Part 5

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

=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.

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.