Fig. 4.Distribution ofPteralopex atrata;P. atrata atrata() andP. atrata anceps(). For names of islands seeFig. 2.
Fig. 4.Distribution ofPteralopex atrata;P. atrata atrata() andP. atrata anceps(). For names of islands seeFig. 2.
Key to Subspecies ofPteralopex atrata
Pteralopex atrata atrataThomas
1888.Pteralopex atrataThomas, 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 atratais 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 withPteropus, that this behavior probably was a reaction from fear rather than an indication of general aggressiveness on the part ofPteralopex.
Pteralopex atrata ancepsAndersen
1909.Pteralopex ancepsAndersen, 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.
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 ancepswas 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 ofancepsand described the ways in which it differed from the dentition ofatrata. Although he (1912:437) concluded thatancepsandatratarepresented "two stages of specialization of ... dentition," there apparently are few, if any, dental differences between the two subspecies. Teeth of adults ofancepsdiffer from teeth of the immature type ofancepsas 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 subspeciesatrata(see Andersen, 1912:438, fig. 22) and not crowded as in the type ofanceps. Distance between individual cheek-teeth apparently increases with growth of the cranium and mandible.
Adults ofP. a. ancepsthat 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 toPteropusin the Solomon Islands
PteropusBrisson
1762.PteropusBrisson, Regnum animale ..., ed. 2, p. 153.
Remarks.—More species (seven) and subspecies (12) ofPteropusoccur in the Solomon Islands than of any other chiropteran genus. Other kinds ofPteropus, 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 (1964a, 1964b) have offered some additional remarks but groupings and suggested relationships of species ofPteropusalmost 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" ofPteropusin 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 areP. hypomelanus,P. admiralitatum,P. tonganus, andP. howensis. The first and second species were placed in thePteropus hypomelanusgroup 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 byP. mahaganusandP. 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). BothP. mahaganusandP. woodfordican be included in thePteropus scapulatusgroup of Andersen (1912:402).
Pteropus hypomelanus
Pteropus hypomelanusis 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).
Fig. 5.Distribution ofPteropus hypomelanus luteus(),Pteropus admiralitatum solomonis(),Pteropus a. colonus(),Pteropus a. goweri(),Pteropus tonganus geddiei(), andPteropus howensis(). For names of islands seeFig. 2.
Fig. 5.Distribution ofPteropus hypomelanus luteus(),Pteropus admiralitatum solomonis(),Pteropus a. colonus(),Pteropus a. goweri(),Pteropus tonganus geddiei(), andPteropus howensis(). For names of islands seeFig. 2.
Pteropus hypomelanus luteusAndersen
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 ofPteropus hypomelanusfrom eastern New Guinea and three nearby islands (Conflict Islands, Trobriand Islands, and Woodlark Island) asP. 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 typicalP. 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 forluteus, other subspecies ofP. hypomelanuswere known in dark phase as well as pale and intermediate phases of coloration (Andersen, 1912:122). The reported occurrence ofP. h. luteuson 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. goweriis known only from Gower (Ndai) Island, notably removed from the western chain of islands inhabited byP. a. colonusandP. a. solomonis. Only one other subspecies, from the Admiralty Islands, is known.
Pteropus admiralitatum solomonisThomas
1904.Pteropus solomonisThomas, 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) consideredPteropus admiralitatum, and especially the subspeciesP. a. solomonis, to be the easternmost "representative" ofPteropus hypomelanus. In comparison withP. hypomelanus luteus,P. a. solomonisdiffers 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 subspeciesP. a. solomonishas been recorded from a "chain" of islands that included Vella Lavella, Simbo, Ghizo, Ganongga, Banika, and Guadalcanal (seeFig. 5).
Pteropus admiralitatum colonusAndersen
1908.Pteropus colonusAndersen, 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 colonusis the largest of the three subspecies that occur in the Solomon Islands. It closely resemblesP. 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 inP. admiralitatum colonusis smaller than inP. 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 goweriTate
1934.Pteropus goweriTate, 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 goweriwas described from six specimens collected in 1930 by the Whitney South Sea Expedition (Tate, 1934:1). This subspecies closely resembles the other two subspecies ofP. admiralitatum(colonusandsolomonis) found in the Solomon Islands. Color and length of forearm (see key onp. 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 howensisTroughton
1931.Pteropus howensisTroughton, 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, —.
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.—ApparentlyPteropus howensisis confined to Ontong Java (Lord Howe Islands) located northeastward of the main body of islands that constitute the Solomon Archipelago (seeFig. 5). According to A. J. Nicholson, who collected the specimens listed above,P. howensisis 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 ofPteropus howensisdeposited 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 ofPteropusknown from Melanesia is not clear. Troughton (1931:204, 206) comparedP. howensiswithP. hypomelanusandP. admiralitatumand found that it resembled each of them. Tate (1934:2) noted that the skull ofP. admiralitatum goweriwas similar to that ofP. howensisin structure. The latter species is, however, larger (length of forearm 122 according to Troughton, 1931:205) than any subspecies ofP. admiralitatum(length of forearm 108-112). Also, the cheek-teeth ofP. howensisthat I have studied are relatively larger than those of eitherP. hypomelanusorP. admiralitatum. Furthermore, inP. howensisthere 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 ofP. howensisresemble those in a dull-colored specimen ofP. tonganusfrom 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 tonganushas 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 (1964a) recently has reported on the species in the New Hebrides.
Pteropus tonganus geddieiMacGillivary
1860.Pteropus geddieiMacGillivary, 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,inSarasin 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, SantaCruz Islands, Samoan Islands and Fiji Islands) rather than with those to the west (New Guinea). The subspeciesP. 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. bascilicusThomas 1915, apparently closely related togeddiei, 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 rayneriis endemic to the Solomon Islands. It is divisible into seven subspecies (seeFig. 6), which, exceptingP. r. rennelliandP. 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 andP. r. cognatus, known from two of the southernmost islands, has the shortest (about 121).
Fig. 6.Distribution ofPteropus rayneri:P. r. rayneri();P. r. grandis();P. r. lavellanus();P. r. monoensis();P. r. rubianus();P. r. cognatus();P. r. rennelli(). For names of islands seeFig. 2.
Fig. 6.Distribution ofPteropus rayneri:P. r. rayneri();P. r. grandis();P. r. lavellanus();P. r. monoensis();P. r. rubianus();P. r. cognatus();P. r. rennelli(). For names of islands seeFig. 2.
Adult males ofPteropus raynerihave well-developed tufts of hair on eachside 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 rayneriGray
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.
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 rayneriwas 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. rayneriis known from Guadalcanal and Malaita (seeFig. 6), and is of almost the same size asP. 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 inP. r. raynerithe rump is brightly colored and therefore contrasts strongly with the dark brown back. A specimen ofraynerifrom 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 grandisThomas
1887.Pteropus grandisThomas, 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).
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 grandisis the largest subspecies of the species. It is also the widest ranging subspecies, being found on six islands (seeFig. 6).
Although the specimens listed above agree well with descriptions of color given by Thomas (1887a: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 inthe 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 inP. r. grandisas ranging from 29.5 to 31.5. Ears of specimens that I examined varied from 31.0 to 37.0.
Pteropus rayneri rubianusAndersen
1908.Pteropus rubianusAndersen, 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.
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 forPteropus 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 ofP. r. rubianusandP. r. lavellanus. Sanborn (1931:16) allocated it to the subspeciesrubianuson the basis of length of forearm.
Andersen's descriptions (1908:366; 1912:256) ofrubianuswere 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 lavellanusAndersen
1908.Pteropus lavellanusAndersen, 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.
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 lavellanusinhabits islands geographically near those from whichP. r. rubianusis known (seeFig. 6) and in most respects the two subspecies closely resemble each other.P. r. lavellanusis 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 asP. r. rubianus, resembled the subspecieslavellanusin color and probably represents an intergrade between the two populations.
The color ofP. r. lavellanusis close to that ofP. r. rubianusexcept 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 ofP. r. rubianusunder 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 monoensisLawrence
1945.Pteropus rayneri monoensisLawrence, 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 monoensisis the most recently described subspecies ofP. rayneri. Lawrence (1945:63) judged that in most ways this bat is intermediate betweenP. r. grandisandP. r. lavellanus. Coloration ofmonoensisindicates affinity with the former, whereas length of forearm (145-148) approaches that in the latter. The small skull, narrow palate, and whitish rump patch ofmonoensisare differences that distinguish it fromgrandisandlavellanus. 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 ofP. 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 andmonoensismay therefore be confined to Mono Island.
Pteropus rayneri cognatusAndersen
1908.Pteropus cognatusAndersen, 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 ofPteropus rayneri cognatusfirst were reported under the namePteropus rayneribased on three specimens (one from San Cristobal and two from Guadalcanal). Because the description was based mostly on the two specimens from Guadalcanal, the namerayneriis applicable to the bats from that island. Andersen (1908:365) thought that specimens that he studied, from San Cristobal, were specifically distinct fromP. rayneriand he proposed the namePteropus cognatusfor them. Later, Hill (1962:9) reducedcognatusto subspecific status underP. rayneri.
PresentlyP. r. cognatusis known only from San Cristobal and the small adjacent island of Ugi (seeFig. 6).
Pteropus rayneri rennelliTroughton
1929.Pteropus rennelliTroughton, 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 rennelliwas known from but a single specimen. Hill (1962:7) reported two additional specimens and pointed out thatP. r. cognatusandP. r. rennelliprobably represent the extremes of an east-west cline in size.P. r. rennelliandP. r. cognatusdiffer 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 subspeciesrennelliandcognatusis close, both geographically and genetically. Longer forearm, longer metacarpals, and longer mandibular tooth-row serve to differentiaterennellifromcognatus.
Pteropus woodfordiThomas
1888.Pteropus woodfordiThomas, 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 austiniLawrence, Proc. New England Zool. Club, 23:59, March 26, from Florida.