_Pteropus tonganus_ has at least three subspecies, one of which has been
recorded from the Solomons. The species ranges from a small island off
the eastern coast of New Guinea, where there is an endemic subspecies,
eastward to Tonga and the New Hebrides (Laurie and Hill, 1954:33-34).
Felten (1964_a_) recently has reported on the species in the New
Hebrides.
=Pteropus tonganus geddiei= MacGillivary
1860. _Pteropus geddiei_ MacGillivary, Zoologist, 18:7134,
September, type from Aneitum Island, New Hebrides; 1912,
Andersen, Catalogue of the Chiroptera ... British Museum,
1:189; 1931, Sanborn, Publ. Field Mus. Nat. Hist., Zool.
Ser., 18:13, February 12, from Rennell Island in the Solomons.
1914. _Pteropus tonganus geddiei_, Revilliod, _in_ Sarasin and
Roux, Nova Caledonia (A), 1:341; 1954, Laurie and Hill, List
of land mammals of New Guinea, Celebes and adjacent islands,
p. 34, June 30.
_Specimens examined._--None.
_Remarks._--_Pteropus tonganus geddiei_, as far as is known, is the
widest ranging subspecies of this genus. It is the only megachiropteran
in the Solomon Islands having affinities with bats to the southeast (the
New Hebrides, Santa Cruz Islands, Samoan Islands and Fiji Islands)
rather than with those to the west (New Guinea). The subspecies _P.
tonganus geddiei_, which ranges from the Solomons to the New Hebrides
(about 500 miles straight-line distance), is said to be remarkably
uniform throughout its range. Sanborn (1931:14) compared color and size
in specimens from the Solomon Islands and the New Hebrides and found
little variation. Another subspecies, _P. t. bascilicus_ Thomas 1915,
apparently closely related to _geddiei_, is known from Dampier [=
Kar-kar] Island off the northeastern coast of New Guinea and therefore
farther westward from the New Hebrides than are the Solomon Islands.
Additional remarks on the distribution of this species are in the
section on Zoogeography and Speciation.
=Pteropus rayneri=
_Pteropus rayneri_ is endemic to the Solomon Islands. It is divisible
into seven subspecies (see Fig. 6), which, excepting _P. r. rennelli_
and _P. r. cognatus_, are strikingly colored--the mantle, back, and rump
being of different colors. Differences in color and size provide
characters differentiating the subspecies (see key, p. 793). Recorded
lengths of forearms do not overlap between any two subspecies. _P. r.
grandis_, northernmost in distribution, has the longest (about 170)
forearm and _P. r. cognatus_, known from two of the southernmost
islands, has the shortest (about 121).
[Illustration: FIG. 6. Distribution of _Pteropus rayneri_: _P. r.
rayneri_ ([RTW]); _P. r. grandis_ ([RW]); _P. r. lavellanus_ ([BC]);
_P. r. monoensis_ ([BW]); _P. r. rubianus_ ([TW]); _P. r. cognatus_
([LW]); _P. r. rennelli_ ([LTW]). For names of islands see Fig. 2.]
Adult males of _Pteropus rayneri_ have well-developed tufts of hair on
each side of the neck where a gland is located (see Andersen,
1912:259). Apparently these glands are not present in females as none
were found in specimens studied by me or those reported by Sanborn
(1931:16). Evidently, these glands are associated with sexual maturity
in males because neither Sanborn nor I found them in subadult males.
=Pteropus rayneri rayneri= Gray
1870. _Pteropus rayneri_ (part), Gray, Catalogue of monkeys, lemurs
and fruit-eating bats ... British Museum, p. 108, cotypes from
Guadalcanal; 1878, Dobson, Catalogue of the Chiroptera ...
British Museum, p. 33; 1879, Trouessart, Rev. Mag. Zool., 6:204;
1879, Trouessart, Ann. Sci. Nat. Zool, 8:16; 1887, Thomas, Proc.
Zool. Soc. London, p. 322, March 15; 1888, Thomas, Proc. Zool.
Soc. London, p. 472, December 4; 1898, Trouessart, Catalogus
Mammalium ..., 1:78; 1912, Andersen, Catalogue of the Chiroptera
... British Museum, p. 254; 1931, Sanborn, Publ. Field Mus. Nat.
Hist., Zool. Ser., 18:15, February 12, from Guadalcanal
and Malaita.
1954. _Pteropus rayneri rayneri_, Laurie and Hill, List of land
mammals of New Guinea, Celebes and adjacent islands, p. 35,
June 30.
1899. _Pteropus (Spectrum) rayneri_ (part), Matschie, Die
Megachiroptera ... naturkunde, p. 22; 1904, Trouessart,
Catalogus Mammalium ..., Suppl., p. 51.
_Specimens examined_ (four males and one female; one embryo in
alcohol).--Guadalcanal in July and November, USNM 278700-02, USNM
278142, USNM 278714.
_Measurements._--Measurements of three males and one female are,
respectively, as follows: Length of head and body, --, 210, 214, 215;
hind foot, --, 33, 39, 42; ear, --, 23, 23, 23; length of forearm, --,
138, 136, 134; greatest length of skull, 61.5, 59.2, 61.6, 61.2;
condylobasal length, 61.4, 58.2, 60.3, 60.0; zygomatic breadth, 36.6,
35.3, 35.4, 36.5; breadth of braincase, 23.7, 22.5, 22.6, 24.1; breadth
across first upper molars, --, 16.9, 16.7, 16.8; width of M1, 3.4, 3.5,
3.5, 3.5; length of maxillary tooth-row, 22.4, 22.1, 23.6, 23.2; length
of mandibular tooth-row, 26.4, 25.5, 25.9, 25.6.
_Remarks._--_Pteropus rayneri_ was named on the basis of two specimens
(cotypes) obtained on Guadalcanal and listed as "male" and "female";
according to Andersen (1912:254), however, both are females.
_P. r. rayneri_ is known from Guadalcanal and Malaita (see Fig. 6), and
is of almost the same size as _P. r. cognatus_, which is known from San
Cristobal and Ugi, only about 40 miles to the southeast. In the latter
subspecies the back and rump are the same color (Prouts Brown), whereas
in _P. r. rayneri_ the rump is brightly colored and therefore contrasts
strongly with the dark brown back. A specimen of _rayneri_ from Malaita
was reported by Sanborn (1931:15) as unusually small and having a
dark-colored rump patch. In the specimens examined from Guadalcanal,
there is noticeable variation in color of the mantle that does not seem
related to age or sex. In two specimens (adult male and female) the
mantle is Cinnamon-Rufous tinged with Russet, strongly contrasting with
the crown, which is Ochraceous-Tawny and has scattered silvery hairs.
Another specimen has a darker mantle (near Chestnut-Brown) and a crown
of about the same color, but with a few scattered Ochraceous-Tawny
hairs.
The skull of one adult male bears an extra peglike tooth posterior to M3
on the right side.
An embryo, in an advanced stage of development, in the collection of the
U. S. National Museum, measures: Length of head and body, 98; hind foot,
30; ear, 8.5; length of forearm, 48 (this may be the same specimen
listed by Sanborn and Nicholson, 1950:329).
=Pteropus rayneri grandis= Thomas
1887. _Pteropus grandis_ Thomas, Ann. Mag. Nat. Hist., ser. 5,
19:147, March, type from Shortland; 1887, Thomas, Proc. Zool.
Soc. London, p. 320, March 15, from Alu and Shortland; 1897,
Trouessart, Catalogus Mammalium ..., 1:80, from "I. Salomonis";
1899, Matschie, Die Megachiroptera ... naturkunde, p. 15; 1904,
Trouessart, Catalogus Mammalium ..., Suppl., p. 49; 1907, Miller,
Bull. U. S. Nat. Mus., 57:58, June 29; 1912, Andersen, Catalogue
of the Chiroptera ... British Museum, 1:259, from Bougainville;
1931, Sanborn, Publ. Field Mus. Nat. Hist., Zool. Ser., 18:16,
February 12, from Choiseul, and Santa Ysabel; 1936, Troughton,
Rec. Australian Mus., 19:348, April 7; 1953, Pohle, Z.
Säugetierk., 17:128, October 27.
1954. _Pteropus rayneri grandis_, Laurie and Hill, List of land
mammals of New Guinea, Celebes and adjacent islands, p. 35,
June 30.
_Specimens examined_ (six males and 10 females; five in
alcohol).--Choiseul in March, 23580, 23644, 23593; Bougainville in July,
August, September, and October, USNM 276926-7, USNM 276968, USNM
277091-9.
_Measurements._--Average and extreme measurements of four males and
seven females are as follows: Length of head and body, 281 (260-302);
hind foot, 52.3 (50-58); ear, 33.1 (31-37); length of forearm, 173
(168-180). Average and extreme measurements of skulls of three males and
six females are as follows: Greatest length of skull, 73.7 (71.3-77.7);
condylobasal length, 73.1 (70.5-77.4); zygomatic breadth, 40
(36.4-41.5); breadth across first-upper molars, 20.9 (18.3-22.1); length
of maxillary tooth-row, 28.1 (26.9-29.9); length of mandibular
tooth-row, 31.8 (29.7-32.7).
_Remarks._--_Pteropus rayneri grandis_ is the largest subspecies of the
species. It is also the widest ranging subspecies, being found on six
islands (see Fig. 6).
Although the specimens listed above agree well with descriptions of
color given by Thomas (1887_a_:147) and Andersen (1912:259, 263-264),
some individual variation is noticeable. In bats not yet fully grown
(judging from small size, unfused epiphyses, and lack of wear on teeth),
numerous scattered hairs on the sides of the face and crown are buffy.
In adults the face and crown are blackish. With regard to individual
variation in color of mantle and rump patch, specimens with the
following combinations were noted (1) mantle Brick Red, rump patch
bright, basal three-quarters of hairs white, tips Warm Buff (2) mantle
darker, near Hessian Brown, rump patch dark, Chestnut along edges,
center Ochraceous-Tawny (3) mantle Brick Red, rump patch intermediate
between the two other types. Size of rump patch also is variable. In
some specimens it extends onto the upper parts of the thighs whereas in
other specimens it does not.
Sanborn (1931:16) reported an extra tooth, behind the last lower molar,
in a specimen from Choiseul. In one of three specimens in the Bishop
Museum, m3 is lacking. Judging from Troughton's (1936:346) remarks, size
of individuals varies considerably. Specimens that he examined from
Bougainville had longer forearms (up to 177) and larger hind feet
(54-57) than those examined by me from Choiseul. On the other hand,
specimens listed above from Bougainville agree well with those from
Choiseul. In many specimens in the U. S. National Museum, length of the
right- and left-forearm differ. For example, in No. 276926 the right
forearm measures 180 whereas the left is 174; in No. 277098 the right is
172 and the left is 167. Troughton (1936:346) gave standard ear
measurement in _P. r. grandis_ as ranging from 29.5 to 31.5. Ears of
specimens that I examined varied from 31.0 to 37.0.
=Pteropus rayneri rubianus= Andersen
1908. _Pteropus rubianus_ Andersen, Ann. Mag. Nat. Hist., ser. 8,
2:366, October, type from Rubiana; 1912, Andersen, Catalogue
of the Chiroptera ... British Museum, 1:255; 1931, Sanborn,
Publ. Field Mus. Nat. Hist., Zool. Ser., 18:15, February 12,
from Narovo (Simbo).
1954. _Pteropus rayneri rubianus_, Laurie and Hill, List of land
mammals of New Guinea, Celebes and adjacent islands, p. 35,
June 30.
1888. _Pteropus grandis_ (part), Thomas, Proc. Zool. Soc. London,
p. 470, December 4, from Rubiana; 1899, Matschie, Die
Megachiroptera ... naturkunde, p. 15; 1904, Trouessart,
Catalogus Mammalium ..., Suppl., p. 49.
_Specimens examined_ (two males and one female).--Kolombangara, in
February, 23458-60.
_Measurements._--Measurements of two males and one female are,
respectively, as follows: Length of head and body, 253, 265, 251; hind
foot, 53, 50, 50; ear, 30, 31, 32; length of forearm, 158, 161, 160;
greatest length of skull, 70.2, 67.4, --; condylobasal length, 67.0, --,
68.4; zygomatic breadth, 40.0, 39.4, 40.7; breadth across first upper
molars, 19.4, 20.4, 19.9; length of mandible, 53.9, 49.4, 51.3.
_Remarks._--Kolombangara Island is a new locality for _Pteropus rayneri
rubianus_; heretofore this subspecies was known only from Rubiana and
Narovo islands (Andersen, 1908:366; Sanborn, 1931:15). The coloration of
a specimen from Narovo Island was described as between that of _P. r.
rubianus_ and _P. r. lavellanus_. Sanborn (1931:16) allocated it to the
subspecies _rubianus_ on the basis of length of forearm.
Andersen's descriptions (1908:366; 1912:256) of _rubianus_ were of a
specimen stored in alcohol. Coloration of the museum skins examined by
me is as follows: Dorsum from shoulders to rump near Vandyke Brown;
crown and mantle Brick Red; face close to Mummy Brown; rump patch and
thighs close to Warm Buff, strongly contrasting with back and mantle;
base of hairs dark, Seal Brown; venter dark; chest about same as back
but paler laterally (to Ochraceous Tawny); throat Brick Red.
=Pteropus rayneri lavellanus= Andersen
1908. _Pteropus lavellanus_ Andersen, Ann. Mag. Nat. Hist., ser. 8,
2:366, October, type from Vella Lavella; 1912, Andersen,
Catalogue of the Chiroptera ... British Museum, 1:259; 1931,
Sanborn, Publ. Field Mus. Nat. Hist., Zool. Ser., 18:16,
February 12, from Ghizo and Ronongo.
1954. _Pteropus rayneri lavellanus_, Laurie and Hill, List of land
mammals of New Guinea, Celebes and adjacent islands, p. 36,
June 30.
_Specimens examined_ (one male and one female).--Vella Lavella in
November, 23192, 23142.
_Measurements._--Measurements of a male and a female are, respectively,
as follows: Length of head and body, 286, 282; hind foot, 55, 56; ear,
30, 30; length of forearm, 156, 155; greatest length of skull, 72.9,
67.6; condylobasal length, 71.8, 64.2; zygomatic breadth, 38.4, 37.9;
breadth across first upper molars, 19.9, 19.8; length of mandible, 54.6,
50.8.
_Remarks._--_Pteropus rayneri lavellanus_ inhabits islands
geographically near those from which _P. r. rubianus_ is known (see Fig.
6) and in most respects the two subspecies closely resemble each other.
_P. r. lavellanus_ is slightly the smaller (average length of forearm
about 156 instead of 160) and darker. A bat from Narovo [Simbo] Island,
only a few miles from Vella Lavella, identified by Sanborn (1931:16) on
basis of its size as _P. r. rubianus_, resembled the subspecies
_lavellanus_ in color and probably represents an intergrade between the
two populations.
The color of _P. r. lavellanus_ is close to that of _P. r. rubianus_
except that the crown, mantle, and foreneck are near Chestnut-Brown, the
basal portions of hair black, and the fur of the venter, from sternum to
pectoral region, is dark, almost black (compare with description of _P.
r. rubianus_ under account of that subspecies).
Measurements of the male examined are greater than those of the female
studied. Andersen (1912:259) noted that the canine teeth are heavier in
males than in females.
=Pteropus rayneri monoensis= Lawrence
1945. _Pteropus rayneri monoensis_ Lawrence, Proc. New England
Zool. Club, 23:63, March 26, type from Mono (Treasury); 1954,
Laurie and Hill, List of land mammals of New Guinea, Celebes
and adjacent islands, p. 36, June 30.
_Specimens examined._--None.
_Remarks._--_Pteropus rayneri monoensis_ is the most recently described
subspecies of _P. rayneri_. Lawrence (1945:63) judged that in most ways
this bat is intermediate between _P. r. grandis_ and _P. r. lavellanus_.
Coloration of _monoensis_ indicates affinity with the former, whereas
length of forearm (145-148) approaches that in the latter. The small
skull, narrow palate, and whitish rump patch of _monoensis_ are
differences that distinguish it from _grandis_ and _lavellanus_. The
relatively isolated position of Mono Island may have been important in
establishment of the distinctive features of this bat.
Lawrence (1945:65) quoted a collector as stating: "They [individuals of
_P. r. monoensis_] rest quietly during the day in the tops of
heavy-leaved, tall jungle trees, and start flying about dusk, looking
for feeding spots. There is usually quite a flight for fifteen to twenty
minutes at twilight...."
No additional specimens of this subspecies have been collected on small
adjacent islands and _monoensis_ may therefore be confined to Mono
Island.
=Pteropus rayneri cognatus= Andersen
1908. _Pteropus cognatus_ Andersen, Ann. Mag. Nat. Hist., ser. 8,
2:365, October 1, type from San Cristobal; 1912, Andersen,
Catalogue of the Chiroptera ... British Museum, 1:251; 1931,
Sanborn, Publ. Field Mus. Nat. Hist., Zool. Ser., 18:15,
February 12, from San Cristobal and Ugi; 1954, Laurie and Hill,
List of land mammals of New Guinea, Celebes and adjacent islands,
p. 35, June 30.
1962. _Pteropus rayneri cognatus_, Hill, The natural history of
Rennell Island, British Solomon Islands, 4:9, February.
1870. _Pteropus rayneri_ (part), Gray, Catalogue of monkeys, lemurs
and fruit-eating bats ... British Museum, p. 108, from San
Cristobal; 1878, Dobson, Catalogue of the Chiroptera ... British
Museum, p. 33.
1904. _Pteropus_ (_Spectrum_) _rayneri_ (part), Trouessart,
Catalogus Mammalium ..., Suppl., p. 51.
_Specimens examined._--None.
_Remarks._--Specimens of _Pteropus rayneri cognatus_ first were reported
under the name _Pteropus rayneri_ based on three specimens (one from San
Cristobal and two from Guadalcanal). Because the description was based
mostly on the two specimens from Guadalcanal, the name _rayneri_ is
applicable to the bats from that island. Andersen (1908:365) thought
that specimens that he studied, from San Cristobal, were specifically
distinct from _P. rayneri_ and he proposed the name _Pteropus cognatus_
for them. Later, Hill (1962:9) reduced _cognatus_ to subspecific status
under _P. rayneri_.
Presently _P. r. cognatus_ is known only from San Cristobal and the
small adjacent island of Ugi (see Fig. 6).
=Pteropus rayneri rennelli= Troughton
1929. _Pteropus rennelli_ Troughton, Rec. Australian Mus., 17:193,
September 4, type from Rennell Island; 1954, Laurie and Hill,
List of land mammals of New Guinea, Celebes and adjacent islands,
p. 35, June 30.
1962. _Pteropus rayneri rennelli_, Hill, The natural history of
Rennell Island, British Solomon Islands, 4:7, February.
_Specimens examined._--None.
_Remarks._--Until recently, _Pteropus rayneri rennelli_ was known from
but a single specimen. Hill (1962:7) reported two additional specimens
and pointed out that _P. r. cognatus_ and _P. r. rennelli_ probably
represent the extremes of an east-west cline in size. _P. r. rennelli_
and _P. r. cognatus_ differ from other subspecies of the species in
lacking tricolored pelage on the dorsum, but their short rostrum clearly
indicates affinity with other members of this complex group in the
Solomon Islands (Hill, 1962:8).
The relationship of the subspecies _rennelli_ and _cognatus_ is close,
both geographically and genetically. Longer forearm, longer metacarpals,
and longer mandibular tooth-row serve to differentiate _rennelli_ from
_cognatus_.
=Pteropus woodfordi= Thomas
1888. _Pteropus woodfordi_ Thomas, Ann. Mag. Nat. Hist., ser. 6,
1:156, February, type from Guadalcanal; 1888, Thomas, Proc. Zool.
Soc. London, p. 472, December 4; 1898, Trouessart, Catalogus
Mammalium ..., 1:78; 1907, Elliot, Field Columbian Mus., Zool.
Ser., 8:491; 1912, Andersen, Catalogue of the Chiroptera ...
British Museum, 1:410, from New Georgia and Guadalcanal; 1931,
Sanborn, Publ. Field Mus. Nat. Hist., Zool. Ser., 18:19, February
12, from Kolombangara; 1947, Sanborn and Beecher, Jour. Mamm.,
28:389, November 19, from Banika and Guadalcanal; 1954, Laurie
and Hill, List of land mammals of New Guinea, Celebes and
adjacent islands, p. 39, June 30.
1899. _Pteropus (Sericonycteris) woodfordi_, Matschie, Die
Megachiroptera ... naturkunde, p. 83; 1904, Trouessart, Catalogus
Mammalium ..., Suppl., p. 54.
1945. _Pteropus austini_ Lawrence, Proc. New England Zool. Club,
23:59, March 26, from Florida.
_Specimens examined_ (four males and three females; five in alcohol and
two skin-onlys).--Fauro, in April, 23727, 23790; Guadalcanal in May and
June, 23823, 23931; Pavuvo (Russell Islands) in August and October, USNM
277887, USNM 283872-3.
_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.
[Illustration: FIG. 7. Distribution of _Pteropus woodfordi_ ([BW])
and _P. mahaganus_ ([BC]). For names of islands see Fig. 2.]