarcely a single character which is not either developed to a
certain extent or at least distinctly foreshadowed in _Pteropus
pselaphon_, _pilosus_, _tuberculatus_, or _leucopterus_." In summary,
Andersen thought several species of _Pteropus_ had undergone
evolutionary development resembling that in _Pteralopex_, and that the
latter, with its massive, cuspidate cheek-teeth, could be considered a
highly modified _Pteropus_. For this hypothesis to be plausible, one
must assume that the originally complex cheek-teeth of pteropodids
became simple and, at least in the case of _Pteralopex_, secondarily
became complex once again. According to present-day theory of
evolutionary development, his hypothesis is improbable. Thomas
(1888b:475) probably was correct when he considered _Pteralopex_ an
isolated relic.
Although _Pteralopex_ usually is listed after _Pteropus_ in phylogenetic
arrangements (see, for example, Sanborn, 1931:21; Pohle, 1953:129;
Laurie and Hill, 1954:40), I have placed _Pteralopex_ before _Pteropus_.
=Pteralopex atrata=
Two subspecies of _Pteralopex atrata_ (_P. a. atrata_ and _P. a.
anceps_) have been named; specimens of both are rare in museum
collections. Thomas (1888_a_:155) described adults of _atrata_. Sanborn
(1931:21) examined the one additional specimen known to me and reported
that it agreed with Thomas' description.
Andersen (1909_b_:266) used a subadult female ("nearly fully grown") as
the holotype of _anceps_. At least five additional specimens, all
adults, of _anceps_ now are housed in various collections. Judging from
these individuals, the holotype of _anceps_ was only four-fifths grown
and because he used an immature individual, Andersen's (1912:437)
criteria for distinguishing the two subspecies mostly are invalid.
[Illustration: FIG. 4. Distribution of _Pteralopex atrata_;
_P. atrata atrata_ ([RW]) and _P. atrata anceps_ ([BC]).
For names of islands see Fig. 2.]
Key to Subspecies of _Pteralopex atrata_
1. Length of forearm 139-144 mm.; dorsal surface of distal
one-fourth of tibia and entire metatarsus naked; known only
from Guadalcanal and Santa Ysabel islands
_Pteralopex atrata atrata_
1'. Length of forearm 162-166 mm.; dorsal surface of distal
one-fourth of tibia and entire metatarsus furred; known only
from Bougainville and Choiseul islands
_Pteralopex atrata anceps_
=Pteralopex atrata atrata= Thomas
1888. _Pteralopex atrata_ Thomas, Ann. Mag. Nat. Hist., ser. 6,
1:155, February, type from Guadalcanal; 1888, Thomas, Proc. Zool.
Soc. London, p. 475, December 4; 1896, Heude, Mém. Hist. Nat.
Emp. China, 3:179; 1897, Trouessart, Catalogus Mammalium ...,
1:83; 1907, Miller, Bull. U. S. Nat. Mus., 57:60, June 29; 1912,
Andersen, Catalogue of the Chiroptera ... British Museum, 1:439;
1931, Sanborn, Publ. Field Mus. Nat. Hist., Zool. Ser., 18:21,
February 12, from Santa Ysabel.
1954. _Pteralopex atrata atrata_, Laurie and Hill, List of land
mammals of New Guinea, Celebes and adjacent islands, p. 40,
June 30.
1899. _Pteropus (Pteralopex) atrata_, Matschie, Die Megachiroptera
... naturkunde, p. 11; 1904, Trouessart, Catalogus Mammalium ...,
Suppl., p. 49.
_Specimens examined._--None.
_Remarks._--_Pteralopex atrata atrata_ is known from four specimens from
Guadalcanal and one from Santa Ysabel (Sanborn, 1931:21).
Sanborn (_loc. cit._) reported that a specimen wounded at night, while
feeding on young green coconuts, was the only fruit bat that attempted
to attack the collectors. Troughton (1936:348) has suggested, on the
basis of his experiences with _Pteropus_, that this behavior probably
was a reaction from fear rather than an indication of general
aggressiveness on the part of _Pteralopex_.
=Pteralopex atrata anceps= Andersen
1909. _Pteralopex anceps_ Andersen, Ann. Mag. Nat. Hist., ser. 8,
3:266, March, type from Bougainville; 1912, Andersen, Catalogue
of the Chiroptera ... British Museum, 1:437; 1936, Troughton,
Rec. Australian Mus., 14:348, April 7; 1953, Pohle,
Z. Säugetierk., 17:129, October 27.
1954. _Pteralopex atrata anceps_, Laurie and Hill, List of land
mammals of New Guinea, Celebes and adjacent islands, p. 40,
June 30.
_Specimens examined_ (three males, two females; one skull-only and one
in alcohol).--Choiseul in March, 23682; Bougainville in July, USNM
276973-74, USNM 276928, USNM 277112.
_Measurements._--Measurements of three males and one female are,
respectively, as follows: Length of head and body, 280, 271, 261, 255;
hind foot, 50, 54, 52, 59; ear, 23, 23, 26, 22; length of forearm, 160,
162, 166, 171; greatest length of skull, 77.6, 77.9, 78.9, 77.0;
condylobasal length, 74.3, 74.3, 75.5, 73.8; zygomatic breadth, 42.2,
45.4, 43.1, 42.6; breadth across upper canines, 18.7, 21.1, 19.0, 19.0;
breadth across first upper molars, 22.2, 25.3, 22.9, 22.0; length of
maxillary tooth-row, 29.3, 29.8, 28.9, 28.2; length of mandibular
tooth-row, 32.8, 32.8, 32.1, 31.4.
_Remarks._--Heretofore, _Pteralopex atrata anceps_ was not known from
Choiseul. The specimen from that island agrees well with specimens in
the U. S. National Museum from Cape Torokina, Bougainville.
The type specimen of this subspecies is a subadult and is smaller than
the specimens examined by me; Andersen (1912:440) gave length of forearm
of the type as 137 (as opposed to 164 in adults). He (1912:438) figured
the dentition of _anceps_ and described the ways in which it differed
from the dentition of _atrata_. Although he (1912:437) concluded that
_anceps_ and _atrata_ represented "two stages of specialization of ...
dentition," there apparently are few, if any, dental differences between
the two subspecies. Teeth of adults of _anceps_ differ from teeth of the
immature type of _anceps_ as follows: in adults the anterior basal ledge
of P4 extends onto the labial surface, whereas in the type it does not;
and maxillary and mandibular teeth in adults are spaced as in the
subspecies _atrata_ (see Andersen, 1912:438, fig. 22) and not crowded as
in the type of _anceps_. Distance between individual cheek-teeth
apparently increases with growth of the cranium and mandible.
Adults of _P. a. anceps_ that I examined are darker than the subadult
type. The mantle in these adults is black, whereas it is seal-brown in
the type (Andersen, 1912:439).
An adult female was lactating when obtained on Bougainville in July
(USNM 276928).
Key to _Pteropus_ in the Solomon Islands
1. Premolars having distinct basal ledges;
molars 2.5-4.0 wide 2
1'. Premolars lacking definite basal ledges;
molars 1.0-2.4 wide 14
2(1). Rostrum unshortened (orbit to anterior tip of nasals
about one-third greatest length of skull); dorsal
surface of tibiae nearly naked 3
2'. Rostrum shortened (orbit to anterior tip of nasals less
than one-third greatest length of skull); dorsal surface
of tibiae usually at least partially furred 8
3(2). Forearm more than 128 4
3'. Forearm less than 128 5
4(3). Forearm about 155; venter and dorsum nearly black,
mantle pale yellow =P. tonganus geddiei=, p. 798
4'. Forearm 128-136; venter and dorsum near Mars Brown,
mantle Ochraceous or Cream-Buff
=P. hypomelanus luteus=, p. 796
5(3'). Mantle dark, russet or cinnamon, not strongly
contrasting with color of back
=P. admiralitatum solomonis=, p. 796
5'. Mantle pale, Ochraceous-Buff or Cream-Buff, strongly
contrasting with color of back 6
6(5'). Mantle Ochraceous-Orange to Ochraceous-Buff, hairs pale
basally; forearm 108-111 =P. admiralitatum goweri=, p. 797
6'. Mantle Ochraceous to Cream-Buff, but hairs dark brown
basally; forearm 110-122 7
7(6'). Length of forearm 110-112
=P. admiralitatum colonus=, p. 796
7'. Length of forearm about 122 =P. howensis=, p. 797
8(2'). Forearm more than 145 9
8'. Forearm less than 144 12
9(8). Forearm more than 162 10
9'. Forearm less than 162 11
10(9). Forearm 167-173 =P. rayneri grandis=, p. 801
10'. Forearm about 164 =P. rayneri rubianus=, p. 802
11(9'). Flanks and lower belly brightly colored, Burnt Sienna
to Sanford's Brown; forearm less than 150
=P. rayneri monoensis=, p. 803
11'. Flanks and lower belly darker, near tawny; forearm
more than 150, =P. rayneri lavellanus=, p. 802
12(8'). Pelage of dorsum tricolored; rump brightly colored;
forearm 139-141, =P. rayneri rayneri=, p. 800
12'. Pelage of dorsum bicolored; rump dark; forearm
less than 135 13
13(12'). Mantle tawny with some Ochraceous-Buff; forearm
about 130, =P. rayneri rennelli=, p. 804
13'. Mantle russet, lacking Ochraceous-Buff; forearm
about 121, =P. rayneri cognatus=, p. 803
14(1'). Forearm more than 131; dorsum Tawny Olive
=P. mahaganus=, p. 806
14'. Forearm less than 100; dorsum dark brown
=P. woodfordi=, p. 804
=Pteropus= Brisson
1762. _Pteropus_ Brisson, Regnum animale ..., ed. 2, p. 153.
_Remarks._--More species (seven) and subspecies (12) of _Pteropus_ occur
in the Solomon Islands than of any other chiropteran genus. Other kinds
of _Pteropus_, as yet unknown, may live there.
The relationships among the species of these large fruit-eating bats,
commonly termed "flying foxes," are obscure and the genus is in need of
revision. The basic, definitive work is still that of Andersen (1912).
Tate (1942) and Felten (1964_a_, 1964_b_) have offered some additional
remarks but groupings and suggested relationships of species of
_Pteropus_ almost entirely are the products of Kund Andersen. According
to present-day concepts of variation and speciation, Andersen's criteria
are artificial.
Basically, there are three "species-groups" of _Pteropus_ in the Solomon
Islands. The first is composed of species in which the rostrum is
"unshortened" (its length about one third of greatest length of skull),
and the cheek-teeth are of moderate size (M1 is 2.8-3.2 wide). The
species are _P. hypomelanus_, _P. admiralitatum_, _P. tonganus_, and _P.
howensis_. The first and second species were placed in the _Pteropus
hypomelanus_ group by Andersen (1912:98).
In the second group the rostrum is "shortened" (its length less than one
third of greatest length of skull) and the cheek-teeth are of moderate
to large size (M1 3.3-4.1 wide). _Pteropus rayneri_, endemic to the
Solomons and represented there by at least seven subspecies, fits into
this category.
The third group is represented by _P. mahaganus_ and _P. woodfordi_.
Both species are endemic to the Solomon Islands. In these species the
rostrum is unshortened but the cheek-teeth are greatly reduced,
especially in width (M1 is 1.0-2.2 wide). Both _P. mahaganus_ and _P.
woodfordi_ can be included in the _Pteropus scapulatus_ group of
Andersen (1912:402).
=Pteropus hypomelanus=
_Pteropus hypomelanus_ is a wide-ranging species of flying fox having at
least seven subspecies; three occur in southeastern Asia, two on and
near Celebes, and two in New Guinea and islands adjacent to the
southeastern coast of New Guinea, including one island in the Solomons
(Ellerman and Morrison-Scott, 1966:95; Laurie and Hill, 1954:32-33).
[Illustration: FIG. 5. Distribution of _Pteropus hypomelanus luteus_
([TW]), _Pteropus admiralitatum solomonis_ ([RW]), _Pteropus a.
colonus_ ([BW]), _Pteropus a. goweri_ ([LW]), _Pteropus tonganus
geddiei_ ([RTW]), and _Pteropus howensis_ ([BC]). For names of
islands see Fig. 2.]
=Pteropus hypomelanus luteus= Andersen
1908. Pteropus hypomelanus luteus Andersen, Ann. Mag. Nat. Hist.,
ser. 8, 2:362, October, type from Kiriwini Island, Trobriand
Islands; 1912, Andersen, Catalogue of the Chiroptera ... British
Museum, 1:128; 1947, Sanborn and Beecher, Jour. Mamm., 28:388,
November 19, from Banika Island, Russell Islands.
_Specimens examined._--None.
_Remarks._--Andersen (1908:362) identified specimens of _Pteropus
hypomelanus_ from eastern New Guinea and three nearby islands (Conflict
Islands, Trobriand Islands, and Woodlark Island) as _P. hypomelanus
luteus_. Sanborn and Beecher (1947:388) identified a female from Banika
Island in the Solomons as of this subspecies although this specimen was
darker and had a slightly smaller skull than typical _P. hypomelanus
luteus_. They noted that the pelage of the venter of the female was
uniformly dark rather than the typical Ochraceous-Buff to Cream-Buff;
the specimen was regarded as a dark phase of the subspecies. Although
not recorded previously for _luteus_, other subspecies of _P.
hypomelanus_ were known in dark phase as well as pale and intermediate
phases of coloration (Andersen, 1912:122). The reported occurrence of
_P. h. luteus_ on Banika Island extended the known geographic range
about 450 miles eastward from Woodlark Island.
=Pteropus admiralitatum=
Three subspecies, all about the same size but differing in coloration,
have been described from the Solomon Islands. _P. a. goweri_ is known
only from Gower (Ndai) Island, notably removed from the western chain of
islands inhabited by _P. a. colonus_ and _P. a. solomonis_. Only one
other subspecies, from the Admiralty Islands, is known.
=Pteropus admiralitatum solomonis= Thomas
1904. _Pteropus solomonis_ Thomas, Novit. Zool., 11:597, type from
Ghizo Island; 1912, Andersen, Catalogue of the Chiroptera ...
British Museum, 1:149; 1931, Sanborn, Publ. Field Mus. Nat. Hist.,
Zool. Ser., 18:12, February 12, from Ronongo (Ganongga), Vella
Lavella, and Narovo (Simbo) islands; 1947, Sanborn and Beecher,
Jour. Mamm., 28:389, November 19, from Banika and Guadalcanal
islands.
1954. _Pteropus admiralitatum solomonis_, Laurie and Hill, List of
land mammals of New Guinea, Celebes and adjacent islands, p. 33,
June 30.
_Specimens examined._--None.
_Remarks._--Andersen (1912:149) considered _Pteropus admiralitatum_, and
especially the subspecies _P. a. solomonis_, to be the easternmost
"representative" of _Pteropus hypomelanus_. In comparison with _P.
hypomelanus luteus_, _P. a. solomonis_ differs mostly in size, being
much smaller (length of forearm about 110 rather than 134). It is now
known that both species occur on Banika Island in the Solomons.
The subspecies _P. a. solomonis_ has been recorded from a "chain" of
islands that included Vella Lavella, Simbo, Ghizo, Ganongga, Banika, and
Guadalcanal (see Fig. 5).
=Pteropus admiralitatum colonus= Andersen
1908. _Pteropus colonus_ Andersen, Ann. Mag. Nat. Hist., ser. 8,
2:363, October, type from Shortland Island; 1912, Andersen,
Catalogue of the Chiroptera ... British Museum, 1:150; 1931,
Sanborn, Publ. Field Mus. Nat. Hist., Zool. Ser., 18:12,
February 12, from Mono Island.
1954. _Pteropus admiralitatum colonus_, Laurie and Hill, List of
land mammals of New Guinea, Celebes and adjacent islands, p. 33,
June 30.
1887. _Pteropus hypomelanus_ (part), Thomas, Proc. Zool. Soc.
London, p. 471, December 4; 1898, Trouessart, Catalogus Mammalium
..., 1:82, from "I. Salomonis."
1899. _Pteropus (Spectrum) hypomelanus_ (part), Matschie, Die
Megachiroptera ... naturkunde, p. 24.
_Specimens examined._--None.
_Remarks._--_Pteropus admiralitatum colonus_ is the largest of the three
subspecies that occur in the Solomon Islands. It closely resembles _P.
hypomelanus luteus_, except in being smaller throughout (see Andersen,
1912:151-152, for measurements) and darker on the underparts.
This bat has been found in a group of small islands (Alu, Mono, and
Shortland) about 30 miles south of Bougainville. Because of this
proximity and because yet another subspecies of this species occurs
northward of Bougainville, it is interesting that neither Troughton
(1936) nor Pohle (1953) included the species in their faunal lists for
Bougainville.
Andersen (1912:152) indicated that the M1 in _P. admiralitatum colonus_
is smaller than in _P. a. solomonis_, the subspecies found in islands to
the southeast (4.4-4.5 and 5.2, respectively), but Sanborn (1931:13)
studied specimens of these two subspecies that overlapped in size of M1.
=Pteropus admiralitatum goweri= Tate
1934. _Pteropus goweri_ Tate, Amer. Mus. Novit., 718:1, May 4, type
from Gower (Ndai) Island.
1954. _Pteropus admiralitatum goweri_, Laurie and Hill, List of
land mammals of New Guinea, Celebes and adjacent islands, p. 33,
June 30.
_Specimens examined._--None.
_Remarks._--_Pteropus admiralitatum goweri_ was described from six
specimens collected in 1930 by the Whitney South Sea Expedition (Tate,
1934:1). This subspecies closely resembles the other two subspecies of
_P. admiralitatum_ (_colonus_ and _solomonis_) found in the Solomon
Islands. Color and length of forearm (see key on p. 793) seem to be the
only reliable criteria for distinguishing between these subspecies. The
longitude of Gower Island, 160° 34' E, was incorrectly listed in Laurie
and Hill (1954:152) as 159° 34' E.
=Pteropus howensis= Troughton
1931. _Pteropus howensis_ Troughton, Proc. Linn. Soc. New South
Wales, 56:204, June 24, type from Lord Howe Islands (Ontong Java);
1950, Sanborn and Nicholson, Fieldiana:Zool., 31:329, August 31.
_Specimens examined_ (one male, three females, and two sex unknown; two
embryos in alcohol).--Liuniuwu, Lord Howe Islands (Ontong Java) in
August, USNM 278703-6, USNM 279715-6.
_Measurements._--Average and extreme measurements of one male and three
females are as follows: Length of head and body, 185.2 (176-196); hind
foot, 34.5 (33-36); ear, 21.5 (21-23); forearm not measured [broken in
all specimens examined]. Cranial measurements of a male and a female
are, respectively, as follows: Greatest length of skull, 55.3, 53.8;
condylobasal length, 54.2, 52.8; palatal length, 26.7, 26.0; zygomatic
breadth, 30.6, 29.9; breadth of braincase, 19.9, 19.2; breadth across
first upper molars, 14.3, 14.3; length of maxillary tooth-row, 20.7,
19.6; length of mandibular tooth-row, 23.1, --.
_Remarks._--Apparently _Pteropus howensis_ is confined to Ontong Java
(Lord Howe Islands) located northeastward of the main body of islands
that constitute the Solomon Archipelago (see Fig. 5). According to A. J.
Nicholson, who collected the specimens listed above, _P. howensis_ is
not abundant in Ontong Java. He related this circumstance to the fact
that these small islands are nothing more than parts of a coral atoll
used almost entirely for the production of coconuts (see Sanborn and
Nicholson, 1950:329).
Specimens of _Pteropus howensis_ deposited in the U. S. National Museum
agree well in most ways with the original description of the species by
Troughton (1931:204-205). Slight variation in color is evident; in two
specimens, the mantle, just posterior to the ears, is Ochraceous-Buff.
The relationship of this species to other kinds of _Pteropus_ known from
Melanesia is not clear. Troughton (1931:204, 206) compared _P. howensis_
with _P. hypomelanus_ and _P. admiralitatum_ and found that it resembled
each of them. Tate (1934:2) noted that the skull of _P. admiralitatum
goweri_ was similar to that of _P. howensis_ in structure. The latter
species is, however, larger (length of forearm 122 according to
Troughton, 1931:205) than any subspecies of _P. admiralitatum_ (length
of forearm 108-112). Also, the cheek-teeth of _P. howensis_ that I have
studied are relatively larger than those of either _P. hypomelanus_ or
_P. admiralitatum_. Furthermore, in _P. howensis_ there is a small but
distinct cusp located medio-posteriorly on P4 (most noticeable in young
individuals) that is more reduced or undeveloped in specimens of the
other two species. Cheek-teeth of _P. howensis_ resemble those in a
dull-colored specimen of _P. tonganus_ from Fiji Island with which I
compared the specimens listed above.
Weights and crown-rump lengths of the two embryos (in an advanced stage
of development) examined were 20 and 29 grams and 43 and 51 mm.
(apparently these are the specimens listed by Sanborn and Nicholson,
1950:329).