Part 2

_Rousettus amplexicaudatus_ has at least three subspecies, one of which is endemic to the Solomon Islands. The species is wide-ranging, being known from as far west as Thailand (Ellerman and Morrison-Scott, 1966:93) and as far east as the Solomons.

=Rousettus amplexicaudatus hedigeri= Pohle

1953. _Rousettus amplexicaudatus hedigeri_ Pohle, Z. Säugetierk., 17:127, October 27, type from Bougainville.

1887. _Cynonycteris brachyotis_, Thomas, Proc. Zool. Soc. London, p. 323, March 15; 1888, Thomas, Proc. Zool. Soc. London, p. 475, December 4, from Fauro.

1889. _Xantharpyia brachyotis_, Matschie, Die Megachiroptera ... naturkunde, p. 68, from Guadalcanal.

1912. _Rousettus brachyotis_, Andersen, Catalogue of the Chiroptera ... British Museum, 1:809; 1931, Sanborn, Publ. Field Mus. Nat. Hist., Zool. Ser., 18:11, February 12, from Santa Ysabel.

_Specimens examined_ (20 males and 21 females; all in alcohol; ten crania extracted and cleaned).--Guadalcanal in May, 23863, 23915; Fauro in April, 23804-5; Malaita in June, 24079; Choiseul in March, 23563-4, 23616, 23627, 23630, 23632-3, 23642, 23658, 23663-4, 23680, 23692-3, 23713, 23722; Kolombangara in January and February, 23343, 23366, 23382-4, 23389-90, 23408-9, 23424, 23455, 23471-4, 23501.

_Measurements._--Average and extreme external measurements of 13 males and 18 females are, respectively, as follows: Length of head and body, 104.4 (99-118), 108.6 (104-117); tail vertebrae, 16.8 (13-19), 17.6 (15-24); hind foot, 18.0 (16-19), 16.2 (12-18); ear, 15.9 (15-17), 15.0 (14-16); length of forearm, 70.1 (66.0-74.1), 68.1 (65.0-69.1). Average and extreme measurements of skulls of five males and five females are, respectively, as follows: Greatest length of skull, 33.2 (33.0-33.7), 31.5 (30.9-32.1); condylobasal length, 31.3 (30.9-31.9), 30.1 (29.3-30.8); palatal length, 14.0 (13.3-14.8), 13.3 (13.0-13.7); zygomatic breadth, 20.8 (19.8-21.8), 19.4 (18.7-20.8); length of maxillary tooth-row, 11.0 (10.9-11.3), 10.3 (10.1-10.6); length of mandibular tooth-row, 12.6 (12.4-12.9), 11.8 (11.7-12.2).

_Remarks._--The specimens from Choiseul, Kolombangara, and Malaita islands provide new records of distribution for _Rousettus amplexicaudatus hedigeri_ (Fig. 3). It was described as smaller than _R. a. brachyotis_ Dobson, which is known from New Guinea, Amboina, and the Bismarck Archipelago (Pohle, 1953:127-128). Andersen (1912:809) gave the range of length of forearm in _R. a. brachyotis_ as 73-81, whereas Pohle (1953:127) gave the length of forearm of the type specimen of _R. a. hedigeri_ (adult male) as 67. Measurements of specimens examined by me indicate that _hedigeri_ occurs throughout the Solomon Islands. Cranial measurements of my specimens and Pohle's type are less than those of _R. a. brachyotis_ (see Andersen, 1912:48).

Sanborn (1931:11) noted that the forearms of three males examined by him were longer than that of a female. Mean and range for length of forearm of males and females listed herein, respectively, are 70.1 (66.0-74.1) and 68.1 (65.0-69.1). Also, each of seven cranial measurements taken by me averaged more in males than in females. Sagittal and lambdoidal crests are more prominent in males than in females.

TABLE 1. A Summary of Breeding Data for Females of _Rousettus amplexicaudatus hedigeri_ Collected December to June.

===========+===========+==============+===========+============= | Total | Number | | Number of MONTH | number | adult [F][F] | Number | immature | collected | collected | lactating | individuals -----------+-----------+--------------+-----------+------------- December | 3 | 3 | 3 | 0 January | 11 | 11 | 8 | 0 February | 6 | 0 | -- | 1 March | 16 | 1 | 0 | 9 April | 2 | 2 | 0 | 0 June | 1 | 1 | 0 | 0 -----------+-----------+--------------+-----------+-------------

As shown in Table 1, adult females obtained in December and January were lactating when captured whereas those obtained in March, April, and June were not. More than half of the individuals collected in March were immature (judging from small size, unfused epiphyses, and lack of wear on teeth). The immature individuals probably had been nursing in December and January.

=Pteralopex= Thomas

1888. _Pteralopex_ Thomas, Ann. Mag. Nat. Hist., ser. 6, 1:155, February 1.

1762. _Pteropus_ Brisson, Regnum animale ..., ed. 2, p. 153.

_Pteralopex_, with one species and two subspecies, is the only megachiropteran genus endemic to the Solomons. Thomas (1888b:475) considered this unusual bat a relic, isolated from the time when pteropodids had cuspidate cheek-teeth. Although two workers (Matschie, 1899:11; Simpson, 1945:54) have synonymized _Pteralopex_ with _Pteropus_, I regard _Pteralopex_ as a morphologically distinct genus.

Individuals of _Pteralopex_ can be distinguished from all species of _Pteropus_ in the Solomon Islands by the following features: wing membranes originate along dorsal midline; braincase diminutive relative to rest of skull; sagittal crest pronounced; cheek-teeth cuspidate, broad and massive; i2 about 10 times larger than i1; upper canines with well-developed secondary cusp; postorbital process fused with zygomatic arch, forming complete bony ring around orbit.

Andersen (1909a:216; 1912:436) considered the relationships of _Pteralopex_ and _Pteropus_ and concluded that _Pteropus pselaphon_ Lay, 1829, from the Sulphur Islands east of Taiwan, and _Pteropus samoensis_ Peale, 1848, from the Samoan Islands, were the "closest" living relatives of _Pteralopex_. He stated further that _Pteralopex_ "presents in fact scarcely a single character which is not either developed to a certain extent or at least distinctly foreshadowed in _Pteropus pselaphon_, _pilosus_, _tuberculatus_, or _leucopterus_." In summary, Andersen thought several species of _Pteropus_ had undergone evolutionary development resembling that in _Pteralopex_, and that the latter, with its massive, cuspidate cheek-teeth, could be considered a highly modified _Pteropus_. For this hypothesis to be plausible, one must assume that the originally complex cheek-teeth of pteropodids became simple and, at least in the case of _Pteralopex_, secondarily became complex once again. According to present-day theory of evolutionary development, his hypothesis is improbable. Thomas (1888b:475) probably was correct when he considered _Pteralopex_ an isolated relic.

Although _Pteralopex_ usually is listed after _Pteropus_ in phylogenetic arrangements (see, for example, Sanborn, 1931:21; Pohle, 1953:129; Laurie and Hill, 1954:40), I have placed _Pteralopex_ before _Pteropus_.

=Pteralopex atrata=

Two subspecies of _Pteralopex atrata_ (_P. a. atrata_ and _P. a. anceps_) have been named; specimens of both are rare in museum collections. Thomas (1888_a_:155) described adults of _atrata_. Sanborn (1931:21) examined the one additional specimen known to me and reported that it agreed with Thomas' description.

Andersen (1909_b_:266) used a subadult female ("nearly fully grown") as the holotype of _anceps_. At least five additional specimens, all adults, of _anceps_ now are housed in various collections. Judging from these individuals, the holotype of _anceps_ was only four-fifths grown and because he used an immature individual, Andersen's (1912:437) criteria for distinguishing the two subspecies mostly are invalid.

Key to Subspecies of _Pteralopex atrata_

1. Length of forearm 139-144 mm.; dorsal surface of distal one-fourth of tibia and entire metatarsus naked; known only from Guadalcanal and Santa Ysabel islands _Pteralopex atrata atrata_

1'. Length of forearm 162-166 mm.; dorsal surface of distal one-fourth of tibia and entire metatarsus furred; known only from Bougainville and Choiseul islands _Pteralopex atrata anceps_

=Pteralopex atrata atrata= Thomas

1888. _Pteralopex atrata_ Thomas, Ann. Mag. Nat. Hist., ser. 6, 1:155, February, type from Guadalcanal; 1888, Thomas, Proc. Zool. Soc. London, p. 475, December 4; 1896, Heude, Mém. Hist. Nat. Emp. China, 3:179; 1897, Trouessart, Catalogus Mammalium ..., 1:83; 1907, Miller, Bull. U. S. Nat. Mus., 57:60, June 29; 1912, Andersen, Catalogue of the Chiroptera ... British Museum, 1:439; 1931, Sanborn, Publ. Field Mus. Nat. Hist., Zool. Ser., 18:21, February 12, from Santa Ysabel.

1954. _Pteralopex atrata atrata_, Laurie and Hill, List of land mammals of New Guinea, Celebes and adjacent islands, p. 40, June 30.

1899. _Pteropus (Pteralopex) atrata_, Matschie, Die Megachiroptera ... naturkunde, p. 11; 1904, Trouessart, Catalogus Mammalium ..., Suppl., p. 49.

_Specimens examined._--None.

_Remarks._--_Pteralopex atrata atrata_ is known from four specimens from Guadalcanal and one from Santa Ysabel (Sanborn, 1931:21).

Sanborn (_loc. cit._) reported that a specimen wounded at night, while feeding on young green coconuts, was the only fruit bat that attempted to attack the collectors. Troughton (1936:348) has suggested, on the basis of his experiences with _Pteropus_, that this behavior probably was a reaction from fear rather than an indication of general aggressiveness on the part of _Pteralopex_.

=Pteralopex atrata anceps= Andersen

1909. _Pteralopex anceps_ Andersen, Ann. Mag. Nat. Hist., ser. 8, 3:266, March, type from Bougainville; 1912, Andersen, Catalogue of the Chiroptera ... British Museum, 1:437; 1936, Troughton, Rec. Australian Mus., 14:348, April 7; 1953, Pohle, Z. Säugetierk., 17:129, October 27.

1954. _Pteralopex atrata anceps_, Laurie and Hill, List of land mammals of New Guinea, Celebes and adjacent islands, p. 40, June 30.

_Specimens examined_ (three males, two females; one skull-only and one in alcohol).--Choiseul in March, 23682; Bougainville in July, USNM 276973-74, USNM 276928, USNM 277112.

_Measurements._--Measurements of three males and one female are, respectively, as follows: Length of head and body, 280, 271, 261, 255; hind foot, 50, 54, 52, 59; ear, 23, 23, 26, 22; length of forearm, 160, 162, 166, 171; greatest length of skull, 77.6, 77.9, 78.9, 77.0; condylobasal length, 74.3, 74.3, 75.5, 73.8; zygomatic breadth, 42.2, 45.4, 43.1, 42.6; breadth across upper canines, 18.7, 21.1, 19.0, 19.0; breadth across first upper molars, 22.2, 25.3, 22.9, 22.0; length of maxillary tooth-row, 29.3, 29.8, 28.9, 28.2; length of mandibular tooth-row, 32.8, 32.8, 32.1, 31.4.

_Remarks._--Heretofore, _Pteralopex atrata anceps_ was not known from Choiseul. The specimen from that island agrees well with specimens in the U. S. National Museum from Cape Torokina, Bougainville.

The type specimen of this subspecies is a subadult and is smaller than the specimens examined by me; Andersen (1912:440) gave length of forearm of the type as 137 (as opposed to 164 in adults). He (1912:438) figured the dentition of _anceps_ and described the ways in which it differed from the dentition of _atrata_. Although he (1912:437) concluded that _anceps_ and _atrata_ represented "two stages of specialization of ... dentition," there apparently are few, if any, dental differences between the two subspecies. Teeth of adults of _anceps_ differ from teeth of the immature type of _anceps_ as follows: in adults the anterior basal ledge of P4 extends onto the labial surface, whereas in the type it does not; and maxillary and mandibular teeth in adults are spaced as in the subspecies _atrata_ (see Andersen, 1912:438, fig. 22) and not crowded as in the type of _anceps_. Distance between individual cheek-teeth apparently increases with growth of the cranium and mandible.

Adults of _P. a. anceps_ that I examined are darker than the subadult type. The mantle in these adults is black, whereas it is seal-brown in the type (Andersen, 1912:439).

An adult female was lactating when obtained on Bougainville in July (USNM 276928).

Key to _Pteropus_ in the Solomon Islands

1. Premolars having distinct basal ledges; molars 2.5-4.0 wide 2

1'. Premolars lacking definite basal ledges; molars 1.0-2.4 wide 14

2(1). Rostrum unshortened (orbit to anterior tip of nasals about one-third greatest length of skull); dorsal surface of tibiae nearly naked 3

2'. Rostrum shortened (orbit to anterior tip of nasals less than one-third greatest length of skull); dorsal surface of tibiae usually at least partially furred 8

3(2). Forearm more than 128 4

3'. Forearm less than 128 5

4(3). Forearm about 155; venter and dorsum nearly black, mantle pale yellow =P. tonganus geddiei=, p. 798

4'. Forearm 128-136; venter and dorsum near Mars Brown, mantle Ochraceous or Cream-Buff =P. hypomelanus luteus=, p. 796

5(3'). Mantle dark, russet or cinnamon, not strongly contrasting with color of back =P. admiralitatum solomonis=, p. 796

5'. Mantle pale, Ochraceous-Buff or Cream-Buff, strongly contrasting with color of back 6

6(5'). Mantle Ochraceous-Orange to Ochraceous-Buff, hairs pale basally; forearm 108-111 =P. admiralitatum goweri=, p. 797

6'. Mantle Ochraceous to Cream-Buff, but hairs dark brown basally; forearm 110-122 7

7(6'). Length of forearm 110-112 =P. admiralitatum colonus=, p. 796

7'. Length of forearm about 122 =P. howensis=, p. 797

8(2'). Forearm more than 145 9

8'. Forearm less than 144 12

9(8). Forearm more than 162 10

9'. Forearm less than 162 11

10(9). Forearm 167-173 =P. rayneri grandis=, p. 801

10'. Forearm about 164 =P. rayneri rubianus=, p. 802

11(9'). Flanks and lower belly brightly colored, Burnt Sienna to Sanford's Brown; forearm less than 150 =P. rayneri monoensis=, p. 803

11'. Flanks and lower belly darker, near tawny; forearm more than 150, =P. rayneri lavellanus=, p. 802

12(8'). Pelage of dorsum tricolored; rump brightly colored; forearm 139-141, =P. rayneri rayneri=, p. 800

12'. Pelage of dorsum bicolored; rump dark; forearm less than 135 13

13(12'). Mantle tawny with some Ochraceous-Buff; forearm about 130, =P. rayneri rennelli=, p. 804

13'. Mantle russet, lacking Ochraceous-Buff; forearm about 121, =P. rayneri cognatus=, p. 803

14(1'). Forearm more than 131; dorsum Tawny Olive =P. mahaganus=, p. 806

14'. Forearm less than 100; dorsum dark brown =P. woodfordi=, p. 804

=Pteropus= Brisson

1762. _Pteropus_ Brisson, Regnum animale ..., ed. 2, p. 153.

_Remarks._--More species (seven) and subspecies (12) of _Pteropus_ occur in the Solomon Islands than of any other chiropteran genus. Other kinds of _Pteropus_, as yet unknown, may live there.

The relationships among the species of these large fruit-eating bats, commonly termed "flying foxes," are obscure and the genus is in need of revision. The basic, definitive work is still that of Andersen (1912). Tate (1942) and Felten (1964_a_, 1964_b_) have offered some additional remarks but groupings and suggested relationships of species of _Pteropus_ almost entirely are the products of Kund Andersen. According to present-day concepts of variation and speciation, Andersen's criteria are artificial.

Basically, there are three "species-groups" of _Pteropus_ in the Solomon Islands. The first is composed of species in which the rostrum is "unshortened" (its length about one third of greatest length of skull), and the cheek-teeth are of moderate size (M1 is 2.8-3.2 wide). The species are _P. hypomelanus_, _P. admiralitatum_, _P. tonganus_, and _P. howensis_. The first and second species were placed in the _Pteropus hypomelanus_ group by Andersen (1912:98).

In the second group the rostrum is "shortened" (its length less than one third of greatest length of skull) and the cheek-teeth are of moderate to large size (M1 3.3-4.1 wide). _Pteropus rayneri_, endemic to the Solomons and represented there by at least seven subspecies, fits into this category.

The third group is represented by _P. mahaganus_ and _P. woodfordi_. Both species are endemic to the Solomon Islands. In these species the rostrum is unshortened but the cheek-teeth are greatly reduced, especially in width (M1 is 1.0-2.2 wide). Both _P. mahaganus_ and _P. woodfordi_ can be included in the _Pteropus scapulatus_ group of Andersen (1912:402).

=Pteropus hypomelanus=

_Pteropus hypomelanus_ is a wide-ranging species of flying fox having at least seven subspecies; three occur in southeastern Asia, two on and near Celebes, and two in New Guinea and islands adjacent to the southeastern coast of New Guinea, including one island in the Solomons (Ellerman and Morrison-Scott, 1966:95; Laurie and Hill, 1954:32-33).

=Pteropus 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.