Part 3

_Remarks._--_Pteropus admiralitatum goweri_ was described from six specimens collected in 1930 by the Whitney South Sea Expedition (Tate, 1934:1). This subspecies closely resembles the other two subspecies of _P. admiralitatum_ (_colonus_ and _solomonis_) found in the Solomon Islands. Color and length of forearm (see key on p. 793) seem to be the only reliable criteria for distinguishing between these subspecies. The longitude of Gower Island, 160° 34' E, was incorrectly listed in Laurie and Hill (1954:152) as 159° 34' E.

=Pteropus howensis= Troughton

1931. _Pteropus howensis_ Troughton, Proc. Linn. Soc. New South Wales, 56:204, June 24, type from Lord Howe Islands (Ontong Java); 1950, Sanborn and Nicholson, Fieldiana:Zool., 31:329, August 31.

_Specimens examined_ (one male, three females, and two sex unknown; two embryos in alcohol).--Liuniuwu, Lord Howe Islands (Ontong Java) in August, USNM 278703-6, USNM 279715-6.

_Measurements._--Average and extreme measurements of one male and three females are as follows: Length of head and body, 185.2 (176-196); hind foot, 34.5 (33-36); ear, 21.5 (21-23); forearm not measured [broken in all specimens examined]. Cranial measurements of a male and a female are, respectively, as follows: Greatest length of skull, 55.3, 53.8; condylobasal length, 54.2, 52.8; palatal length, 26.7, 26.0; zygomatic breadth, 30.6, 29.9; breadth of braincase, 19.9, 19.2; breadth across first upper molars, 14.3, 14.3; length of maxillary tooth-row, 20.7, 19.6; length of mandibular tooth-row, 23.1, --.

_Remarks._--Apparently _Pteropus howensis_ is confined to Ontong Java (Lord Howe Islands) located northeastward of the main body of islands that constitute the Solomon Archipelago (see Fig. 5). According to A. J. Nicholson, who collected the specimens listed above, _P. howensis_ is not abundant in Ontong Java. He related this circumstance to the fact that these small islands are nothing more than parts of a coral atoll used almost entirely for the production of coconuts (see Sanborn and Nicholson, 1950:329).

Specimens of _Pteropus howensis_ deposited in the U. S. National Museum agree well in most ways with the original description of the species by Troughton (1931:204-205). Slight variation in color is evident; in two specimens, the mantle, just posterior to the ears, is Ochraceous-Buff.

The relationship of this species to other kinds of _Pteropus_ known from Melanesia is not clear. Troughton (1931:204, 206) compared _P. howensis_ with _P. hypomelanus_ and _P. admiralitatum_ and found that it resembled each of them. Tate (1934:2) noted that the skull of _P. admiralitatum goweri_ was similar to that of _P. howensis_ in structure. The latter species is, however, larger (length of forearm 122 according to Troughton, 1931:205) than any subspecies of _P. admiralitatum_ (length of forearm 108-112). Also, the cheek-teeth of _P. howensis_ that I have studied are relatively larger than those of either _P. hypomelanus_ or _P. admiralitatum_. Furthermore, in _P. howensis_ there is a small but distinct cusp located medio-posteriorly on P4 (most noticeable in young individuals) that is more reduced or undeveloped in specimens of the other two species. Cheek-teeth of _P. howensis_ resemble those in a dull-colored specimen of _P. tonganus_ from Fiji Island with which I compared the specimens listed above.

Weights and crown-rump lengths of the two embryos (in an advanced stage of development) examined were 20 and 29 grams and 43 and 51 mm. (apparently these are the specimens listed by Sanborn and Nicholson, 1950:329).

=Pteropus tonganus=

_Pteropus tonganus_ has at least three subspecies, one of which has been recorded from the Solomons. The species ranges from a small island off the eastern coast of New Guinea, where there is an endemic subspecies, eastward to Tonga and the New Hebrides (Laurie and Hill, 1954:33-34). Felten (1964_a_) recently has reported on the species in the New Hebrides.

=Pteropus tonganus geddiei= MacGillivary

1860. _Pteropus geddiei_ MacGillivary, Zoologist, 18:7134, September, type from Aneitum Island, New Hebrides; 1912, Andersen, Catalogue of the Chiroptera ... British Museum, 1:189; 1931, Sanborn, Publ. Field Mus. Nat. Hist., Zool. Ser., 18:13, February 12, from Rennell Island in the Solomons.

1914. _Pteropus tonganus geddiei_, Revilliod, _in_ Sarasin and Roux, Nova Caledonia (A), 1:341; 1954, Laurie and Hill, List of land mammals of New Guinea, Celebes and adjacent islands, p. 34, June 30.

_Specimens examined._--None.

_Remarks._--_Pteropus tonganus geddiei_, as far as is known, is the widest ranging subspecies of this genus. It is the only megachiropteran in the Solomon Islands having affinities with bats to the southeast (the New Hebrides, Santa Cruz Islands, Samoan Islands and Fiji Islands) rather than with those to the west (New Guinea). The subspecies _P. tonganus geddiei_, which ranges from the Solomons to the New Hebrides (about 500 miles straight-line distance), is said to be remarkably uniform throughout its range. Sanborn (1931:14) compared color and size in specimens from the Solomon Islands and the New Hebrides and found little variation. Another subspecies, _P. t. bascilicus_ Thomas 1915, apparently closely related to _geddiei_, is known from Dampier [= Kar-kar] Island off the northeastern coast of New Guinea and therefore farther westward from the New Hebrides than are the Solomon Islands. Additional remarks on the distribution of this species are in the section on Zoogeography and Speciation.

=Pteropus rayneri=

_Pteropus rayneri_ is endemic to the Solomon Islands. It is divisible into seven subspecies (see Fig. 6), which, excepting _P. r. rennelli_ and _P. r. cognatus_, are strikingly colored--the mantle, back, and rump being of different colors. Differences in color and size provide characters differentiating the subspecies (see key, p. 793). Recorded lengths of forearms do not overlap between any two subspecies. _P. r. grandis_, northernmost in distribution, has the longest (about 170) forearm and _P. r. cognatus_, known from two of the southernmost islands, has the shortest (about 121).

Adult males of _Pteropus rayneri_ have well-developed tufts of hair on each side of the neck where a gland is located (see Andersen, 1912:259). Apparently these glands are not present in females as none were found in specimens studied by me or those reported by Sanborn (1931:16). Evidently, these glands are associated with sexual maturity in males because neither Sanborn nor I found them in subadult males.

=Pteropus rayneri rayneri= Gray

1870. _Pteropus rayneri_ (part), Gray, Catalogue of monkeys, lemurs and fruit-eating bats ... British Museum, p. 108, cotypes from Guadalcanal; 1878, Dobson, Catalogue of the Chiroptera ... British Museum, p. 33; 1879, Trouessart, Rev. Mag. Zool., 6:204; 1879, Trouessart, Ann. Sci. Nat. Zool, 8:16; 1887, Thomas, Proc. Zool. Soc. London, p. 322, March 15; 1888, Thomas, Proc. Zool. Soc. London, p. 472, December 4; 1898, Trouessart, Catalogus Mammalium ..., 1:78; 1912, Andersen, Catalogue of the Chiroptera ... British Museum, p. 254; 1931, Sanborn, Publ. Field Mus. Nat. Hist., Zool. Ser., 18:15, February 12, from Guadalcanal and Malaita.

1954. _Pteropus rayneri rayneri_, Laurie and Hill, List of land mammals of New Guinea, Celebes and adjacent islands, p. 35, June 30.

1899. _Pteropus (Spectrum) rayneri_ (part), Matschie, Die Megachiroptera ... naturkunde, p. 22; 1904, Trouessart, Catalogus Mammalium ..., Suppl., p. 51.

_Specimens examined_ (four males and one female; one embryo in alcohol).--Guadalcanal in July and November, USNM 278700-02, USNM 278142, USNM 278714.

_Measurements._--Measurements of three males and one female are, respectively, as follows: Length of head and body, --, 210, 214, 215; hind foot, --, 33, 39, 42; ear, --, 23, 23, 23; length of forearm, --, 138, 136, 134; greatest length of skull, 61.5, 59.2, 61.6, 61.2; condylobasal length, 61.4, 58.2, 60.3, 60.0; zygomatic breadth, 36.6, 35.3, 35.4, 36.5; breadth of braincase, 23.7, 22.5, 22.6, 24.1; breadth across first upper molars, --, 16.9, 16.7, 16.8; width of M1, 3.4, 3.5, 3.5, 3.5; length of maxillary tooth-row, 22.4, 22.1, 23.6, 23.2; length of mandibular tooth-row, 26.4, 25.5, 25.9, 25.6.

_Remarks._--_Pteropus rayneri_ was named on the basis of two specimens (cotypes) obtained on Guadalcanal and listed as "male" and "female"; according to Andersen (1912:254), however, both are females.

_P. r. rayneri_ is known from Guadalcanal and Malaita (see Fig. 6), and is of almost the same size as _P. r. cognatus_, which is known from San Cristobal and Ugi, only about 40 miles to the southeast. In the latter subspecies the back and rump are the same color (Prouts Brown), whereas in _P. r. rayneri_ the rump is brightly colored and therefore contrasts strongly with the dark brown back. A specimen of _rayneri_ from Malaita was reported by Sanborn (1931:15) as unusually small and having a dark-colored rump patch. In the specimens examined from Guadalcanal, there is noticeable variation in color of the mantle that does not seem related to age or sex. In two specimens (adult male and female) the mantle is Cinnamon-Rufous tinged with Russet, strongly contrasting with the crown, which is Ochraceous-Tawny and has scattered silvery hairs. Another specimen has a darker mantle (near Chestnut-Brown) and a crown of about the same color, but with a few scattered Ochraceous-Tawny hairs.

The skull of one adult male bears an extra peglike tooth posterior to M3 on the right side.

An embryo, in an advanced stage of development, in the collection of the U. S. National Museum, measures: Length of head and body, 98; hind foot, 30; ear, 8.5; length of forearm, 48 (this may be the same specimen listed by Sanborn and Nicholson, 1950:329).

=Pteropus rayneri grandis= Thomas

1887. _Pteropus grandis_ Thomas, Ann. Mag. Nat. Hist., ser. 5, 19:147, March, type from Shortland; 1887, Thomas, Proc. Zool. Soc. London, p. 320, March 15, from Alu and Shortland; 1897, Trouessart, Catalogus Mammalium ..., 1:80, from "I. Salomonis"; 1899, Matschie, Die Megachiroptera ... naturkunde, p. 15; 1904, Trouessart, Catalogus Mammalium ..., Suppl., p. 49; 1907, Miller, Bull. U. S. Nat. Mus., 57:58, June 29; 1912, Andersen, Catalogue of the Chiroptera ... British Museum, 1:259, from Bougainville; 1931, Sanborn, Publ. Field Mus. Nat. Hist., Zool. Ser., 18:16, February 12, from Choiseul, and Santa Ysabel; 1936, Troughton, Rec. Australian Mus., 19:348, April 7; 1953, Pohle, Z. Säugetierk., 17:128, October 27.

1954. _Pteropus rayneri grandis_, Laurie and Hill, List of land mammals of New Guinea, Celebes and adjacent islands, p. 35, June 30.

_Specimens examined_ (six males and 10 females; five in alcohol).--Choiseul in March, 23580, 23644, 23593; Bougainville in July, August, September, and October, USNM 276926-7, USNM 276968, USNM 277091-9.

_Measurements._--Average and extreme measurements of four males and seven females are as follows: Length of head and body, 281 (260-302); hind foot, 52.3 (50-58); ear, 33.1 (31-37); length of forearm, 173 (168-180). Average and extreme measurements of skulls of three males and six females are as follows: Greatest length of skull, 73.7 (71.3-77.7); condylobasal length, 73.1 (70.5-77.4); zygomatic breadth, 40 (36.4-41.5); breadth across first-upper molars, 20.9 (18.3-22.1); length of maxillary tooth-row, 28.1 (26.9-29.9); length of mandibular tooth-row, 31.8 (29.7-32.7).

_Remarks._--_Pteropus rayneri grandis_ is the largest subspecies of the species. It is also the widest ranging subspecies, being found on six islands (see Fig. 6).

Although the specimens listed above agree well with descriptions of color given by Thomas (1887_a_:147) and Andersen (1912:259, 263-264), some individual variation is noticeable. In bats not yet fully grown (judging from small size, unfused epiphyses, and lack of wear on teeth), numerous scattered hairs on the sides of the face and crown are buffy. In adults the face and crown are blackish. With regard to individual variation in color of mantle and rump patch, specimens with the following combinations were noted (1) mantle Brick Red, rump patch bright, basal three-quarters of hairs white, tips Warm Buff (2) mantle darker, near Hessian Brown, rump patch dark, Chestnut along edges, center Ochraceous-Tawny (3) mantle Brick Red, rump patch intermediate between the two other types. Size of rump patch also is variable. In some specimens it extends onto the upper parts of the thighs whereas in other specimens it does not.

Sanborn (1931:16) reported an extra tooth, behind the last lower molar, in a specimen from Choiseul. In one of three specimens in the Bishop Museum, m3 is lacking. Judging from Troughton's (1936:346) remarks, size of individuals varies considerably. Specimens that he examined from Bougainville had longer forearms (up to 177) and larger hind feet (54-57) than those examined by me from Choiseul. On the other hand, specimens listed above from Bougainville agree well with those from Choiseul. In many specimens in the U. S. National Museum, length of the right- and left-forearm differ. For example, in No. 276926 the right forearm measures 180 whereas the left is 174; in No. 277098 the right is 172 and the left is 167. Troughton (1936:346) gave standard ear measurement in _P. r. grandis_ as ranging from 29.5 to 31.5. Ears of specimens that I examined varied from 31.0 to 37.0.

=Pteropus rayneri rubianus= Andersen

1908. _Pteropus rubianus_ Andersen, Ann. Mag. Nat. Hist., ser. 8, 2:366, October, type from Rubiana; 1912, Andersen, Catalogue of the Chiroptera ... British Museum, 1:255; 1931, Sanborn, Publ. Field Mus. Nat. Hist., Zool. Ser., 18:15, February 12, from Narovo (Simbo).

1954. _Pteropus rayneri rubianus_, Laurie and Hill, List of land mammals of New Guinea, Celebes and adjacent islands, p. 35, June 30.

1888. _Pteropus grandis_ (part), Thomas, Proc. Zool. Soc. London, p. 470, December 4, from Rubiana; 1899, Matschie, Die Megachiroptera ... naturkunde, p. 15; 1904, Trouessart, Catalogus Mammalium ..., Suppl., p. 49.

_Specimens examined_ (two males and one female).--Kolombangara, in February, 23458-60.

_Measurements._--Measurements of two males and one female are, respectively, as follows: Length of head and body, 253, 265, 251; hind foot, 53, 50, 50; ear, 30, 31, 32; length of forearm, 158, 161, 160; greatest length of skull, 70.2, 67.4, --; condylobasal length, 67.0, --, 68.4; zygomatic breadth, 40.0, 39.4, 40.7; breadth across first upper molars, 19.4, 20.4, 19.9; length of mandible, 53.9, 49.4, 51.3.

_Remarks._--Kolombangara Island is a new locality for _Pteropus rayneri rubianus_; heretofore this subspecies was known only from Rubiana and Narovo islands (Andersen, 1908:366; Sanborn, 1931:15). The coloration of a specimen from Narovo Island was described as between that of _P. r. rubianus_ and _P. r. lavellanus_. Sanborn (1931:16) allocated it to the subspecies _rubianus_ on the basis of length of forearm.

Andersen's descriptions (1908:366; 1912:256) of _rubianus_ were of a specimen stored in alcohol. Coloration of the museum skins examined by me is as follows: Dorsum from shoulders to rump near Vandyke Brown; crown and mantle Brick Red; face close to Mummy Brown; rump patch and thighs close to Warm Buff, strongly contrasting with back and mantle; base of hairs dark, Seal Brown; venter dark; chest about same as back but paler laterally (to Ochraceous Tawny); throat Brick Red.

=Pteropus rayneri lavellanus= Andersen

1908. _Pteropus lavellanus_ Andersen, Ann. Mag. Nat. Hist., ser. 8, 2:366, October, type from Vella Lavella; 1912, Andersen, Catalogue of the Chiroptera ... British Museum, 1:259; 1931, Sanborn, Publ. Field Mus. Nat. Hist., Zool. Ser., 18:16, February 12, from Ghizo and Ronongo.

1954. _Pteropus rayneri lavellanus_, Laurie and Hill, List of land mammals of New Guinea, Celebes and adjacent islands, p. 36, June 30.

_Specimens examined_ (one male and one female).--Vella Lavella in November, 23192, 23142.

_Measurements._--Measurements of a male and a female are, respectively, as follows: Length of head and body, 286, 282; hind foot, 55, 56; ear, 30, 30; length of forearm, 156, 155; greatest length of skull, 72.9, 67.6; condylobasal length, 71.8, 64.2; zygomatic breadth, 38.4, 37.9; breadth across first upper molars, 19.9, 19.8; length of mandible, 54.6, 50.8.

_Remarks._--_Pteropus rayneri lavellanus_ inhabits islands geographically near those from which _P. r. rubianus_ is known (see Fig. 6) and in most respects the two subspecies closely resemble each other. _P. r. lavellanus_ is slightly the smaller (average length of forearm about 156 instead of 160) and darker. A bat from Narovo [Simbo] Island, only a few miles from Vella Lavella, identified by Sanborn (1931:16) on basis of its size as _P. r. rubianus_, resembled the subspecies _lavellanus_ in color and probably represents an intergrade between the two populations.

The color of _P. r. lavellanus_ is close to that of _P. r. rubianus_ except that the crown, mantle, and foreneck are near Chestnut-Brown, the basal portions of hair black, and the fur of the venter, from sternum to pectoral region, is dark, almost black (compare with description of _P. r. rubianus_ under account of that subspecies).

Measurements of the male examined are greater than those of the female studied. Andersen (1912:259) noted that the canine teeth are heavier in males than in females.

=Pteropus rayneri monoensis= Lawrence

1945. _Pteropus rayneri monoensis_ Lawrence, Proc. New England Zool. Club, 23:63, March 26, type from Mono (Treasury); 1954, Laurie and Hill, List of land mammals of New Guinea, Celebes and adjacent islands, p. 36, June 30.

_Specimens examined._--None.

_Remarks._--_Pteropus rayneri monoensis_ is the most recently described subspecies of _P. rayneri_. Lawrence (1945:63) judged that in most ways this bat is intermediate between _P. r. grandis_ and _P. r. lavellanus_. Coloration of _monoensis_ indicates affinity with the former, whereas length of forearm (145-148) approaches that in the latter. The small skull, narrow palate, and whitish rump patch of _monoensis_ are differences that distinguish it from _grandis_ and _lavellanus_. The relatively isolated position of Mono Island may have been important in establishment of the distinctive features of this bat.

Lawrence (1945:65) quoted a collector as stating: "They [individuals of _P. r. monoensis_] rest quietly during the day in the tops of heavy-leaved, tall jungle trees, and start flying about dusk, looking for feeding spots. There is usually quite a flight for fifteen to twenty minutes at twilight...."

No additional specimens of this subspecies have been collected on small adjacent islands and _monoensis_ may therefore be confined to Mono Island.

=Pteropus rayneri cognatus= Andersen

1908. _Pteropus cognatus_ Andersen, Ann. Mag. Nat. Hist., ser. 8, 2:365, October 1, type from San Cristobal; 1912, Andersen, Catalogue of the Chiroptera ... British Museum, 1:251; 1931, Sanborn, Publ. Field Mus. Nat. Hist., Zool. Ser., 18:15, February 12, from San Cristobal and Ugi; 1954, Laurie and Hill, List of land mammals of New Guinea, Celebes and adjacent islands, p. 35, June 30.

1962. _Pteropus rayneri cognatus_, Hill, The natural history of Rennell Island, British Solomon Islands, 4:9, February.

1870. _Pteropus rayneri_ (part), Gray, Catalogue of monkeys, lemurs and fruit-eating bats ... British Museum, p. 108, from San Cristobal; 1878, Dobson, Catalogue of the Chiroptera ... British Museum, p. 33.

1904. _Pteropus_ (_Spectrum_) _rayneri_ (part), Trouessart, Catalogus Mammalium ..., Suppl., p. 51.

_Specimens examined._--None.

_Remarks._--Specimens of _Pteropus rayneri cognatus_ first were reported under the name _Pteropus rayneri_ based on three specimens (one from San Cristobal and two from Guadalcanal). Because the description was based mostly on the two specimens from Guadalcanal, the name _rayneri_ is applicable to the bats from that island. Andersen (1908:365) thought that specimens that he studied, from San Cristobal, were specifically distinct from _P. rayneri_ and he proposed the name _Pteropus cognatus_ for them. Later, Hill (1962:9) reduced _cognatus_ to subspecific status under _P. rayneri_.

Presently _P. r. cognatus_ is known only from San Cristobal and the small adjacent island of Ugi (see Fig. 6).

=Pteropus rayneri rennelli= Troughton

1929. _Pteropus rennelli_ Troughton, Rec. Australian Mus., 17:193, September 4, type from Rennell Island; 1954, Laurie and Hill, List of land mammals of New Guinea, Celebes and adjacent islands, p. 35, June 30.

1962. _Pteropus rayneri rennelli_, Hill, The natural history of Rennell Island, British Solomon Islands, 4:7, February.

_Specimens examined._--None.

_Remarks._--Until recently, _Pteropus rayneri rennelli_ was known from but a single specimen. Hill (1962:7) reported two additional specimens and pointed out that _P. r. cognatus_ and _P. r. rennelli_ probably represent the extremes of an east-west cline in size. _P. r. rennelli_ and _P. r. cognatus_ differ from other subspecies of the species in lacking tricolored pelage on the dorsum, but their short rostrum clearly indicates affinity with other members of this complex group in the Solomon Islands (Hill, 1962:8).

The relationship of the subspecies _rennelli_ and _cognatus_ is close, both geographically and genetically. Longer forearm, longer metacarpals, and longer mandibular tooth-row serve to differentiate _rennelli_ from _cognatus_.

=Pteropus woodfordi= Thomas

1888. _Pteropus woodfordi_ Thomas, Ann. Mag. Nat. Hist., ser. 6, 1:156, February, type from Guadalcanal; 1888, Thomas, Proc. Zool. Soc. London, p. 472, December 4; 1898, Trouessart, Catalogus Mammalium ..., 1:78; 1907, Elliot, Field Columbian Mus., Zool. Ser., 8:491; 1912, Andersen, Catalogue of the Chiroptera ... British Museum, 1:410, from New Georgia and Guadalcanal; 1931, Sanborn, Publ. Field Mus. Nat. Hist., Zool. Ser., 18:19, February 12, from Kolombangara; 1947, Sanborn and Beecher, Jour. Mamm., 28:389, November 19, from Banika and Guadalcanal; 1954, Laurie and Hill, List of land mammals of New Guinea, Celebes and adjacent islands, p. 39, June 30.

1899. _Pteropus (Sericonycteris) woodfordi_, Matschie, Die Megachiroptera ... naturkunde, p. 83; 1904, Trouessart, Catalogus Mammalium ..., Suppl., p. 54.

1945. _Pteropus austini_ Lawrence, Proc. New England Zool. Club, 23:59, March 26, from Florida.

_Specimens examined_ (four males and three females; five in alcohol and two skin-onlys).--Fauro, in April, 23727, 23790; Guadalcanal in May and June, 23823, 23931; Pavuvo (Russell Islands) in August and October, USNM 277887, USNM 283872-3.