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
De Beaufort (1951:113) considered bats of "less zoogeographical
importance" than other mammals because the ocean is not an "absolute
barrier to their dispersal." Volant animals are ecologically terrestrial
and therefore are more nearly earthbound than De Beaufort's remarks
would suggest (see Miller, 1966:10). Indeed, many kinds of volant
animals are endemic to the Solomons. Birds, for example, are well
adapted for flight but pose some of the most complex zoogeographic
problems in the area of New Guinea and the Solomon Islands (Mayr,
1940:198; 1942:81-83; Koopman, 1957). Rapid speciation can take place
in any situation where there is a high degree of isolation (Wright,
1931; Lack, 1947). In fact, isolation is a most important factor in
speciation of insular populations (Baker, 1951:55). The one genus, nine
species, and 19 subspecies of megachiropterans that are endemic to the
Solomons (Table 4) obviously indicate that bats, although volant, can be
restricted to one or more islands long enough for new taxa to evolve.
TABLE 4. A Summary of the Kinds of Megachiropteran Bats in the
Solomon Islands and Their Affinities with Faunas of Adjacent
Islands.
===========+========+==========+===========+============+===============
| | | Common | Common to | Common to
| | Endemic | only to | Solomons, | Solomons,
| Totals | to | Solomons | Bismarcks, | New Hebrides,
| | Solomons | and | and | and
| | | Bismarcks | New Guinea | New Caledonia
-----------+--------+----------+-----------+------------+---------------
Genera | 7 | 1 | 0 | 6 | 0
Species | 16 | 9 | 1 | 6 | 1
Subspecies | 20 | 19 | 0 | 0 | 1
-----------+--------+----------+-----------+------------+---------------
The megachiropteran bats of the Solomons have their affinities with the
fauna of New Guinea (Table 4); the Solomons and New Guinea have six
genera and six species in common. Because the two areas never have been
connected (_via_ the Bismarck Archipelago) by dry land, bats probably
have reached the Solomons by flying from island to island (see Durham,
1963:357, 359, 361, 363). Deignan (1963:266) has dismissed voluntary or
involuntary flight as possible explanations for distributions of bats
and birds on islands of the Pacific.
The taxonomic level of endemism can be used as an indicator of antiquity
(Dobzhansky, 1941; Koopman, 1958:429-430). The one megachiropteran genus
(_Pteralopex_) endemic to the Solomons apparently is an ancient relic.
Bats of this monotypic genus occur on Bougainville, Choiseul, Santa
Ysabel, and Guadalcanal (see Fig. 4). These four islands probably were
contiguous during the maximum lowering of sea level in the Pleistocene
(see Durham, 1963:362-363). Bats of the genus _Pteralopex_ are the only
kind in the Solomons having a distribution that can be correlated with
former land connections between islands.
The distributions of 16 species of megachiropterans known from the
Solomons are summarized in Table 5 and in Figure 16. The larger islands
(in terms of surface area and elevation) in general have the highest
number of species (Guadalcanal 10, Choiseul 9, and Bougainville 8). But
Fauro, one of the smallest islands for which data are available, has six
species of megachiropterans whereas San Cristobal and Malaita, two of
the larger islands, have only three and four species, respectively.
Possibly this difference signals the need for additional collecting.
Bougainville and Choiseul, about 60 miles apart, have seven species of
megachiropterans in common (Table 5). Fauro, 25 miles southeast of
Bougainville and 35 miles west of Choiseul, shares five species with
each of these islands (Fig. 16). _Pteralopex atrata_ and _Pteropus
rayneri_ occur on Choiseul and on Bougainville, but not on Fauro.
Individuals of these species are the largest fruit bats in the Solomons,
and their absence on Fauro suggests, therefore, that this small island
is ecologically unsuitable, at least in some months, for the support of
populations of bats that require relatively large amounts of food. The
small size of the island is consistent with this hypothesis, but several
other islands as small as Fauro do support populations of the large
kinds of _Pteropus_, at least in some months.
TABLE 5. A Summary of Distribution of All Species of Megachiropteran
Bats Known from the Solomons. Only Islands Well Known Faunistically
Are Listed.
Column headings:
A: Bougainville I: Vella Lavella
B: Choiseul J: Kolombangara
C: Santa Ysabel K: Russell
D: Ndai L: Guadalcanal
E: Malaita M: San Cristobal
F: Florida N: Ugi
G: Fauro O: Rennell
H: Shortland P: Ontong Java
===================+==+==+==+==+==+==+==+==+==+==+==+==+==+==+==+==
SPECIES | A| B| C| D| E| F| G| H| I| J| K| L| M| N| O| P
-------------------+--+--+--+--+--+--+--+--+--+--+--+--+--+--+--+--
R. amplexicaudatus | X| X| X| | X| | X| | | X| | X| | | |
P. atrata | X| X| X| | | | | | | | | X| | | |
Pt. hypomelanus | | | | | | | | | | | X| | | | |
Pt. admiralitatum | | | | X| | | | X| X| | X| X| | | |
Pt. tonganus | | | | | | | | | | | | | | | X|
Pt. howensis | | | | | | | | | | | | | | | | X
Pt. rayneri | X| X| X| | X| | | | X| X| | X| X| X| X|
Pt. woodfordi | | | | | | | X| | | X| X| X| | | |
Pt. mahaganus | X| | X| | | | | | | | | | | | |
D. inermis | X| X| X| | X| X| X| X| X| | | X| X| X| X|
M. lagochilus | X| X| | | | X| X| | | X| | X| X| | |
M. woodfordi | X| X| | | | | X| X| | | X| X| | | |
M. aurantius | | X| | | | X| | | | | | | | | |
N. albiventer | X| X| X| | | | X| | | X| | X| | | |
N. major | | X| | | | X| | X| | | | X| | | |
N. malaitensis | | | | | X| | | | | | | | | | |
+--+--+--+--+--+--+--+--+--+--+--+--+--+--+--+--
Totals | 8| 9| 6| 1| 4| 4| 6| 4| 3| 5| 4|10| 3| 2| 3| 1
-------------------+--+--+--+--+--+--+--+--+--+--+--+--+--+--+--+--
Santa Ysabel has six species of megachiropterans and 10 occur on
Guadalcanal (Table 5). These two islands, separated by about 100 miles
of water, share five species (_Rousettus amplexicaudatus_, _Pteralopex
atrata_, _Pteropus rayneri_, _Dobsonia inermis_, and _Nyctimene
albiventer_). The Nggela Group, in which Florida is the largest island
and the only one from which bats have been collected, is 50 miles
southeast of Santa Ysabel and 30 miles north of Guadalcanal (Fig. 16).
Four species of megachiropterans are known from Florida (_Dobsonia
inermis_, _Macroglossus lagochilus_, _Melonycteris aurantius_, and
_Nyctimene major_). Three of these are known from Guadalcanal and one
occurs on Santa Ysabel. This situation resembles the one involving
Fauro, Bougainville, and Choiseul because none of the large bats
(_Pteropus_ and _Pteralopex_) is known from Florida, even though two
species of large bats that occur on Santa Ysabel to the northwest occur
also on Guadalcanal to the south. Possibly Florida and the smaller
islands that comprise the Nggela Group are ecologically unsuitable for
large bats, or perhaps these small islands can support only limited
numbers of individuals during part of a year.
[Illustration: FIG. 16. The number of megachiropteran species known
from individual islands (number within a circle) is compared with
the number of species common to two different islands (number
without a circle). For names of islands see Fig. 2.]
Some of the small islands in the Solomons have populations of large
fruit bats. For example, _Pteropus admiralitatum_ and _P. hypomelanus_
have been reported from the small islands in the Russell Group (Table
5). Possibly these species do not live concurrently in the Russells;
specimens of the two were obtained in different years. Two small
megachiropterans, _P. woodfordi_ and _Melonycteris woodfordi_, also
inhabit the Russells. Shortland, a small island about 15 miles south of
Bougainville, supports one large bat, _P. admiralitatum_, as well as
smaller megachiropterans.
Kolombangara and Vella Lavella are about the same size and are separated
by about 15 miles of water. _Rousettus amplexicaudatus_, _Pteropus
rayneri_, _P. woodfordi_, _Macroglossus lagochilus_, and _Nyctimene
albiventer_ have been collected on Kolombangara but only _P.
admiralitatum_, _P. rayneri_, and _Dobsonia inermis_ have been found on
Vella Lavella. The difference in the known megachiropteran faunas is
more striking when one compares each island with adjacent islands. Two
species on Vella Lavella occur also on Choiseul, which is about 35 miles
northeastward, and two species occur also on Shortland, which is 120
miles northwestward (Fig. 16). Four of the five megachiropterans on
Kolombangara also have been found on Choiseul, about 50 miles northward
(Table 5). _Pteropus rayneri_ is the only megachiropteran known from
both Kolombangara and Vella Lavella, even though the islands are
separated by only a few miles of water. Inadequate data possibly account
for the differences in the megachiropteran fauna, but I suspect that
some other factors are involved. Although Vella Lavella and Kolombangara
do have one species (_P. rayneri_) in common, a different subspecies
occurs on each island--_rubianus_ on Kolombangara and _lavellanus_ on
Vella Lavella (Fig. 17 and Table 6). This indicates that some factor or
factors are operating to keep megachiropterans from moving frequently or
easily from one island to the other.
Each of several subspecies of species in the genus _Pteropus_ are known
from one or two small islands separated by only a few miles from other
islands on which different subspecies occur (see Fig. 6). Judging from
this kind of distribution, these bats do not move frequently from island
to island. Possibly this is because they cannot easily cross water
barriers, or are not inclined to do so because food is abundantly
available throughout the year on their home island. Because "flying
foxes" frequently are seen in flight over water several hundred yards
from shore, the first factor probably is unimportant--at least where
short distances are involved. It seems most likely that when abundant
food is available these bats have no reason to move even moderate
distances.
[Illustration: FIG. 17. The number of subspecies of megachiropterans
known from individual islands (number within a circle) is compared
with the number of subspecies common to different islands (number
without a circle). For names of islands see Fig. 2.]
Distributions of subspecies of polytypic species are summarized in Table
6 and Figure 17. Generally, more subspecies are known from the larger
islands than from the smaller islands (Guadalcanal with 5, Bougainville,
Choiseul, and Santa Ysabel with 4, Fauro with 2.) The distributions of
some subspecies can be used to judge the differential effectiveness of
water gaps between islands. The distribution of _Pteropus rayneri
lavellanus_ and _P. rayneri rubianus_ is an example.
Choiseul and Santa Ysabel are separated by about 50 miles of water (see
Fig. 17) but have three subspecies in common (_Pteropus rayneri
grandis_, _Dobsonia inermis minimus_, and _Nyctimene albiventer
minor_.) Choiseul is about 50 miles from Kolombangara and about 35 miles
from Vella Lavella, but shares no subspecies with these smaller islands
although some species are shared (Tables 5 and 6). From these data one
can conclude that exchange of genes between populations on Choiseul and
populations on Santa Ysabel is frequent but for some reason exchange of
genes between populations on Vella Lavella and Choiseul and Kolombangara
and Choiseul is infrequent. A series of small islands (Rob Roy, Wagina,
and the Arnavon Islands, not named on the maps) connect Choiseul and
Santa Ysabel in stepping-stone fashion (see Fig. 17). Possibly these
small islands enhance movement of megachiropterans between Choiseul and
Santa Ysabel.
TABLE 6. A Summary of Distribution of Polytypic Species of
Megachiropteran Bats in the Solomon Islands. Only Islands
Well Known Faunistically Are Listed.
Column headings:
A: Bougainville I: Vella Lavella
B: Choiseul J: Kolombangara
C: Santa Ysabel K: Russell
D: Ndai L: Guadalcanal
E: Malaita M: San Cristobal
F: Florida N: Ugi
G: Fauro O: Rennell
H: Shortland P: Ontong Java
===================+==+==+==+==+==+==+==+==+==+==+==+==+==+==+==+==
SUBSPECIES | A| B| C| D| E| F| G| H| I| J| K| L| M| N| O| P
-------------------+--+--+--+--+--+--+--+--+--+--+--+--+--+--+--+--
P. atrata atrata | X| X| | | | | | | | | | | | | |
P. atrata anceps | | | X| | | | | | | | | X| | | |
Pt. a. solomonis | | | | | | | | | X| | X| X| | | |
Pt. a. colonus | | | | | | | | X| | | | | | | |
Pt. a. grandis | | | | X| | | | | | | | | | | |
Pt. r. rayneri | | | | | X| | | | | | | X| | | |
Pt. r. grandis | X| X| X| | | | | | | | | | | | |
Pt. r. rubianus | | | | | | | | | | X| | | | | |
Pt. r. lavellanus | | | | | | | | | X| | | | | | |
Pt. r. monoensis | | | | | | | | | | | | | | | | X
Pt. r. cognatus | | | | | | | | | | | | | X| X| |
Pt. r. rennelli | | | | | | | | | | | | | | | X|
D. i. inermis | X| | | | X| X| X| X| X| | | X| X| X| X|
D. i. minimus | | X| X| | | | | | | | | | | | |
N. a. bougainville | X| | | | | | | | | X| | X| | | |
N. a. minor | | X| X| | | | X| | | | | | | | |
+--+--+--+--+--+--+--+--+--+--+--+--+--+--+--+--
Totals | 4| 4| 4| 1| 2| 1| 2| 2| 3| 2| 1| 5| 2| 2| 2| 1
-------------------+--+--+--+--+--+--+--+--+--+--+--+--+--+--+--+--
Florida, of the Nggela Group, is approximately halfway between Santa
Ysabel and Guadalcanal. _Pteralopex atrata anceps_ occurs on Santa
Ysabel and on Guadalcanal but is unknown from Florida. Fauro lies
between Bougainville and Choiseul. _Pteralopex atrata atrata_ and
_Pteropus rayneri grandis_ occur on Choiseul and on Bougainville but are
unknown from Fauro. As suggested earlier, small islands like Fauro and
Florida possibly cannot support large fruit bats, although they probably
would utilize these small islands when in transit between larger
islands.
Fauro apparently is important to the distribution of the two subspecies
of _Dobsonia inermis_ and _Nyctimene albiventer_ in the Solomons (see
Figs. 9 and 13). In both species, one subspecies is found in the eastern
chain of islands and one subspecies is found in the western chain.
Specimens of _Dobsonia inermis_ from Fauro and Bougainville can be
identified as the subspecies _inermis_ whereas those from Choiseul are
assignable to the subspecies _minimus_. _Nyctimene albiventer
bougainville_ occurs on Bougainville but specimens of _N. albiventer_
from Fauro and Choiseul can be identified as the subspecies _minor_.
Although interchange of genes occurs between populations on Bougainville
and Fauro in the case of _D. inermis_, the population of _N. albiventer_
on Fauro is at least partially isolated from the population on
Bougainville.
Rennell and Ontong Java are relatively isolated from other islands in
the Solomons (see Fig. 17). Only one kind of bat (_Pteropus howensis_)
is known from Ontong Java and apparently is endemic to that atoll.
_Pteropus tonganus geddiei_, one of the megachiropterans that occurs on
Rennell (Table 5), also is found in the New Hebrides and on New
Caledonia (Table 4). This makes _P. t. geddiei_ the only megachiropteran
bat in the Solomons that is more closely related to bats on islands to
the southeast of the Solomons than to bats on other islands of the
Solomons, the Bismarcks, or New Guinea, to the north and west. The other
species of megachiropterans (_Dobsonia inermis_ and _Pteropus rayneri_)
on Rennell are found also on other islands in the Solomons. It is to be
noted that Mayr (1931) regarded the avifauna of Rennell as most nearly
like that of the New Hebrides and New Caledonia. He suggested that the
prevailing winds from the southeast have been important for birds that
have reached Rennell. The New Hebrides and New Caledonia are four and a
half times farther from Rennell than are San Cristobal and Guadalcanal.
On first consideration a person might doubt that the winds would be
favorable enough to compensate for the great distance between Rennell
and the New Hebrides and New Caledonia. Darlington (1938) has used the
formula X n/m to obtain a comparison of barriers of different widths.
[X = the probability of an individual crossing a barrier of width m;
the probability of an individual crossing a similar barrier of width n
is the ratio n/m.] If this formula is applied here, one finds that winds
from the southeast (that is, from the New Hebrides and New Caledonia)
would have to be more than 100 times more favorable than winds from the
northeast (from Guadalcanal and San Cristobal) in order to compensate
for the distance of Rennell from the New Hebrides and New Caledonia.
Even so, tropical storms with unusually strong winds, frequent during
some parts of the year, possibly account for the present distributional
pattern of bats and birds that live on Rennell.
Whatever the means by which bats of the species _P. tonganus_ reached
Rennell, the fact remains that specimens from Rennell cannot be
distinguished from specimens of _P. tonganus geddiei_ from the New
Hebrides and New Caledonia, more than 500 miles to the southeast.
NOTE: An important and interesting paper on zoogeography of bats, which
was published too late to be included here, is: Krzanowski, A., 1967,
The magnitude of islands and the size of bats (Chiroptera), Acta Zool.
Cracoviensia, 12:281-348.
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