Chapter II: Part 2

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

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

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

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

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

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

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

=Pteropus rayneri monoensis= Lawrence

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

_Specimens examined._--None.

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

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

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

=Pteropus rayneri cognatus= Andersen

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

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

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

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

_Specimens examined._--None.

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

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

=Pteropus rayneri rennelli= Troughton

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

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

_Specimens examined._--None.

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

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

=Pteropus woodfordi= Thomas

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

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

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

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

_Measurements._--External measurements of two males and two females are, respectively, as follows: Length of head and body, 152, 128, 132, 155; hind foot, 29, 26, 31, 28; ear, 16, 14, 14, 17; length of forearm, 79, 76, 86, 90.

_Remarks._--Heretofore, _Pteropus woodfordi_ was known from New Georgia, Guadalcanal, Kolombangara, and Banika (see Fig. 7); specimens from Fauro and Pavuvo islands, listed above, provide new northern localities of record for this species.

Judging by small size and unfused epiphyses, a bat obtained in April and another obtained in June are subadults. Specimens of adults, examined by me, agree well with the descriptions of _P. woodfordi_ by Thomas (1888_a_:156) and Andersen (1912:407-409), but are slightly smaller than specimens listed by Sanborn and Beecher (1947:389). Color of pelage in this species seems to vary. Adults seen have a pale head and mantle, contrasting strongly with the dark back. Andersen (1912:409) and Lawrence (1945:61) discussed individuals that had scattered silvery hairs mixed with dark fur dorsally and darker mantles that did not contrast noticeably with the rest of the dorsum.

Lawrence (1945:389) named _Pteropus austini_ as a new species closely related to _P. woodfordi_ and other species of the _P. scapulatus_ group of Andersen (1912:402) and Tate (1942:336). Sanborn and Beecher (1947:389), studied a series of _P. woodfordi_ from Banika and Guadalcanal and found that skulls of two subadults agreed well with cranial characteristics ascribed to _P. austini_, which was based on two subadults. Lawrence (1945:61) stated also that "the interfemoral membrane is entirely absent medially in _austini_, while in _woodfordi_ it is present as a barely discernible ridge 8 mm. wide." Andersen (1912:408) had earlier reported that in the type of _woodfordi_ the interfemoral membrane was "undeveloped in [the] centre." In 13 adults (in alcohol) studied by Sanborn and Beecher (1947:389), as well as in adults examined by me, the uropatagium is not present. In size, however, these specimens agree with dimensions given for _woodfordi_ by Thomas (1888_a_:156) and Andersen (1912:410); for example, length of forearm is 93-99. According to Lawrence (1945:59) _austini_, in which the interfemoral membrane is lacking, is smaller than _woodfordi_ and has a forearm of about 84. In two juveniles of _P. woodfordi_ in the U. S. National Museum, the medially-developed interfemoral membrane is about 7 wide. One specimen has small but distinct calcars whereas the other (slightly larger) apparently lacks calcars. This suggests individual variation in the presence or absence, as well as in the size, of the uropatagium in _Pteropus woodfordi_.

Sanborn and Beecher (1947:389) decided that "until fully adult specimens showing the characters of _austini_ are available, it best be considered a synonym of _woodfordi_." For the following reasons I agree with these authors: (1) _austini_ is known from only two specimens, both of which are apparently subadults; (2) _austini_ is reported to have a forearm 84 long and no interfemoral membrane, whereas _woodfordi_ has a forearm about 96 long and an interfemoral membrane that is only slightly developed; (3) specimens that agree in size and cranial characters with the type of _woodfordi_ but that lack an interfemoral membrane have been obtained; and (4) skulls of subadults of _woodfordi_ agree with the description of skulls of _austini_.

Sanborn (1931:19) reported that specimens of _Pteropus woodfordi_ were obtained at night, while feeding on young green coconuts. Lawrence (1945:62) reported that in the late afternoon a collector found individuals of _austini_ [= _woodfordi_] in the fronds of a coconut tree, apparently feeding on pollen shoots. Sanborn and Beecher (1947:388) have reported malaria (_Plasmodium_) in _P. woodfordi_ obtained on Guadalcanal. They suggested that malaria might have rendered one individual helpless because when it was found, on the ground, no wounds were evident and parasites were present in the blood.

)
and _P. mahaganus_ ([BC]). For names of islands see Fig. 2.]

=Pteropus mahaganus= Sanborn

1931. _Pteropus mahaganus_ Sanborn, Publ. Field Mus. Nat. Hist.,
Zool. Ser., 18:19, February 12, type from Santa Ysabel, also
reported from Bougainville; 1954, Laurie and Hill, List of
land mammals of New Guinea, Celebes and adjacent islands,
p. 39, June 30.

_Specimens examined_ (one male and two females; one in alcohol).--Bougainville, in August and October, USNM 276972, USNM 277104-5.

_Measurements._--Measurements of one male and two females are, respectively, as follows: Length of head and body, 180, 204, 198; hind foot, 42, 38, 44; ear, 25, 23, 22; length of forearm, 134, 138, 140. Measurements of the skull of the male and one female are, respectively, as follows: Greatest length of skull, 52.5, 55.8; condylobasal length, 50.9, 54.3; palatal length, 24.1, 26.0; zygomatic breadth, 28.9, 32.5; breadth across first upper molars, 14.4, 15.0; width of M1, 2.2, 2.2; length of maxillary tooth-row, 17.4, 18.4; length of mandibular tooth-row, 20.1, 21.4.

_Remarks._--Sanborn (1931:19-21) described _Pteropus mahaganus_ on basis of six specimens, five from Santa Ysabel and one from Bougainville. The latter was in poor condition and only provisionally allocated to this species. The specimens examined by me (listed above) confirm the occurrence of _P. mahaganus_ on Bougainville.

Sanborn (1931:20) described _mahaganus_ as "similar to and about the size of [_Pteropus scapulatus_] from Australia, but lighter in color," and considered it, along with _P. woodfordi_, a member of the _Pteropus scapulatus_ group of Andersen (1912:402) and Tate (1942:336). I would judge, however, that _P. mahaganus_ and _P. woodfordi_ are much more closely related to one another than to _P. scapulatus_ of Australia. The only significant characteristic that the latter has in common with the two species from the Solomons is small cheek-teeth. In fact, teeth of _scapulatus_ are relatively smaller than teeth of either _mahaganus_ or _woodfordi_. Also, in _scapulatus_ the upper canines are widely separated due to lateral expansion of the palate at that point, whereas in _mahaganus_ and _woodfordi_ the width across the upper canines is relatively much less.

=Dobsonia= Palmer

1898. _Dobsonia_ Palmer, Proc. Biol. Soc. Washington, 12:114,
April 30.

1810. _Cephalotes_ (part) É. Geoffroy, Ann. du Mus. d'Hist. Nat.,
15:104.

_Dobsonia_, a genus of large to medium-sized fruit bats, occurring from Celebes to the Solomon Islands, contains at least nine species. One species and its two subspecies are endemic to the Solomons.

_Dobsonia_ differs from all other genera of megachiropteran bats in the Solomons by combining absence of a small claw on the second digit and presence of external tail vertebrae.

The cranium of _Dobsonia_ resembles, in some ways, the cranium of _Rousettus_ as well as that of _Pteropus_. Even so, in _Dobsonia_ the rostrum is shorter and the cheek-teeth, especially in the upper jaw, are more crowded. The anterior part of the mandible is narrow and the lower incisors are diminutive and often concealed by the flesh of the gum.

=Dobsonia inermis=

In a review of the genus _Dobsonia_, Andersen (1909_c_:532) named and described _D. inermis_ and _D. nesea_ from the Solomons. Specimens of _Dobsonia inermis_ from San Cristobal and Ugi were said to differ from specimens of _D. nesea_ from Alu, Shortland, and Rubiana in having perpendicular as opposed to anteriorly slanted upper canines. Andersen (1909_c_:532) reported that the two species were of "... the same general size." Troughton (1936:348-349) studied specimens of _Dobsonia_ from Bougainville and Santa Ysabel and, because of individual variation in proclivity of the upper canines, concluded that _D. nesea_ was conspecific with _D. inermis_. He (p. 349) noted that the ears were shorter in _inermis_ than in _nesea_, but the size of teeth showed insular variation and a "... confusing amount of intergradation ... [that obscures] ... diagnostic importance."

Specimens of _Dobsonia_ from Choiseul are smaller (externally and cranially) than those from Alu, Shortland, Rubiana, Bougainville, Fauro, Vella Lavella, Guadalcanal, Florida, Ugi, San Cristobal, and Rennell. Specimens from Santa Ysabel (see Fig. 8) are intermediate in size between those from Choiseul and the other islands listed. Judging from available specimens, two subspecies of _Dobsonia inermis_ occur in the Solomons. Specimens from Choiseul (see A, Fig. 8), which are smaller than those from other islands, represent one subspecies (heretofore unrecognized), whereas specimens from other islands (except Santa Ysabel) represent a second subspecies. Specimens from Santa Ysabel are slightly larger than those on Choiseul and are regarded as intergrades between the two subspecies. Specimens from Rennell, Ugi, San Cristobal, Florida, Fauro, and Guadalcanal are slightly smaller than those from Bougainville, Vella Lavella, Shortland, and Rubiana, but the differences are not great enough to warrant recognition of two subspecies. Therefore, the subspecific name _nesea_ is arranged as a synonym of _inermis_, which has priority, and the latter name is used for specimens of _Dobsonia inermis_ from the Solomon islands other than Choiseul and Santa Ysabel. Additional remarks on the distribution of this species are in the section on Zoogeography and Speciation.

Pohle (1953:130) suggested that _Dobsonia inermis_ (as well as _D. crenulata_ and _D. praedatrix_) is conspecific with _D. viridis_, but Laurie and Hill (1954:41) did not adopt his suggestion. I have not seen adequate series of _crenulata_, _praedatrix_, and _viridis_ (none of which occurs in the Solomons) to judge systematic relationships of these kinds; therefore I follow Laurie and Hill.

=Dobsonia inermis inermis= Andersen

1909. _Dobsonia inermis_ Andersen, Ann. Mag. Nat. Hist., ser. 8,
4:532, December, type from San Cristobal; 1912, Andersen,
Catalogue of the Chiroptera ... British Museum, 1:475.

1936. _Dobsonia inermis inermis_, Troughton, Rec. Australian Mus.,
14:349, April 7, from Santa Ysabel; 1954, Laurie and Hill, List
of land mammals of New Guinea, Celebes and adjacent islands,
p. 41, June 30; 1956, Hill, The natural history of Rennell
Island, British Solomon Islands, 1:74, November 28, from
Rennell Island.

1878. _Cephalotes peroni_ (part), Dobson, Catalogue of the
Chiroptera ... British Museum, p. 91; 1879, Trouessart, Rev.
Mag. Zool., 3:208; 1887, Thomas, Proc. Zool. Soc. London,
p. 323, March 15, from Ugi and San Cristobal; 1888, Thomas,
Proc. Zool. Soc. London, p. 476, December 4; 1897, Trouessart,
Catalogus Mammalium ..., 1:87.

1899. _Dobsonia peroni_ (part), Trouessart, Catalogus Mammalium
..., 2:1278.

1909. _Dobsonia nesea_ Andersen, Ann. Mag. Nat. Hist., ser. 8,
4:532, December 1, type from Shortland Island; 1912, Andersen,
Catalogue of the Chiroptera ... British Museum, 1:476, from
Shortland and Rubiana; 1931, Sanborn, Publ. Field Mus. Nat.
Hist., Zool. Ser., 18:22, February 12, from San Cristobal.

1936. _Dobsonia inermis nesea_, Troughton, Rec. Australian Mus.,
14:348, April 7, from Bougainville; 1953, Pohle, Z. Säugetierk.,
17:130, October 27; 1954, Laurie and Hill, List of land mammals
of New Guinea, Celebes and adjacent islands, p. 41, June 30,
from New Georgia.

_Specimens examined_ (13 males and three females; three in alcohol, crania extracted and cleaned).--Fauro in April, 23728, 23740, 23751; Vella Lavella in November, 23134-36, 23141, 23145, 23147, 23149, 23151, 23153; Guadalcanal in May and June, 23865, 23914, 24008; Florida in October, 24416.

_Measurements._--See tables 2 and 3.

_Remarks._--Heretofore, _Dobsonia inermis inermis_ was unreported from Fauro, Vella Lavella, Guadalcanal, and Florida. Apparently the subspecies occurs on most islands of the archipelago (see Fig. 9).

In coloration and most cranial dimensions the specimens listed above agree with specimens of _D. i. inermis_ from Alu, Shortland, and Rubiana (Andersen, 1909_c_:532; 1912:475, 476), Bougainville (Troughton, 1936:348, 349), and Rennell (Hill, 1963:74). The forearm in the adult male holotype of "_nesea_," from Shortland, is 109.5 as opposed to 109.0 in an adult female topotype of _inermis_ from Ugi (Andersen, 1912:478) in the southeastern part of the archipelago (see Fig. 9). Forearms of specimens of _D. i. inermis_ from Vella Lavella are 107 to 112 (measurements from labels because forearms of these specimens were broken and therefore could not be remeasured). Forearms of specimens from Fauro, Florida, Guadalcanal, and Rennell are 103.6 to 110.0 (see Hill, 1956:74). Variation in length of forearm probably is not significant because no cline is evident (see Fig. 9 and Table 2).

In 1964, 11 specimens of _Dobsonia inermis_ were collected on Choiseul. They are smaller, externally and cranially, than specimens of _D. inermis_ from Sun Cristobal, Ugi, Rennell, Guadalcanal, Florida, Rubiana, Vella Lavella, Shortland, Alu, Bougainville, and Fauro, and may be named and described as follows:

=Dobsonia inermis minimus=, new subspecies

_Type._--Adult male skin and skull, in good condition (originally stored in 70 per cent alcohol for about one year), no. BBM-BSIP 23716, Bernice P. Bishop Museum; from Choiseul Island, British Solomon Islands Protectorate; obtained on 20 March 1964 by Philip Temple, original number 1524.

_Distribution._--Choiseul Island (type locality); intergrades from Santa Ysabel also assigned to this subspecies.

_Diagnosis._--Size small for species; wing membranes, feet, and ears black; dorsal surface of interfemoral membrane sparsely set with silvery hairs, other membranes naked; hair soft, medium length (10 on mantle, 5 on crown), black hairs and scattered white hairs on face and crown; fur of dorsal surface of mantle composed of whitish hairs having faint olive cast imparting general color of Buffy-Citrine; hair of venter short (about 5), soft, and fine; general coloration Buffy-Citrine; cranium delicate; rostrum narrow in dorsal aspect (nasals not expanded laterally); forehead (junction of nasals and frontals) pronounced in lateral aspect; teeth resembling those of other subspecies of _D. inermis_ but slightly smaller.

_Comparisons._--From adults of _Dobsonia inermis inermis_, which occurs on Rennell, San Cristobal, Ugi, Malaita, Florida, Guadalcanal, Rubiana, Vella Lavella, Shortland, Alu, Bougainville, and Fauro, _minimus_ differs in being smaller. Average length of mandible 31.2 and 33.4. For other measurements see Table 2.

From _Dobsonia praedatrix_, which occurs on New Britain, New Ireland, and Duke of York (northward of the Solomons), _minimus_ differs in being smaller in all dimensions; length of forearm averaging 100.5 as opposed to 116.0, and greatest length of skull 42.4 as opposed to 50.0.

), _D. i. minimus_ ([TW]), and
intergrades assigned to _minimus_ ([BC]). Capital letters are used
to relate groups of specimens to the island or islands from which
they were collected; spatial distribution of specimens indicated in
the scatter diagram thus is shown in the inset map. Specimens from
Santa Ysabel and Bougainville are deposited in the Australian
Museum. The type specimen of _D. i. inermis_ is labeled "E." For
names of islands see Fig. 2.]

) and _D. inermis minimus_ ([RW]). For names of islands see
Fig. 2.]

TABLE 2. Average and Extreme Measurements of Two Subspecies of
_Dobsonia inermis_.

================+====================================+===================
| _D. i. minimus_ | _D. i. inermis_
+------------------+-----------------+-------------------
| | | Guadalcanal,
MEASUREMENT | Choiseul | Santa Ysabel | Fauro, Vella
| 4 [M], 2 [F] | 1 [M], 2 [F] |Lavella, Florida
| | | 9 [M], 2 [F]
----------------+------------------+-----------------+-------------------
Length of head | | |
and body |174.5 (170 -180 )| |174.5 (160 -190 )
Tail vertebrae | 28.5 (24 - 33 )| 23.0 (21.5-24.0)| 30.4 ( 25 - 35 )
Hind foot | 25.3 (25.1- 25.9)| 23.5 (23.5-24.0)| 29.3 ( 26.0- 31.6)
Ear | 21.5 (21.0- 22.9)| 21.1 (21.0-21.5)| 23.1 ( 19 - 25 )
Length of | | |
forearm |100.5 (98.1-104.0)|105.3 (104 -107 )|108.4 (105 -112 )
2nd metacarpal | 43.6 (42.6- 45.2)| | 48.1 ( 45.9- 50.9)
3rd metacarpal | 61.5 (59.8- 62.9)| | 67.6 ( 65.2- 68.5)
4th metacarpal | 57.5 (56.5- 58.5)| | 62.5 ( 58.7- 65.5)
5th metacarpal | 59.0 (57.0- 60.5)| | 64.4 ( 61.8- 66.0)
| | |
Greatest length | | |
of skull | 42.4 (42.1- 43.5)| 44.0 (43.0-45.6)| 45.9 ( 45.2- 47.4)
Condylobasal | | |
length | 40.4 (39.5- 41.3)| 41.7 (41.1-42.6)| 43.6 ( 43.1- 45.0)
Zygomatic | | |
breadth | 25.6 (24.9- 26.8)| 26.5 (25.7-27.6)| 27.9 ( 27.2- 28.5)
Breadth of | | |
braincase | 16.8 (16.5- 17.4)| 18.0 (17.1-19.9)| 17.9 ( 16.7- 19.0)
Breadth across | | |
upper canines | 8.4 ( 8.1- 8.7)| | 9.2 ( 9.2- 9.5)
Breadth across | | |
first upper | | |
molars | 12.1 (11.8- 12.6)| | 13.1 ( 12.6- 13.3)
Length of | | |
maxillary | | |
tooth-row | 15.6 (15.5- 15.8)| 16.4 (16.0-17.2)| 16.4 ( 15.9- 17.0)
Length of | | |
mandibular | | |
tooth-row | 17.1 (16.8- 17.6)| 17.8 (17.3-18.4)| 18.2 ( 17.8- 19.4)
----------------+------------------+-----------------+-------------------

_Measurements._--Comparative measurements of the subspecies _inermis_ and _minimus_ are given in Table 2. Some measurements of the type are as follows: Length of head and body, 147; tail vertebrae, 31; hind foot, 25; ear, 21; length of forearm, 99.5; 2nd metacarpal, 42.8; 3rd metacarpal, 62.7; 4th metacarpal, 58.5; 5th metacarpal, 59.1; greatest length of skull, 42.2; condylobasal length, 40.6; zygomatic breadth, 25.8; breadth of braincase, 16.8; length of maxillary tooth-row, 15.8; length of mandible, 31.2.

_Remarks._--_Dobsonia inermis minimus_ is the smallest subspecies of _Dobsonia inermis_. Specimens from Santa Ysabel, southeastward of Choiseul, are slightly larger than the type and paratypes of _minimus_. As can be seen in the scatter diagram (Fig. 8), a male from Santa Ysabel is as large as one male and most females of _D. i. inermis_. The other three specimens from Santa Ysabel also are slightly larger than specimens of _minimus_ from Choiseul, but are much smaller than specimens of _D. i. inermis_, and, therefore, are referred to _D. i. minimus_.

Although there is a cline in size of _Dobsonia inermis_ from Choiseul to Florida (generally southward; Fig. 9), no cline in size is apparent between Choiseul and Fauro (generally westward). Specimens of _D. inermis_ from Fauro are average for the subspecies _inermis_; there is no evidence, in the small series available, of intergradation between _minimus_ on Choiseul and _inermis_ on Fauro.

_Specimens examined_ (eight males and three females, all originally in alcohol; seven crania, all adults, extracted and cleaned).--Choiseul in March, 23565, 23628, 23637, 23665-67, 23640, 23714, 23716 (holotype), 23717, 23720. Ellis LeG. Troughton kindly examined and measured nos. AM-M. 3693[M], AM-M. 3694[M], AM-M. 3937[F], and AM-M. 3940[F], from Santa Ysabel in the Australian Museum.

Subfamily Macroglossinae

=Macroglossus= F. Cuvier

1824. _Macroglossus_ F. Cuvier, Des dents des mammiferes ...
zoologiques, p. 248.

1840. _Kiodotus_ Blyth, _in_ Cuvier's animal kingdom ..., p. 69.

1891. _Carponycteris_ Lydekker, _in_ Flower and Lydekker, mammals
living and extinct, p. 654.

1902. _Odontonycteris_ Jentink, Notes Leyden Mus., 23:140, July 15.

_Macroglossus_, the widest-ranging genus of macroglossine bats, occurs from southeastern Asia to the southern islands of the Solomon Archipelago (see Ellerman and Morrison-Scott, 1966:101; Laurie and Hill, 1954:44). One species, known also from Celebes and New Guinea, occurs in the Solomons and is represented there by an endemic subspecies.

Numerous generic names have been applied, at one time or another, to bats now considered as _Macroglossus_. Trouessart (1904:65) and Miller (1907:70) listed the one bat of this genus occurring in the Solomons under _Carponycteris_ and _Kiodotus_, respectively. Andersen (1911:642; 1912:767) and, later, Sanborn (1931:22) identified this bat as _Macroglossus lagochilus microtus_. Troughton (1936:350), reporting an extension of range of this species in the Solomons, used the generic name _Odontonycteris_ without explanation. Andersen (1912:754) pointed out that Jentink originally established the name _Odontonycteris_ on the basis of an extra premolar in each upper jaw as opposed to the usual two in _Macroglossus_, and arranged _Odontonycteris_ as a synonym of _Macroglossus_ because "in no genus of Megachiroptera are dental anomalies of so frequent occurrence as in _Macroglossus_, and on no point of the jaws are these anomalies ... so often met with as on that occupied by the molar series." Sanborn (1931:22) and Phillips (1966:27) noted variation in number of incisors in _Macroglossus_ as well as in _Melonycteris_, another macroglossine genus. All of the more recent workers (Ellerman and Morrison-Scott, 1966; Pohle, 1953; Laurie and Hill, 1954) use the name _Macroglossus_.

=Macroglossus lagochilus=

_Macroglossus lagochilus_ has at least three subspecies, one of which is endemic to the Solomons. The species ranges from Celebes on the west to the Solomon Islands on the east, occurring not only in New Guinea but also on many of the small adjacent islands (see Laurie and Hill, 1954:44).

=Macroglossus lagochilus microtus= Andersen

1911. _Macroglossus lagochilus microtus_ Andersen, Ann. Mag. Nat.
Hist., Ser. 8, 7:642, June, type from Guadalcanal, additional
specimens from Florida; 1912, Andersen, Catalogue of the
Chiroptera ... British Museum, 1:767; 1931, Sanborn, Publ.
Field Mus. Nat. Hist., Zool. Ser., 18:22, February 12, from San
Cristobal; 1953, Pohle, Z. Säugetierk., 17:130, October 27,
from Bougainville; 1954, Laurie and Hill, List of land mammals
of New Guinea, Celebes and adjacent islands, p. 44, June 30.

1888. _Macroglossus australis_ (part). Thomas, Proc. Zool. Soc.
London, p. 476, December 4, from Guadalcanal.

1904. _Carponycteris nana_ (part), Trouessart, Catalogus Mammalium
..., Suppl., p. 65.

1907. _Kiodotus_ sp., Miller, Bull. U. S. Nat. Mus., 57:70,
June 29.

1936. _Odontonycteris lagochilus microtus_, Troughton, Rec.
Australian Mus., 14:350, April 7, from Bougainville.

_Specimens examined_ (14 males and 16 females; in alcohol).--Choiseul in March, 23654-57, 23614, 23629, 23643, 23645, 23647, 23677-79, 23684; Vella Lavella in December, 23277-79, 23283-84; Fauro in April, 23765; Guadalcanal in May and June, 23830, 23864, 23935; Kolombangara in January, 23385, 23399, 23397, 23407, 23420-21; Santa Ysabel in June, 24067; Malaita in June, 24067.

_Measurements._--Average and extreme external measurements of 14 males and 15 females are as follows: Length of head and body, 68.3 (63-72); tail vertebrae present but scarcely perceptible and therefore not measured; hind foot, 11.4 (9.0-12.9); ear, 12.0 (10.0-12.9); length of forearm, 37.6 (36.2-39.9).

_Remarks._--The distribution of _Macroglossus lagochilus microtus_ has not been well known. Specimens herein reported from Choiseul, Fauro, and Vella Lavella provide new records of distribution. As shown on Figure 10, the subspecies occurs throughout the Solomon Islands.

_Macroglossus lagochilus microtus_ differs slightly from _M. l. nanus_ Matschie, the subspecies of the Bismarck Archipelago and Admiralty Islands to the north of the Solomons. _M. l. nanus_ averages slightly larger than _microtus_ (see Andersen, 1912:768-769, for comparative measurements) but otherwise closely resembles it.

Individual variation is evident in several measurements of the specimens at hand (in length of forearm, for example) but no clines are apparent. Four females obtained in March were lactating, as was one taken in December and one taken in January.

=Melonycteris= Dobson

1877. _Melonycteris_ Dobson, Proc. Zool. Soc. London, p. 119,
June 1.

1877. _Cheiropteruges_ Ramsay, Proc. Linn. Soc. New South Wales,
2:19, July.

1887. _Nesonycteris_ Thomas, Ann. Mag. Nat. Hist., ser. 5, 14:147,
February.

The genus _Melonycteris_ is known from three species, two apparently endemic to the Solomon Islands and the third occurring in eastern New Guinea and the Bismarck Archipelago (Laurie and Hill, 1954:45).

Heretofore, the generic name _Nesonycteris_ has been applied to the species in the Solomons, whereas _Melonycteris_ has been restricted to the one species in the Bismarck Archipelago and New Guinea. Andersen (1912:792) judged that _Nesonycteris_ was clearly distinct from _Melonycteris_ on the basis of two characters (loss of a claw on the second digit and loss of the inner, lower incisors). On the other hand, he noted striking similarities in general cranial features, dentition, palatal ridges, tongue, and external appearance of the two genera. Pohle (1953:131) synonymized the two but Laurie and Hill (1954:45) considered them distinct. I have suggested previously (Phillips, 1966:26, 27) that characteristics used to distinguish between _Melonycteris_ and _Nesonycteris_ are of less than generic value. Variability of number of incisors in the upper jaw of specimens of _Melonycteris_ (and in other macroglossine genera, as well) indicates a lack of selective pressure for either increase or decrease in number of incisors. Furthermore, the loss of the small claw on the second digit might not be important because, as Bader and Hall (1960:15) have pointed out, limbs of bats vary more in phenotypic expression than do other parts of the skeletal structure.

The discovery of a new species (_Melonycteris aurantius_) in the Solomon Islands sheds additional light on the problem. Although _M. aurantius_ possesses the distinguishing characteristics of the genus "Nesonycteris," the species closely resembles _Melonycteris_ in other features. Similarity in structure of hair of _Melonycteris_ and _Nesonycteris_, as first reported by Benedict (1957:293), also supports the argument for synonymy (see Phillips, 1966:26).

_Melonycteris aurantius_ lacks a small claw on the second digit and has only two lower incisors. In these ways this species is like _woodfordi_, which also is restricted to the Solomons. On the other hand, the structure of the skull of _M. aurantius_ is like that of _M. melanops_, which is the species found in the Bismarck Archipelago.

Although _melanops_ is not yet known from the Solomon Islands, I have included it in the following key.

Key to Known Species of _Melonycteris_

1. Ventral surface darker than dorsum, but not strongly
contrasting with it; lacking a small claw on the
second digit, 2

1´. Ventral surface nearly black, strongly contrasting with
dorsum; small claw on second digit,
=Melonycteris melanops=

2(1´). Pelage bright, Cinnamon-Rufous; postorbital region of
skull expanded (about 8.3 wide),
=Melonycteris aurantius=, p. 816

2´. Pelage dark, near Wood-Brown or Cinnamon; postorbital
region of skull constricted (about 7.5),
=Melonycteris woodfordi=, p. 816

and _M. woodfordi_ [LW]. For names of islands see Fig. 2.]

=Melonycteris aurantius= Phillips

1966. _Melonycteris aurantius_ Phillips, Jour. Mamm., 47:23-27,
March 12, type from Florida Island, additional specimens from
Choiseul Island.

_Specimens examined_ (six females; three in alcohol).--Florida in October, 24440; Choiseul in March, 23615, 23617, 23558, 23694, 23681.

_Measurements._--Average and extreme measurements of six females are as follows: Length of head and body, 80.8 (77-106); hind foot, 17.2 (16.0-18.7); ear, 12.7 (11.5-14.0); length of forearm, 49.3 (42.9-53.8). Average and extreme measurements of skulls of five females are as follows: Greatest length of skull, 31.8 (30.8-33.3); condylobasal length, 29.7 (28.6-32.4); zygomatic breadth, 18 (17.2-20.0); breadth of braincase, 12.6 (12.4-13.2); postorbital breadth, 8.3 (8.0-8.9); length of maxillary tooth-row, 10.1 (9.4-10.4); length of mandibular tooth-row, 11.7 (10.8-12.2).

_Remarks._--On Choiseul Island _Melonycteris aurantius_ was taken at the same locality as its congener, _Melonycteris woodfordi_.

Externally, _M. aurantius_ resembles _M. woodfordi_. These species are the same size, but the former is brighter in color (nearly orange in adults) than the latter, which is Wood-Brown dorsally. Internally, differences between _M. aurantius_ and _M. woodfordi_ are more obvious. In the skull of _M. aurantius_, the postorbital region is expanded (measuring about 8.3), whereas in _M. woodfordi_ the postorbital region is constricted. Furthermore, in lateral aspect the posterior portion of the skull of _M. aurantius_ is down-turned and the angle of the facial axis with the basicranial axis is much more acute than in _M. woodfordi_.

The number of upper incisors is highly variable in the six specimens of _M. aurantius_ that I have examined. In two specimens an extra tooth has erupted just anterior to I2 and there is a total of six upper incisors. In two other specimens an extra tooth has erupted in front of I2 on one side but not the other. I could find no trace of an extra tooth in the remaining two specimens.

Practically nothing is known about the natural history of _M. aurantius_, or, indeed, that of either of the other two species of this genus. One field collector (Temple, _in litt._) for the Bishop Museum reported that he obtained both _M. aurantius_ and _M. woodfordi_ in the same mist net in one night. The holotype, an adult female, was lactating when obtained in October.

=Melonycteris woodfordi= (Thomas)

1887. _Nesonycteris woodfordi_ Thomas, Ann. Mag. Nat. Hist., ser.
5, 14:147, February, type from Shortland Island; 1887, Thomas,
Proc. Zool. Soc. London, p. 324, March 15; 1888, Thomas, Proc.
Zool. Soc. London, p. 476, December 4; 1898, Trouessart,
Catalogus Mammalium ..., 1:90; 1899, Matschie, Die Megachiroptera
... naturkunde, p. 91; 1904, Trouessart, Catalogus Mammalium ..,
Suppl., p. 66; 1907, Miller, Bull. U. S. Nat. Mus., 57:74,
June 29; 1912, Andersen, Catalogue of the Chiroptera ... British
Museum, 1:792, from Alu, Shortland, Fauro, and Guadalcanal; 1931,
Sanborn, Publ. Field Mus. Nat. Hist., Zool. Ser., 18:23, February
12, from Russell Island (Pavuvo); 1954, Laurie and Hill, List of
land mammals of New Guinea, Celebes and adjacent islands, p. 45,
June 30.

1953. _Melonycteris woodfordi_, Pohle, Z. Säugetierk., 17:130,
October 27, from Bougainville Island; 1966, Phillips, Jour. Mamm.,
47:23, March 12, from Choiseul.

_Specimens examined_ (three males and one female; in alcohol).--Choiseul, in April, 23413-14, 23434, 23275.

_Measurements._--Average and extreme measurements of three males and one female are as follows: Length of head and body, 86.1 (83.1-91.0); hind foot, 19.6 (17.2-22.2); ear, 11.3 (10.8-11.7); length of forearm, 54.4 (52.1-57.7).

_Remarks._--Specimens of _Melonycteris woodfordi_ from Choiseul constitute a new locality of occurrence for the species. Apparently _M. woodfordi_ occurs throughout the Solomons (see Fig. 11).

Thomas (1887_a_:147) named _Nesonycteris woodfordi_ in a preliminary report that appeared before the publication of the more detailed description of the genus and species (1887_b_:323-324). In the second paper he stated that the anterior projections of the premaxillary bones are separated distinctly in both _Nesonycteris_ and _Melonycteris_. According to Thomas (1887_b_:323), it was by some "accident" that Dobson (1878:4) reported the anterior projections of the premaxillary bones in _Melonycteris melanops_ to be united. Writing at a later date, Andersen (1912:785) reported that in _Melonycteris melanops_ the premaxillary bones have "simple contact with each other." Furthermore, in Andersen's (1912:791) illustration of _M. woodfordi_ the premaxillary bones are in contact anteriorly. In specimens of _woodfordi_ and _melanops_ examined by me, the premaxillary bones are in contact. In _M. aurantius_ the premaxillary bones are not in contact, and it differs from _woodfordi_ in several other respects.

In _M. woodfordi_, as in other macroglossine bats, there is variability in dentition. One specimen examined has a total of three upper incisors, and another had an extra peglike tooth just anterior to I1.

Subfamily Nyctimeninae

=Nyctimene= Borkhausen

1797. _Nyctimene_ Borkhausen, Deutsche fauna ..., 1:86.

1810. _Cephalotes_ É. Geoffroy, Ann. du Mus. d'Hist. Nat., 15:104.

1811. _Harpyia_ Illiger, Prodr. Syst. Mamm. et Avium, p. 118.

1837. _Gelasinus_ Temminck, Monographe de Mammalia ..., 2:100.

Tube-nosed bats of the genus _Nyctimene_ occur from Celebes on the west to the Santa Cruz Islands on the east. Heretofore, two species (_N. albiventer_ and _N. major_), each with an endemic subspecies, were known from the Solomon Islands. Both species occur also in New Guinea and on many adjacent islands. A new species of _Nyctimene_, apparently endemic to the Solomons, and a new subspecies of _N. albiventer_ are named beyond.

_Nyctimene_ is related closely to _Cynopterus_ and the "Cynopterus group" of Andersen (1912:691). Because _Nyctimene_ is a highly specialized bat, Miller (1907:75) placed it in a subfamily separate from that of _Cynopterus_ and its allies.

Andersen (1912:696, 697) placed the species of _Nyctimene_ previously known from the Solomons in two groups, the "papuanus" group and the "cephalotes" group, on the basis of difference in length of forearm and length of maxillary tooth-row. Because of its short forearm (about 58), _N. albiventer_ is in the _papuanus_ group; and _N. major_, because of its long forearm (about 74), is in the _cephalotes_ group.

Key to Species of Nyctimene in the Solomons

1. Forearm longer than 70; males grayish-brown, females
pale gray, =N. major scitulus=, p. 825

1'. Forearm shorter than 70; males dark brown, females
pale brown, 2

2(1'). Forearm about 65, =N. malaitensis=, p. 822

2'. Forearm less than 61, =N. albiventer=, p. 818

=Nyctimene albiventer=

This species occurs throughout New Guinea and on many adjacent islands, including the Bismarck Archipelago and the Admiralty and Solomon islands. The species varies geographically and five subspecies are recognized. The two subspecies in the Solomons resemble _N. albiventer papuanus_, the subspecies that ranges from eastern New Guinea to New Britain. _N. albiventer bougainville_ occurs in the western chain of islands of the Solomons, whereas another subspecies, named as new beyond, occurs in the eastern chain of islands (see Fig. 12).

Sexual dichromatism is striking. As Andersen (1912:690) previously reported, females generally are paler, more brownish than males, which are dark and have a better defined black dorsal stripe.

=Nyctimene albiventer bougainville= Troughton

1936. _Nyctimene bougainville_ Troughton, Rec. Australian Mus.,
19:349, April 7, type from Bougainville.

1954. _Nyctimene albiventer bougainville_, Laurie and Hill, List of
land mammals of New Guinea, Celebes and adjacent islands, p. 46,
June 30.

1953. _Nyctimene papuanus bougainville_, Pohle, Z. Säugetierk.,
17:130, October 27.

_Specimens examined_ (nine males, one female; nine in alcohol, nine crania extracted and cleaned).--Bougainville in December, AM-M. 5786 (paratype); Guadalcanal in May, 23812, 23815, 23827; Kolombangara in January and February, 23369, 23381, 23388, 23406, 23444, 23456.

_Measurements._--See Table 3.

_Remarks._--Heretofore, _Nyctimene albiventer bougainville_ was not known from Kolombangara and Guadalcanal. The subspecies apparently ranges throughout the western chain of the Solomons.

Troughton (1936:350) considered _Nyctimene bougainville_ specifically distinct from its nearest ally, _N. papuanus_. Pohle (1953:130) did not examine specimens of either kind, but on the basis of Troughton's description decided that _N. bougainville_ differed only subspecifically from _N. papuanus_. Laurie and Hill (1954:46) synonymized _bougainville_ and _papuanus_ with _N. albiventer_. However, Troughton (1936:350) pointed out that in addition to size _bougainville_ differed from _papuanus_ by having narrower and longer pm3 and pm4. Judging from specimens examined by me, such is the case, and the difference is even more pronounced in m1.

Specimens of _N. a. bougainville_ from Kolombangara and Guadalcanal agree with a paratype of this subspecies from Bougainville. Geographic variation, if present in the population in the western chain of islands (see Fig. 12), is slight and not notable in the series available. Some individual variation was found, especially in the shape of the interorbital region of the skull. An adult male from Kolombangara is unusually dark, almost black; color of the other specimens (all in alcohol) is consistent according to sex.

_Nyctimene albiventer_ from Choiseul and Santa Ysabel is smaller, in all respects, than _N. albiventer_ from Bougainville, Kolombangara, and Guadalcanal (see Table 3), and therefore may be named and described as follows:

=Nyctimene albiventer minor=, new subspecies

_Type._--Adult male, skin and skull, in good condition (originally stored in alcohol for about one year), no. BSIP 23636, Bernice P. Bishop Museum; from Choiseul Island, British Solomon Islands Protectorate; obtained on 11 March 1964, by Philip Temple, original number 1441.

_Distribution._--Known only from Choiseul and Santa Ysabel islands (see Fig. 12).

_Diagnosis._--Small for _Nyctimene_; wing membranes brown with scattered yellow spots (dried specimens); uropatagium, feet, and ears brown; dorsum of tibia set with hair, ventral surface naked; dorsum of uropatagium sparsely set with pale brown hairs, ventral surface almost bare; fringe of hairs along two centimeters of dorsal and ventral surfaces of trailing edge of wing membrane; proximal third of dorsal surface of forearm sparsely set with hairs; pelage of back soft and thick, of medium length (about 7); hair on crown and nape short (about 4); well-defined black dorsal stripe, extending from uropatagium to shoulders; skull resembling that of other subspecies of _N. albiventer_ but relatively smaller; zygomatic arch delicate, slender anteriorly; P2 small (see Fig. 14). Sexually dichromatic as follows: male--dorsum Hair-Brown, bases of hairs darker; hair on throat sparse, medium length (about 6), Hair-Brown; fur along sides of abdomen Drab; female--dorsum having Buffy-Brown cast, some individual hairs Hair-Brown; shoulders Sayal-Brown; hair on throat sparse, Hair-Brown on throat and midline of abdomen; sides of abdomen Sayal-Brown.

_Comparisons._--From _Nyctimene major scitulus_, the largest member of this genus in the Solomons, _N. a. minor_ differs in being smaller in all measurements taken; forearm averaging 54.8 as opposed to 73.5; greatest length of skull 28.2 as opposed to 37.0, and females pale brown instead of pale gray.

From nine adults of _Nyctimene albiventer bougainville_ from Bougainville, Kolombangara, and Guadalcanal, _minor_ differs as follows: averaging slightly smaller in all dimensions; forearm averaging 54.8 as opposed to 57.9; second metacarpal averaging 27.4 as opposed to 28.3; 5th metacarpal averaging 38.5 as opposed to 40.0; condylobasal length 26.7 as opposed to 28.0; length of mandibular tooth-row 10.3 as opposed to 10.9; mandible smaller (see Fig. 14); dorsal stripe fainter.

From _Nyctimene albiventer papuanus_, known from eastern New Guinea, New Britain, and the Admiralty Islands, _minor_ differs as follows: slightly smaller in most dimensions; forearm averaging 54.8 as opposed to 57.0; length of maxillary tooth-row 8.9 as opposed to 9.8; length of mandibular tooth-row 10.3 as opposed to 11.0; breadth across upper third premolars notably less (7.5 as opposed to 8.4).

_N. a. minor_ differs from _N. albiventer albiventer_ Gray, which occurs about 800 miles to the west of _minor_, in ways made apparent by the description by Andersen (1912:700-701). _N. a. minor_ occurs about 1500 miles eastward of the place from which _N. a. draconilla_ Thomas, a subspecies essentially unknown to me, was named (see Laurie and Hill, 1954:46).

From _Nyctimene sanctacrucis_, known from the Santa Cruz Islands, _minor_ differs as follows: much smaller in all dimensions; forearm averaging 54.8 as opposed to 75; greatest length of skull 28.2 as opposed to 34.5; length of maxillary tooth-row 8.9 as opposed to 12.9.

) and _N. albiventer minor_ ([LW]).
For names of islands see Fig. 2.]

), _N. a. minor_ ([TW]), and intergrades
assigned to _minor_ ([BC]). For names of islands see Fig. 2.]

TABLE 3. Average and Extreme Measurements of _Nyctimene albiventer
bougainville_ and _N. a. minor_.

===============+==================+==================+===================
| _N. a. minor_ | _Intergrades_ | _N. a.
| | | bougainville_
| | |
MEASUREMENT | Choiseul, | Fauro | Kolombangara,
| Santa Ysabel | | Guadalcanal
| 4 [M], 1 [F] | 1 [M], 3 [F] | 8 [M], 1 [F]
---------------+------------------+------------------+-------------------
Length of head | | |
and body |107.0 (105 -109 )|109.2 (105 -112 )|110.0 (106 -117)
Tail vertebrae | 20.0 (19.3- 20.5)| 21.0 ( 20 - 22 )| 19.2 ( 15.5- 23.0)
Hind foot | 14.2 (13.5- 15.0)| | 14.3 ( 13.0- 15.9)
Ear | 11.9 (11.0- 13.0)| | 12.8 ( 11.8- 14.5)
Length of | | |
forearm | 54.8 (54.0- 55.8)| 57.1 (55.9- 59.0)| 57.9 ( 55.8- 59.8)
Greatest | | |
length of | | |
skull | 28.2 (27.2- 28.9)| 28.6 (28.3- 29.7)| 29.7 ( 28.6- 30.1)
Condylobasal | | |
length | 26.7 (26.2- 27.5)| 27.4 (26.6- 28.0)| 28.0 ( 27.8- 28.9)
Palatal length | 11.2 (10.9- 11.9)| 11.6 (11.3- 11.8)| 11.7 ( 11.0- 12.5)
Breadth of | | |
braincase | 12.0 (11.5- 12.4)| 12.0 (11.7- 12.2)| 12.3 ( 12.1- 12.8)
Zygomatic | | |
breadth | 18.9 (18.4- 19.7)| 18.6 (18.4- 19.2)| 19.2 ( 18.7- 20.0)
Interorbital | | |
breadth | 5.0 ( 4.7- 5.6)| 5.3 ( 5.0- 5.6)| 5.1 ( 4.7- 5.5)
Breadth across | | |
first upper | | |
molars | 8.6 ( 8.4- 8.9)| 8.9 ( 8.7- 9.1)| 9.1 ( 8.8- 9.6)
Maxillary | | |
tooth-row | 8.9 ( 8.7- 9.3)| 9.3 ( 9.1- 9.5)| 9.5 ( 9.2- 9.8)
Mandibular | | |
tooth-row | 10.3 (10.0- 10.6)| 10.5 (10.2- 11.1)| 10.9 ( 10.7- 11.4)
---------------+------------------+------------------+-------------------

_Measurements._--Measurements of the two subspecies from the Solomons are given in Table 3. Some measurements of the type are as follows: Length of head and body, 108; tail vertebrae, 20.5; hind foot, 14.7; ear, 11.3; length of forearm, 55.1; 2nd metacarpal, 27.4; 3rd metacarpal, 39.0; 4th metacarpal, 37.5; 5th metacarpal, 39.1; greatest length of skull, 28.6; condylobasal length, 27.5; zygomatic breadth, 18.4; length of maxillary tooth-row, 9.0; length of mandibular tooth-row, 10.4.

_Remarks._--_Nyctimene albiventer minor_ closely resembles _N. albiventer bougainville_, differing from the latter mostly in size. Although adults of _minor_ average only slightly smaller than adults of _bougainville_ (see Table 3), there is only slight overlap (about 0.2 at most) in most minimum dimensions of external and cranial features of _bougainville_ and corresponding maximum dimensions of externals and crania of _minor_. The difference in size is clearly shown in Figs. 13 and 14.

Four specimens of _Nyctimene albiventer_ from Fauro herein are considered to be intergrades between _N. a. bougainville_ and _N. a. minor_. As shown in Table 3, the specimens from Fauro average slightly larger than those of _minor_ from Choiseul and Santa Ysabel and slightly smaller than specimens of _bougainville_ from Kolombangara and Guadalcanal. I have assigned the specimens from Fauro to _N. a. minor_ because they generally are closer to _minor_ in size (see Fig. 13).

_Specimens examined_ (five males and four females; seven in alcohol; seven crania extracted and cleaned).--Choiseul in February and March, 23636 (holotype), 23631, 23540, 23646; Santa Ysabel in February, 23539; Fauro in April, 23742, 23743, 23763, 23764.

One specimen of _Nyctimene_ from Malaita Island is smaller than _Nyctimene major_, which is known from Shortland, Alu, Florida, New Georgia, Guadalcanal, Choiseul, and Malapa (see Fig. 15) and is larger than either of the two subspecies of _Nyctimene albiventer_ known from Bougainville, Fauro, Kolombangara, Guadalcanal, Choiseul, and Santa Ysabel. This specimen represents a previously unknown species and may be named and described as follows:

=Nyctimene malaitensis=, new species

_Type._--Adult female, skin and skull, in good condition (originally stored in alcohol for about one year), no. BSIP 24103, Bernice P. Bishop Museum; from Malaita Island, British Solomon Islands Protectorate; obtained on 1 July 1964, by Peter Shanahan, original no. unknown.

_Distribution._--Known only from Malaita (see Fig. 16).

_Diagnosis._--Size average for genus but larger than closest relative, _Nyctimene albiventer_; wing membranes brown with scattered yellow spots (dried specimen); uropatagium, ears, and feet brown; dorsal surface of tibia set with hair, ventral surface bare; dorsal surface of uropatagium sparsely set with hair, ventral surface having few, scattered hairs; dorsal surface of trailing edge of wing membrane sparsely set with hairs, ventral surface bare; proximal third of upper- and under-surface of forearm set with hair; pelage of back luxuriant and soft (about 10 long); hair on crown and nape shorter than on back (4 to 8); well-defined black dorsal stripe from shoulders to rump (about 2 wide); basal half of most hairs on dorsum Deep Mouse Gray, distal half Light Buff, tips Ochraceous-Tawny; some hairs on back entirely Light Buff; hairs of crown Light Ochraceous Buff tipped with Ochraceous-Tawny; hair on throat and along sides of abdomen Light Ochraceous Buff; hairs of ventral midline Smoke Gray; braincase narrow; zygomatic breadth relatively narrow; well-developed lambdoidal crest in female; rostrum short, wide; upper canines slanted posteriorly; upper incisors large; foramen ovale large (see Fig. 14).

],
(B) _N. a. bougainville_ [specimen 23381 [M]], and
(C) _N. malaitensis_ [specimen 24103 [F]].]

_Comparisons._--From _Nyctimene major scitulus_, the largest kind of _Nyctimene_ in the Solomons, _malaitensis_ differs as follows: smaller in all dimensions (forearm 65 as opposed to 73.5); greatest length of skull 32.4 as opposed to 37.0; length of maxillary tooth-row 10.5 as opposed to 13.0; length of mandibular tooth-row 11.8 as opposed to 14.2.

From nine adults of _Nyctimene albiventer bougainville_ from Bougainville, Kolombangara, and Guadalcanal, _malaitensis_ differs as follows: larger in all dimensions: forearm 65 as opposed to 57.9; greatest length of skull 32.4 as opposed to 29.7; zygomatic breadth 20.4 as opposed to 19.2; and length of maxillary tooth-row 10.5 as opposed to 9.5; length of mandibular tooth-row 11.8 as opposed to 11.1.

From five adults of _Nyctimene albiventer minor_, from Choiseul and Santa Ysabel, _malaitensis_ differs in the same ways it differs from _N. a. bougainville_, but the contrast is even greater when _malaitensis_ and _minor_ are compared.

From _Nyctimene sanctacrucis_, known only from the Santa Cruz Islands, _malaitensis_ differs in being smaller in all dimensions: forearm 65 as opposed to 75; greatest length of skull 32.4 as opposed to 34.5; and length of maxillary tooth-row 10.5 as opposed to 12.9.

_Measurements of the holotype._--Length of head and body, 118; tail vertebrae, 23.0; hind foot, 16.0; ear, 14.0; length of forearm, 65.0; 2nd metacarpal, 33.2; 3rd metacarpal, 46.4; 4th metacarpal, 44.3; 5th metacarpal, 46.0; greatest length of skull, 32.4; condylobasal length, 30.6; palatal length, 13.0; breadth of braincase, 12.5; zygomatic breadth, 20.4; interorbital breadth, 5.5; breadth across first upper molars, 9.5; length of maxillary tooth-row, 10.5; length of mandibular tooth-row, 11.8.

and _N. major scitulus_ [RW]. For names of islands see Fig. 2.]

_Remarks._--In size, _Nyctimene malaitensis_ is intermediate between _N. albiventer_ and _N. major_. Because the type of _malaitensis_ is brown and not pale gray, as are females of _major_, _N. malaitensis_ most likely is more closely related to _N. albiventer_, in which the females are brown. The teeth of the holotype and only known specimen of _malaitensis_ are too worn to be useful in determining the relationships between these species.

When more specimens are available, _N. malaitensis_ may prove to be a subspecies of _N. albiventer_. At present, _malaitensis_ is accorded specific rank in order not to obscure the apparent relationships of _N. albiventer bougainville_ and _N. a. minor_. Additionally, _N. malaitensis_ is given specific rank because (1) it is larger (especially in external dimensions) than the largest subspecies of _N. albiventer_ (compare above measurements with those in Table 3), and (2) _malaitensis_ does not form a cline with either of the two subspecies of _N. albiventer_.

_Specimen examined_ (one female).--Malaita in July, 24103 (holotype).

=Nyctimene major=

This large species of tube-nosed bat has at least four subspecies, one of which (_N. major scitulus_) is endemic to the Solomons. The species occurs throughout eastern New Guinea and on many of the islands adjacent to the eastern coast of New Guinea, including the Trobriand Islands, the Bismarck Archipelago, and the Solomons (see Laurie and Hill, 1954:47). The geographic distribution of the species generally is the same as that of _N. albiventer_.

In _Nyctimene major_, as in _N. albiventer_, most males are grayish-brown, whereas most females are pale gray.

=Nyctimene major scitulus= Andersen

1910. _Nyctimene scitulus_ Andersen, Ann. Mag. Nat. Hist., ser. 8,
6:623, December 1, type from Shortland; 1912, Andersen, Catalogue
of the Chiroptera ... British Museum, 1:711, from Shortland, New
Georgia, Florida, Guadalcanal; 1931, Troughton, Proc. Linnean Soc.
New South Wales, 56:206, July 15; 1931, Sanborn, Publ. Field Mus.
Nat. Hist., 18:22, February 12, from Choiseul and Malapa; 1942,
Tate, Bull. Amer. Mus. Nat. Hist., 80:342, December 31.

1954. _Nyctimene major scitulus_, Laurie and Hill, List of land
mammals of New Guinea, Celebes and adjacent islands, p. 47,
June 30.

1862. _Harpyia pallasi_, Gerrard, Catalogue of the bones ...
British Museum, p. 58.

1870. _Harpyia cephalotes_, Gray, Catalogue of monkeys, lemurs and
fruit-eating bats in the British Museum, p. 121.

1878. _Harpyia major_, Dobson, Catalogue of the Chiroptera ...
British Museum, p. 90; 1879, Trouessart, Rev. Mag. Zool., 3:207;
1887, Thomas, Proc. Zool. Soc. London, p. 323; 1888, Thomas, Proc.
Zool. Soc. London, p. 476; 1897, Trouessart, Catalogus Mammalium
..., 1:87.

1899. _Cephalotes major_, Trouessart, Catalogus Mammalium ...,
2:1277.

1899. _Gelasinus major_, Matschie, Die Megachiroptera ...
naturkunde, p. 84; 1904, Trouessart, Catalogus Mammalium ...,
Suppl., p. 64.

_Specimens examined_ (four males and one female; dried skins with skulls inside).--Florida in October, 24397, 24413, 24418, 24419.

_Measurements._--External measurements of four males and one female are, respectively, as follows: Length of head and body, 134, 128, 134, 134, 136; tail vertebrae, 28, 23, 27, 26, 21; hind foot, 20, 16, 19, 16, 21; ear, 17, 17, 17, 17, 18; length of forearm, 73.8, 68.0, 74.0, 73.6, 78.0.

_Remarks._--_Nyctimene major scitulus_ has been recorded only from the western chain of islands in the Solomons (see Fig. 15). Specimens examined by me agree well in external dimensions and color with specimens described by Andersen (1912:712) and Troughton (1931:206-207).

ZOOGEOGRAPHY AND SPECIATION