Part 4

_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.

=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.

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.