Notes on distribution and taxonomy of australasian bats. I. Pteropodinae and Nyctimeninae (Mammalia, Megachiroptera, Pteropodidae)

34 Pages·2001·4.2 MB·English

by  Bergmans, Wim

Beaufortia
BULLETIN ZOOLOGICAL MUSEUM
UNIVERSITY OFAMSTERDAM
Vol. 51, no. 8 December 10, 2001
Notes on distribution and taxonomyof australasianbats.
I. Pteropodinaeand Nyctimeninae(Mammalia,
Megachiroptera, Pteropodidae)
WimBergmans
Zodlogisch Museum, InstituutvoorBiodiversiteitenEcosysteemdynamica, University ofAmsterdam,
P.O. Box94766, 1090GTAmsterdam, TheNetherlands
Abstract
Hitherto unreported australasian Pteropodinae and Nyctimeninae from the collections of the Zoölogisch Museum in
Amsterdam and some important samples of Nyctimeninae from the Nationaal Natuurhistorisch Museum (formerly:
Rijksmuseum vanNatuurlijke Historie) in Leiden are reported and discussed. Thepublished first occurrenceof Pteropus
vampyrusonKrakatau Islands afterthe last eruption ispreceded by aspecimenfound thereeightyears earlier. Thefruitbat
material collected onBuru IslandbyL.J. Toxopeus in 1921-1922,asfaras deposited inAmsterdam,isreported forthe first
time.New localities are reported forPteropusgriseusgriseus,P.personatus, P.tonganusgeddiei, P.caniceps,Macroglossusminimusnanus,
Syconycteris australispapuana,Nyctimenealbiventer subsp.,Nyctimene keastitozeri,Nyctimeneaello,Nyctimene rabori,and Paranyctimenerap-
tor. Thefirstcaseof(near-)sympatryofMacroglossusminimusand M. sobrinusonJavaispresented.Nyctimenefrom the Talaud
Islands is identified asN. rabori,notN.cephalotes. Nyctimenefrom New Britain are provisionally referred toN. albiventer andN.
vizcaccia. Thefirst specimen of Nyctimene fromBabar Island, Indonesia,isrecordedand describedasa newsubspecies of
Nyctimene keasti. Paranyctimene is considered a subgenus of Nyctimene. A new species and a new subspecies of Nyctimene
(Paranyctimene)fromPapuaNew Guinea andWestPapua, Indonesia,aredescribed. Parasitic Diptera(Nycteribiidae) werecol-
lected from severalofthe studied fruit batspecies. Theyhave been identified and arereportedin their hosts’ accounts.
INTRODUCTION Jong & Bergmans, 1981; Bergmans & Sarbini,
1985; Bergmans & Van Bree, 1986; Boeadi &
The Zoôlogisch Museum of Amsterdam (ZMA) Bergmans, 1987; Bergmans & Rozendaal, 1988;
has a modest but interesting collection of bats Strien, 1996). However, someof theolder mater-
from Australasia, from Laos to the Solomon ial has remainedunreported, e.g., some fruitbats
Islands, with an accent on Indonesia. Over the from the island of Buru collected by L. J.
years, anumber of reports have been published Toxopeus in 1921-1922.The mammalscollected
on parts of this collection (Jentink, 1890-1891; by Toxopeus were sent fromMuseumBuitenzorg
Beaufort, 1911; Oei & Feen, 1958; Oei, 1960; (the present Zoological Museum of Bogor,
Van Bree, 1961; Bergmans, 1975, 1978, 1979, Indonesia; ZMBI) to the ZMA, from where the
1994, 1995; Rookmaaker & Bergmans, 1981; materialwas sent to MrO. Thomasof theBritish
119
Museum (Natural History) (BMNH) for identifi- imens only. Various criteria havebeen applied to
cation. Some new taxa were described, thetypes establish the adulthood of specimens. Among
of which apparently remained in London, and these, fused and ossified sutures between ptery-
parts of the remainder were sent back to the goid and basisphenoid, and between the latter
ZMA and MZBI, respectively. Dammerman and basioccipital are themost important. See for
(1929) wrote a report on the material in the other criteria Bergmans (1988: 76-77). The fol-
MZBI but the ZMA specimens of this collection lowing abbreviations are used throughout:
have never been reported. ale. = preserved in alcohol; a.s.l. = above sea
Further, through exchange and through contri- level; C = canine; cbl =condylobasal length; E =
butions of interested naturalists, the ZMA has earlength, frombasalnotch to tip; gsl = greatest
continued to receive additional Chiroptera from skull length; HF = length of (hind) foot; imm.=
the region. Together, these collections include immature; M— molar;P =premolar; pdp - post-
somenew,rare,orotherwise interesting taxa,e.g., dentalpalate length; pi = palatal length; skull =
from noteworthy new localities. The representa- skull extracted; W = weight; # = collector's or
tives of the subfamilies Pteropodinae and field number.
Nyctimeninae of the family Pteropodidae among
these are reported here. The Pteropodinae AMNH = American Museum of Natural
include some of the formerMacroglossinae (see History, New York; BMNH = British Museum
Bergmans, 1997). It is the author's intention to (Natural History), London; MZBI = Museum
report the specimens of the remaining Zoologicum Bogoriense, Bogor; RMNH =
Pteropodidae and other chiropteran families in Nationaal Natuurhistorisch Museum, Leiden;
one or more following papers. ZMA = Zoôlogisch Museum, Amsterdam; ZMB
= Zoologisches Museum, Berlin.
MATERIALSAND METHODS
TAXONOMIC SECTION
The ZMA material is listed under the species in
the taxonomic section below. Some important SUBFAMILYPTEROPODINAEGray, 1821
samples of Nyctimeninae in the collection of the
NationaalNatuurhistorisch Museum in Leiden- This subfamily, and especially the polyspecific
the former Rijksmuseum van Natuurlijke genusPteropus Erxleben, 1777, is in needof a full
Historié, and for historical and practical reasons revision, the latest being that by Andersen
abbreviated to RMNH - have also been studied (1912b), who divided the genusinto 17 different
and are listed as such. All measurements are groups.Quite recently, apromising newstudy has
given in mm: body and skull measurements, been started, using sequence data from mito-
taken with callipers, to the nearest 0.1 mm,and chondrial and nucleargenesto construct amole-
teeth - always over the crowns - measuredwith a cular phylogeny for Pteropus (Kershnar et al.,
stereomicroscope with micrometer disc, to the 1999). Thepresent paper presents some interest-
nearest 0.05 mm. For the designation of teeth, ing newspecimens andlocalities for taxonomical-
Andersen has been followed(1912: xxvii-xxviii); ly mostly rather well established species. The
capital I, C, P and M indicate permanent species are treated in the order of Andersen's
incisors, canines, premolars and molars, and the species groups.
position of the tooth number, above or below,
indicates upperandlowerteeth,respectively (e.g., PTEROPUSHYPOMELANUSGROUP
P3 and P3 are the third upper and third lower
premolar, respectively). Weights havebeencopied Pteropus griseus griseus E. Geoffroy - St.
from collectors' labels and are given in g. Hilaire, 1810
Geographical coordinateshave been copied from
collectors' labels (and thenmayinclude seconds), MATERIAL
from atlases, or in some cases calculated. Unless INDONESIA: ZMA23.438/39, 1 male, 1 female,ale., Pulau
statedotherwise, measurements are ofadultspec- Sailus Kecil (1 17°27'E 7°35'S), Kepulauan Tengah
120
(Paternoster Islands), 27-VII-1987, leg. J. de Korte; taken of therelationship between thesetwospecies, and
fromacolony in asmallmangrove forestonthereef. does not conclude their synonymy. Corbet et al.
(1992) however, added the essentially Philippine
REMARKS. - The male has a forearm length of Pteropus speciosus Andersen, 1908to the synonymy
119.1, the female one of 113.1. Corbet & Hill of P.griseus. This was based on the observation
(1992) gave asurvey of thelocalities from where that P.speciosus seems to have been retained only
P. griseus has been reported previously (note that because of a supposed sympatry with P. griseus.
Ujung Pandang and Peleng are mentioned in Corbet et al. wrote "speciosus (Philippines) differs
theirtext but not mapped). Pulau Sailus Kecil is from mimus(Sulawesi) only inblackishrather than
an interesting newlocality record. brownish back and in darker underparts".
Heaney etal. (1998) listed speciosus as aspecies but
DISCUSSION.- Theassignation of thenewmater- do not exclude itspossible synonymywithgriseus.
ial to the typical subspecies is tentative, and
baisedmainly on geographical grounds, as suffi- ECTOPARASITES. - The specimens were infested
cient comparison materialis lacking. with the nycteriibid fly Cyclopodia horsfieldi de
The taxonomy of the species is not entirely Meijere, 1899.
clear. Pteropus pallidus Temminck, 1825 from
Banda Island and P. mimusAndersen, 1908from Pteropus ornatus ornatus Gray, 1870
Ujung Pandang (Makassar), Salayer and Peleng
are generally regarded as - atmost - subspecies of MATERIAL
P. griseus (Laurie & Hill, 1954; Goodwin, 1979; NEWCALEDONIA: ZMA 11.717, 11.719, 1 male, 1 female,
Bergmans et al., 1988; Corbet et al., 1992; ale., incomplete skulls, near Thio (21°37'S 166°13'E), 17-
Mickleburgh, et al., 1992; Wilson & Reeder, III-1968,leg. P Guépig.
1993; Koopman, 1994), although there appears
to be somedoubt as to the allocation andprove- REMARKS. - The male appears to be a young
nance of what has been described as P.pallidus. adultand has a forearm length of 142.6and an
Goodwin (1979) wrote that a subspecies of P. uppertoothrow length of 25.2. Thefemale hasa
griseus "possibly" occurs on the Banda islands, forearm length of 139.6, a gsl of 62.4, and an
andFlannery (1995a) doesnot listP.pallidus atall. uppertoothrow length of 24.4.
For their paper on small Philippine Pteropus
species, Klingener & Creighton (1984) studied DISCUSSION. - Both specimens are adulttojudge
series of P. speciosus and P. griseus, among other by their fused skull base sutures, but they are
species. Although their paper is centeredaround small.Felten(1964) gave 144-163for theforearm
somesynonymies, it doesnot containa discussion length range in 22 specimens from New
')Thementioned studyby Klingeneret al.(1984)isstrongly influenced by thepeculiarhypothesis that,astheywrote,
the "Taxonomic studyof megachiropteran batsiscomplicatedbythe factthat the skull continuesto grow indimensions
and shape throughoutlife" or, moreprecisely, "In megachiropterans theskull continues toelongatein adults.This elon-
gation may involve backward and dorsal movementofthe braincase aswell aselongation ofthe rostrumin long-faced
forms."Klingeret al.mention twostudies (Lanza, 1961;Peterson &Fenton, 1970)which aresuggested tobe supportive
of this hypothesis. But neither is.Lanza (1961), who describedpost-embryonal skull growth in two African fruit bat
species, literally concluded that in Epomophoruslabiatus (Temminck, 1837), and implicitly also in E.wahlbergi (Sundevall,
1846), the dimensional development oftherostrum stops at acertain stage (Lanza, 1961: 183). Peterson etal. (1970)
describedthe cranialvariationwithage inHarpyionycteris Thomas, 1896,involving several characters, but in nowaysug-
gestedthatskulls offruitbats continue growing 'forever'. In fact, asexperience shows, there is aratherrestrictedrange,
with both aminimum and amaximum value, ofthe greatest skull length(and the condylobasal length)in adultmales,
and most frequently adiflerent butequallyrestricted range in adult females,in every species offruitbat. Accumulated
measurementsofadultspecimens ofeithersexfromagivenpopulationshow anormalstochasticdistribution.Ithasnever
been substantiated that,asKlingener et al.(1984)seemed tobelieve, skull growth continuesforever.Iftheywereright,
everynow and then anewmaximummeasurementwouldberecorded forevery specific skull dimension. But the uncer-
taintiesareofaless Olympic character.The processofstretchingof the skull in Megachiroptera, involving adecreasein
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Caledonia, sexes combined, 66.7-72.6 for the gsl specimens from the Fiji Islands, and are too few
range in 16of thesespecimens, and24.7-28.0 for to beconclusive.
the upperteethrow length range in 23. Flannery
(1995a) gave 165 as forearm length forone male PTEROPUSCANICEPSGROUP
and 142-154for three females.
Pteropus caniceps Gray, 1871
ECTOPARASITES. - Onespecimen of thenycteribi- MATERIAL
id fly Cyclopodia similis Speiser, 1900was foundon INDONESIA: ZMA21.433, 1 male, ale., skull, Ngele Ngele
the malespecimen. Besar (2°H'20"N 128°12'20"E), 16-XI-1980,leg. H. Moll,
mistnettedundertrees; ZMA22.134, 1 imm. female, ale.,
PTEROPUSMARIANNUSGROUP skull, Bataka (Talaga), Gamkunora (Gunung Gamkunoro:
01°30'N 127°32'E), northwest Halmahera, 3/4-III-1983,
leg.F.G.Rozendaal.
Pteropus tonganus geddiei MacGillivray,
1860
REMARKS. - This species is restricted to
Halmaheraand surrounding islands, from where
MATERIAL
NEW CALEDONIA: ZMA 11.716, 11.718, 1 female, 1 imm. it has beenreported from Bacan, Morotai, and
male, ale., incomplete skulls, near Thio (21°37'0"S 166° Ternate, and to which Ngele Ngele Besar is now
13'00"E), 17-III-1968,leg. P Guépig. added, and from Sula Bessi, Sula Islands (Corbet
etal., 1992).(See Bergmans etal., 1988, fora dis-
REMARKS. - The present female specimen has a cussion of the species' disputed occurrence in
forearm length of 168.0and large nipples. The Sulawesi and Peleng Island.) Theadult male has
immature malehas a forearmlength of 145.2. aforearmlength of 138.3, agslof 68.8, andlarge
testes; the immaturebut nearly adultfemalehasa
DISCUSSION. - Felten (1964) revised Pteropus from forearm length of 129.5, a gsl of 59.4 and a
New Caledonia, and concluded that P. tonganus weight of 267.
fromthatislandrepresented thesubspecies geddiei.
He listed anumberof measurements for thetwo ECTOPARASITES. - The specimen from Ngele
sexes combined. For 16 specimens, from both Ngele Besar yielded one specimen of the nyc-
NewCaledoniaand the Loyalty Islands, he gave teribiid flyEucampsipoda inermis Theodor, 1955.
a forearm length range of 147-166.The female
specimen surpasses this range, which suggests PTEROPUSMELANOPOGONGROUP
that females may attain larger forearm lengths
than males. However, Flannery (1995a) stated Pteropus melanopogon Peters, 1867
that in this species males are considerably larger
than females, withmales maturing at aweight of MATERIAL
600 and females at 450. The forearm lengths in INDONESIA: ZMA 16.368, 1 female, skin, skull, Wa'tra,near
his account (148.9 in one male, and 144-145 in Leksula (03°46'S 126°35'E), Buru, 3-III-1921, L. J.
Toxopeus.
three females) do not include geddiei but typical
what Andersen (1912) called braincase deflection (and entailing, among otherthings, adecrease in relative postorbital
width),isnotaprocess takingplacein adults,butin maturingindividuals. Strongbraincase deflectionis generallyajuve-
nile condition, which in somespecies and tosomedegreemaypersistasaneotenecharacter in adultlife, andthe mea-
sureofthis deflection inadults dependsonthespecies involved (see also Bergmans, 1977 and 1994:81). The measureof
skull bone fusion provides agoodindicationof the growthstage of the skull involved. When all sutures arefused and
have become invisible, in which process the basi-cranial sutures most usually are the last onestoclose, onecan safely
assumethat the skull is full-grownand thatits growthhad stoppedatthe time ofcollection. Skull parts which may con-
tinue to grow afterthis stage has been achieved are the sagittal and occipital crests, and possibly othercomparable fea-
tures,butthesedonotchange the essential form oftheskullassuch. This late growthoftheoccipital crest (whichisalso
notcontinuous) may contributetothegreatestskull length,butitdoesnot growforevereither.Toavoidthe effectofoccip-
ital crest growth ongreatest skull length,many authorsprefer thecondylobasallengthasameasureof skullsize.
122
REMARKS. - The specimen has a forearm length ly from what Andersen published in 1912: 196-
of about 213, and a gsl of over 78.8. The type 204forthreespecimensof melanopogon (fromthree
locality of this species is Ambon, and it has fur- differentislandsand of unknownsex but for one
ther been recorded from Banda, Goram (also female), 190-191 forone maleanda specimen of
spelled Gorom = Gorong), Buru, Seram (as unknownsex of aruensis, and 179-187.5for two
Ceram), Boano, Saparua, Manawoka (as males and two specimens of unknown sex of
Manavolka), Siao, TimorLaut(= Tanimbar), and keyensis. Only Flannery (1995a) mentions new
Taâm. Bergmans et al. (1988) re-identified the specimens, amalekeyensis from Taâm, intheKey
material from Siao reported as P. melanopogon by group, with a forearm of 175, and a female
Jentink (1888) and Laurie etal. (1954) as Pteropus melanopogon from Buru, with a forearm length of
hypomelanus Temminck, 1853. 193.
Synonymizing taxa withoutother than implicit
DISCUSSION. - Flannery (1995a) questioned the arguments, in this case on the one hand
localities Boano, Saparua and Manowoka. They Andersen'sremarks (1912) on the relationship of
were first mentioned by Peters (1868), who did theseforms,andon theotheranapparent wishto
not indicatethematerialonwhichthis wasbased. simplify taxonomy by recognizing lowernumbers
Peters' basis was the Zoologisches Museum in of taxa, is not acommendableprocedure. If any-
Berlin, but he certainly visited and examined thing, it unnecessarily burdens the literaturewith
other collections. In the Berlin museum, unfoundedor insufficiently founded and there-
Andersen (1912) found only one specimen, the fore as such unacceptable taxonomicpropositions
type from Ambon, but he stated to have exam- whichnevertheless haveto be taken into account
ined a specimen from Saparua in the RMNH. by laterstudents. For this reason, and for as long
Even if we cannot trace Peters' material from as anewanalysis of existing andnewmaterialhas
Boanoand Manowokatoday, these localities are not shownotherwise, the present authordoesnot
very probable for the species, and should not be followLaurie etal. (1954) and considersP.melano-
doubtedwithoutreasons. pogon as amonotypic species, and P.aruensisandP.
Andersen(1912)pointed out thatPteropus aruen- keyensis as indépendant species. Itis interesting to
sis Peters, 1868and Pteropus keyensis Peters, 1868, note that no author after Andersen (1912) has
both described by Peters as 'varieties' of melano- examined the material from Tanimbar in the
pogon, were related species, but Peters and museum in London, and that we do not know
Andersen also described in what characters they where this wouldfit in.
differed. Laurie et al. (1954) listed both aruensis Finally, Flannery (1995a) dwells extensively on
and keyensis as subspecies of melanopogon, and thewhereaboutsofWonambay, one of thelocal-
although they did not give any reason for doing ities where the type or type series originated,
so, they were followedin this by alllaterauthors. which he quotes from Laurie et al. (1954) and
The two were originally differentiated on the which he hesitatingly identifies with the modern
basis of fur colours (Peters, 1868). Another ManumbaiSungai. Strien (1996), writing on the
apparent difference between the mentioned mammals of the Aru Islands, presented a map
forms is in size. One would expect a further indicating mammalcollectors, journeys andloca-
analysis of these and possible other differential tions, including Wanumbai (one of the several
characters, but what strikes one when going spellings ofthis name).
through the post-Andersen literature on the
species, is the lack ofany analysis at all, and the PTEROPUSRAYNERIGROUP
very small number of measurements or other
characters thathavebeenpublished. Andindeed, Pteropus chrysoproctus Temminck, 1837
very few specimens seemto have been collected
Material
and studied since Andersen wrote his account
INDONESIA: ZMA 3092, 1 imm. (female?), skin, skull, Tifu
(1912). For example, Koopman (1994) mentioned
(03°41'S 126° 24'E), Huru, 1922,leg. L..J. Toxopeus; ZMA
as forearm length range in Pteropus melanopogon 16.369/70,1 male, 1 female,skins, skulls, Buru, 1921-1922,
sensu lato 179-204.This has been compiled sole- leg.L.J. Toxopeus.
123
REMARKS. - Themaleof thesetwohas aforearm Toxopeus from Buru to the same zoo, where it
length of 166anda gslof 73.6, thefemaleafore- died on 10-VI-1923.This animal has a forearm
arm length of 170anda gslof 76.5. Thisis with- length of 105 and a gsl of 52 (Dammerman,
in theknown dimensionalranges. 1929). It bears number 2155but its precise col-
lecting locality is apparently unknown.Itproba-
DISCUSSION. - The specimen from Tifu was sent bly is in the MZBI.
alive to NaturaArtis Magistra, thezoological gar-
denofAmsterdam, whereit diedon 13-X-1922. Pteropus personatus Temminck, 1825
As it is subadult, it has probably been collected
and sentearlier in 1922.The labels of the other MATERIAL
two specimens bear no other information than INDONESIA:Photographsofafemalewithajuvenile clinging
"Toxopeus, Buru" and unfortunately more pre- toher,caughtand released atNgeleNgele Besar (2°11'20"N
128°12'20"E), 16-XI-1980,by H. Moll; ZMA 22.107, 1
cise data could not be derived from Toxopeus'
male,ale., Bataka (01°24'N 127°30'E),northwest Halma-
itinerary of his expedition (Toxopeus, cf. 1924)
hera,27-111-1983,leg. F.G.Rozendaal,# 179.
nor from Dammerman'sreport (1929) on anoth-
er partof Toxopeus' mammalmaterial.
REMARKS. - The photos of the specimen from
PTEROPUS TEMMINCKIIGROUP Ngele Ngele Besar, an island off Halmahera,
made by the late Dr H. Moll, are very distinct
Pteropus temminckii temminckii Peters, and leave no doubt as to the identity of the
1867 species. Ngele Ngele Besar is a new locality
record for the species. Flannery (1995a) reported
lactating andpregnant femalesonTernate Island
MATERIAL
in January. The juvenile in the photo is quite
INDONESIA: ZMA 3084, 1 female, ale., skull, Ambon
(03°41'S 128°01'E),1913,leg.WillemszGeerooms. large. Somefieldmeasurements of themalefrom
Bataka are: forearm length 97.8, E 23, and W
REMARKS. - This specimen first entered the col- 131. This specimen was caught in the daytime
lection of the Colonial Institute (now Royal from aroost inapapawtree plantation.
Institute for the Tropics) inAmsterdam, andwas
deposited in theZMA inthe 1930s. Ithas a fore- DISCUSSION. - Bergmans etal. (1988) pointed out
arm length of 100.6 and a gsl of 50.4. Dr H. that the alleged occurrence of this species in
Felten identifiedthe specimen as arepresentative Sulawesi is basedon two specimens of uncertain
of the typical form. provenance. They proposed to reject these
records as prove that the species was foundon
Pteropus temminckiiliops Thomas, 1910 that island. Following this, Mickleburgh et al.
(1992) added a question mark to its occurrence
MATERIAL there; its presence in the Dumoga Bone National
INDONESIA: ZMA3080, 1 female, skin, skull, Buru (03°27'S Park in North Sulawesi as recordedby the same
126°40'E),1921-1922,leg.L.J. Toxopeus. authorsmust thereforebe basedonan error.
PTEROPUS VAMPYRUSGROUP
REMARKS - Mr.Toxopeus sent this specimen first
to the zoological garden in Amsterdam, Natura Pteropus vampyrus vampyrus Linnaeus,
Artis Magistra, whereit diedon 15-VIII-1922.As 1758
it is immature, witha forearmlength of92.6 and
a gslof 44.9, itcan beassumedto havebeensent MATERIAL
at most some months earlier in 1922. Dr H. INDONESIA: ZMA 19.354, 1 female, found dead,wing,foot
Felten identified the specimen as Pteropus tem- and skull in collection, at edge of Krakatau, Krakatau
Island,29-VII-1977,leg.J. Regout and donatedbyP.J. van
minckii liops.
Nieuwenhoven; ZMA 21.637, 1 male, ale., Botanical
Dammerman (1929) reported on another Garden, Bogor (06°34'S 106°45'E), 30-XII-1981, leg. W.
female of this species, as Pteropus liops, sent by Bergmans.
124
REMARKS. - Unfortunately, it has not been docu- REMARKS. - Thisspecimen has a forearm length
mentedon whichof the fourparts of the former of 169.7, which adds nothing to the known gen-
Krakatau Island the remains of the female have eral range but supports the observation by
been found. The skull is broken; the forearm Bergmans etal. (1988) thatspecimens fromNorth
length is 182.8, E 36, and HF 55. Sulawesi may be smaller, onaverage, than speci-
mens from south-west Sulawesi.
DISCUSSION. - Thorntonetal. (1988) reported on
an observationof this species on Anak Krakatau DISCUSSION. - The provenance of this and other
in 1986,whichthey consideredastray individual. specimens would suggest that in the region of
However, in 1985 Tidemann et al. (1990) Imandi fruit bat species are frequently trapped
observed a camp of about 250 individuals on for consumption. However, in a recent book on
Sertung. The present specimen preceeds these his experiences in Sulawesi and other islands,
finds with eight years. Together, these occur- Argeloo (2001) reports on fruit bat hunting in
rences suggest that the species may have flown western Minahassa for the market in Manado.
out to Krakatau earlier in the island'srecent his- According to hisobservations, nohuntablequan-
tory, andmoreoften, andmayhavebeenof more tities of fruitbats remainbetweenManado and
importance as disperser of seeds to the island(s) Marisa (at 121°56'E on the south coast of
than the known record wouldsuggest.-^ Minahassa), which would suggest thatat present
The specimen from the Botanical Garden at the provenanceof fruit bats intheImandimarket
Bogor, home to awell-knownprotected colony of is also "west of Marisa, northern Sulawesi". It
this species, hadjust been shotby two guards in furthermore implies ableak futureforthe North
the very early morning, when the author stum- Sulawesian endemic fruit bat Rousettus bidens
bleduponthem; forearmlength about213, E 44, (Jentink, 1879),whichis extravulnerablebecause
W855.Thisis proof thatthespecies ishunted for it lives in large colonies in caves, the north
foodinJava, whileanotherapparent threat to this Sulawesian populations of the rare Neopteryxfrosti
subspecies is the very advanced deforestationin Hayman, 1946, and other Sulawesianendemics.
Java. As such, neither of these threats to this
species appears to havebeen documentedbefore PTEROPUSCONSPICILLATUSGROUP
(see Mickleburgh et al., 1992).
Pteropus conspicillatus chrysauchen
PTEROPUSALECTOGROUP Peters, 1862
Pteropus alecto Temminck, 1837 MATERIAL
INDONESIA:ZMA22.133, 1 imm. female, ale., skull, Bataka
MATERIAL (01°24'N 127°30'E),northwestHalmahera,27-111-1983,leg.
INDONESIA: ZMA23.081, 1 male,ale., boughtat market at F.G.Rozendaal,mistnetted insecondary growthin coconut
Imandi (00°35'N 124°04'E),North Sulawesi, 1985,leg. R. plantation; ZMA 22.135, 1 imm. female, ale., 16-V-1983,
W.R.J. Dekker. island near Labuha (00°35'S 127°28'E), Bacan, leg. F. G.
Rozendaal.
2)In thisconnection,anotheraspect ofbats asseed dispersersshouldalsobe mentioned. Whittaker&Jones(1994),writ-
ingontherole offruitbats in therebuildingof the forestecosystem onKrakatau, and someauthorsbeforethem,appear
totake forgrantedthat seedsare dispersedby fruitbats overlarger distancesonly afterincidental ingestion ofsmall seeds
atoneplace and their subsequent defecation or oral ejection at another. Food does notstay longinthe intestinaltract
(between 15 and 70 minutes, asWhittaker et al. quote),which effectively restricts the successofseed dispersion in this
way. However,onmore thanoneoccasionthepresentauthor hastrapped fruitbats with small seeds sticking totheirfur,
mostprobablybymeansofthe sugaryjuice ofthe fruitsinvolved. Thisnaturallycomesaboutwhile eatingand the seeds
will normallyprobably stay ontill the bats arrive back attheir dayroost, where extensive grooming will end the seeds'
voyage. But itis notimpossible, and in factquitelikely, thatbats carryseedsin thiswayfromonefeeding placeto anoth-
er, and thatthey mayincidentally losetheseseedswherever theyfly oralight.
125
REMARKS. - The present specimen from falls within the known size range for thespecies.
Halmahera has a forearm length of 148.7 (field
measurement: 151), agsl of66.7, andaweight of Pteropus vetulusJouan, 1863
420. Its skull base androstral sutures are distinct
and the specimen is obviously immature. On the MATERIAL
other hand, its skull is nearly full-grown, and its NEW CALEDONIA: ZMA 19.311, 1 male, ale., Blue River
fingerjoints as well.Theotherspecimen is asuck- Reserve (now the Provincial Blue River Park;
22°04722°10'S I66°367166°4RE),3-IX-1967, leg. H. L.
ling, with its milk dentition still in place and its
Bregulla.
permanent canines just breaking through, and a
forearm length of99.3.
REMARKS. - The specimen has a forearm length
DISCUSSION. - Although known since long from of 103.8 and a W of 155. Felten (1964) gave a
the surrounding islands of Bacan, Morotai, Obi, forearm length range of 100-114(mean 106) for
Ternate(Andersen, 1912) and more recently also 14specimens, including the type, but didnot dis-
from Gebe(Corbet et al., 1992), this species has tinguish the sexes; he gave no data on weights.
been recorded for the principal island of the Flannery (1995a) gave as forearm length range
group, Halmahera, only by Peterson et al. in for three males 92.3-100.8 and as weights 120-
1990. The island olf Labuha, Bacan is a new 142.
locality. Itis clear thatthe forearmlength range
for P. conspicillatus in the Moluccas given as 175- Styloctenium wallacei(Gray, 1866)
185 by Corbet et al. (1992), who based this on
dataprovided by Andersen(1912), is incomplete MATERIAL
and inneedof modification. Thepresent author INDONESIA: ZMA 24.256, 1 male, ale., bought at market,
Imandi (00°35'N 124°04'E), North Sulawesi, leg. M.
assumes that in this species males grow larger
Argeloo,9-1-1991;ZMA24.444/46,3males,samedata, 10-
thanfemales,withpossible forearmlength ranges 11-1992;ZMA24.913/15,2 males, 1 female,samedata, 6-
of about 155-174 in females and 163-185 in VII-1994.
males. The ear in thespecimen from Halmahera
is 29 in length (field measurement: 32)and rather REMARKS. - These specimens were presented to
pointed, withanincurvation inthe posteriormar- the ZMA after the publication of the report on
gin, just under the tip. In this respect also, the Sulawesi fruit bats (Bergmans etal., 1988). The
Pteropus group table in Corbet et al. (1992: 57) males have forearm lengths of 92.0, 92.6, 94.0,
needs modification. 99.8, 99.9 and 103.2, respectively. The first three
are below the known range of 95.5-103.2
ECTOPARASITES.- Thespecimen from Batakawas (Bergmans et al., 1988: 30). The female has a
forearm length of 98.0 which falls within the
infestedwith the nycteribiid fly Cyclopodia albertisii
known
Rondani, 1878, and possibly also with range.
Archinycteribia actena Speiser, 1901, of which one
specimen was foundin the bat's container. DISCUSSION. - On the provenance of bats from
the Imandimarket see underPteropusalecto.
PTEROPUSSCAPULATUSGROUP
Neopteryx frosti Hayman, 1946
Pteropus woodfordi Thomas, 1888
MATERIAL
MATERIAL INDONESIA:ZMA24.258, 24.406,2 females,ale., bought at
SOLOMONISLANDS: ZMA11.251, 1 male,ale., nearBelaga, the market ofImandi (00°35'N 124°04'E),NorthSulawesi,
Small (or Little) Gela (09°10'S 160°20'E),Nggela Group, 21-1- and 29-V-1991, respectively, by M. Argeloo; ZMA
VII-1966/IX-1967,leg. M.J.A. deKoster. 24.448, 1imm.male,ale.,sameprovenance, 10-11-1992,leg.
M.Argeloo.
REMARKS. - The specimen is a youngadult and REMARKS. - Neopteryxfrosti is very rare in collec-
witha forearmlength of 93.2 anda gslof42.7 it tions, the present specimens being the 5th to 7th
126
of this species onrecord (Bergmans et al., 1988). (Bhutan? See the map in Corbet et al., 1992),
One female has a forearm length of 106.5, the Burma, Central and South Thailand, southern
other one of 111.0, which enlarges the known Laos and Vietnam, Cambodia, Peninsular
variation range in females from 110-110.6 to Malaysia, Sumatra, Java and Bali; specimens
106.5-111.0.The immaturemale has a forearm from Baliare smaller in somerespects than those
length of 98.1. from Java (Kitchener & Foley, 1985). A second
subspecies,fraternus Chasen & Kloss, 1927inhab-
BIOLOGY. - The two females were pregnant. The its Sipora, Siberut and MentaweiIslands.
onebought on 21January, forearmlength 106.5, For thepresent paper,Hill ( 1983) andCorbetet
hadan embryo witha greatest length of 21, and al. (1992) have been followed.
the onebought on29May, forearmlength 111.0,
had an embryo with a greatestlength of 48.5. Macroglossus minimus minimus
(Geoffroy, 1810)
DISCUSSION. - On the provenance of bats from
the Imandimarket see underPteropus alecto. MATERIAL
INDONESIA: ZMA 16.706a, 1 male, ale., skull, Jakarta
MACROGLOSSUSCUVIER, 1824 (06°08'S 106°45'E),Java, 1908,leg. PN. vanKampen.
Hill (1983) revised the taxonomy ofMacroglossus, REMARKS. - The skull of this specimen is dam-
and retained two species: the small M. minimus aged, its occiput being brokeninto several pieces.
(Geoffroy, 1810) and the somewhat larger M. Its gsl is probably not much more than26.4(and
sobrinus Andersen, 1911, each with several sub- its cbl between 1 and 1.5 shorter). Some other
species. The taxonomy and distribution of this measurements are: distance from orbit to nare
genus were further amended in Corbet et al. 8.75, C'-M2 8.3, and CrM3 10.0. The fused
(1992). sutures of the skull baseand thelarge, descended
InM. minimus, five subspecies were retained by testesprove the specimen tobe adult.Its forearm
Hill in 1983: minimus; lagochilus Matschie, 1899; length is 41.3. The internarial groove is linear,
nanus Matschie, 1899; pygmaeus Andersen, 1911; weak, and does not run downto the lip margin.
and microtusAndersen, 1911 - withbothpygmaeus The dimensionssuggest thatthe specimen repre-
and microtus admittedly possible synonyms of sents the species minimus, the internarial groove
nanus,as McKean(1972) hadsuggested. In 1992, would be atypical (Hill, 1983), but see also
Corbet et al. retained three subspecies only: Kitchener et al. (1985), who describe specimens
lagochilus, minimusand nanus,and suggested more- from Bali combining characters of both minimus
over that lagochilus and minimusare hardly distinct andM. sobrinus.
andmayprovesynonymous.Of thesesubspecies,
minimusis foundinJava, Madura, Bali, Kangean DISCUSSION. - Another adult male from Jakarta
Islands and Lombok (see for Lombok: (ZMA 21.621) has beenidentifiedas M. sobrinus;
Mickleburgh et al., 1992); nanus (including pyg- it has a forearm length of 46.0. So have two
maeus and microtus) is found in New Guineaand femalesfrom Jakarta (ZMA 21.624, 21.626) with
surrounding islands, Queensland, Murray Island forearm lengths of 45.5 and 47.2, respectively.
in the Torres Strait, Bougainville, and Solomon The internarial grooves of these specimens are
Islands; and lagochilus would inhabit Buru (the linear and weak, and they do not reach the lip
type locality) and otherMoluccaIslands, butalso margin. This appears to be the first instance of
Indochina(Thailand, Vietnam), Nias (listed but actual sympatry (although Jakarta still covers a
not mapped by Corbet et al., 1992), Sri Buat huge area) of minimusand sobrinusinJava.
Island, Sirhassen Island; Bunguran Island,
Borneo, Philippines; Sulawesi; Peleng; Sanghir Macroglossus minimus lagochilus
Islands, Timor(see Hill, 1983: 135). Matschie, 1899
For M. sobrinus, the picture is more simple: the MATERIAL
typical subspecies is found in northeast India INDONESIA: ZMA 16.705, 1 male, ale., Gunung Sitoli
127
(01°16'N97°34'E), Nias Island, 1910,leg. J. P. Kleiweg de ZMA 16.702, 1 male, ale., Sumanik, near Singkarah (or
Zwaan; ZMA22.114/21, 6 males, 2 females, ale., Telaga- Singkarak: 00°41'S 100°38'E), Sumatra, 1888, leg. M.
Gamkunora (or Gamkunoro:01°30'N 127°32'E),northwest Weber; ZMA 16.703/04, 2 females, ale., (probably
Halmahera, 3/4-III-1983, leg. F. G. Rozendaal; ZMA Sumanik), Sumatra, probably 1888,leg. M. Weber; ZMA
22.122, 1 male, ale., lowerslopes of Sibela range (00°4'S 16.706b/07, 1female, 1 imm.female, ale., Salatiga(07°15'S
127°35'E), northeast of Ngame, northwest of Wayaua, 110°34'E), Java, before 1923, leg. D. de Lange; ZMA
Bacan (127°30'E00°35'S), 9-V-1983,leg.F. G. Rozendaal. 21.621/26, 3males, 3 females, ale., Paneoran,Jakarta (06°
08'S 106°45'E), Java, 22- and 23-XII-1981, leg. W.
REMARKS. - The specimen from Nias hasa fore- Bergmans; ZMA21.627/30, 1 male, 3 females, ale., Bogor,
Java, 24/29-XII-1981, leg.W.Bergmans; ZMA21.888/89,
arm length of 39.7. Six males from Halmahera
1 male, 1 female, ale., near Parapat (02°43'N98°58'E),
and Bacan have forearmlengths of 37.3 to 39.0 Sumatra, 21/22-X-1982, leg. P. J. H. van Bree; ZMA
(mean 38.3), the two females of 37.5 and 39.0, 21.890, 1 female, ale., base camp GunungLeuser National
suggesting thatthepopulations from thenorthern Park, at river Alas, opposite Ketambe (03°4I'N 97°38'E),
Moluccasarerelatively small. Theiridentification Sumatra, 25/26-X-1982, leg. P J. H. van Bree; ZMA
as lagochilus is given here with reservation, nanus 22.500/04, 2 males, 3 females, ale., Ubud (08°30'S
115°18'E),Bali, 2/10-11-1985,leg. RJ. H. van Bree;ZMA
being the alternative.
23.866/67, 1 male, 1 female, Banjar Alum, Gilimanuk
(08°12'S 114°27'E),Bali, 27-IV- and XI-1988,respectively,
Macroelossus minimus nanus Matschie, leg.B.E.van Helvoort.
1899
REMARKS. - Forearm lengths and knownweights
MATERIAL
(between brackets) of the adults among thepre-
INDONESIA: ZMA 2695, 1 male, spirit, Jayapura (02°37'S
sent specimens are as follows.Sumatra: two males
140°39'E),WestPapua, 15-11-1956,leg.F.Hoekzema; ZMA
2694, 1specimen,sexunknown,ale.,Jayapura,WestPapua, 46.8-48.8, four females fal 44.2-48.8; Java: two
25-VII-1957, leg. F. Hoekzema; ZMA 2307, 1 male, skin, males 46.0-47.1 (W 22.5-23), six females 44.8-
skull in situ, Jamas village (ca. 05°17'S 138°04'E),Asmat 47.2(W of 5 19-22,5); Bali: fourmales44.7-48.0,
area, WestPapua,8-VII-1958,leg.W.R. vanMourik;ZMA threefemales 44.3-48.1.
24.948/52,2males, 3females,alc.,Jirlai village (ca. 06°01'S
134°2'E),Kobroor Island,Aru Islands,IV-1993,leg. M.van
der Wal; ZMA 24.954/57, 4 males, ale., Baun Island DISCUSSION. - The specimens from Bali show all
(06°30'S 134°40'E),Aru Islands, 21-IV-1993, leg. M. van sorts of variation in depth and length of the
der Wal; ZMA 25.429, 1 male, ale., 200 m from Rumei internarialgroove.Infourof theseven specimens
River, ca. 10 km E of Urbinasopen village (Urbinasopon;
00°22'S 131°13'E)and ca. 7 km upstream,alt. 200-250 m, the grooveruns to thelip margin, whichwouldbe
southeastWaigeo,WestPapua, 13-III-1993,leg.M. Argeloo. acharacterof minimus(Corbet etal., 1992),inthe
others it doesnot. Thesamevariationhas already
REMARKS.- Thespecimens from theAru Islands been noted by Kitchener et al. (1985), whoalso
haveforearmlengths of 38.3-40.3(mean 39.1) in allocated their Bali specimens to M. sobrinus
four males and 38.3, 39.4 and 41.0 in three ratherthan minimus.
females. The male from Waigeo has a forearm
length of 41.3and a weight of 17 g. Itpresents ECTOPARASITES. - Specimens ZMA 21.621 and
the first record for that island. The male from 21.629, from Jakarta and Bogor, respectively,
Jayapura has aforearmlength of 40.2.Thespec- each carried a nycteribiid fly, Cyclopodia
imenfromJamaswas founddeadin ahouseafter (Cyclopodia) tenuis Schuurmans Stekhoven &
it had flown into it. Jamas lies inland from the Hardenberg, 1938.
mangrove zone. The forestconsists ofbroadleaf
trees andPandanusand sago palms. Syconycteris australis papuana(Matschie,
1899)
Macroglossus sobrinus sobrinus
Andersen, 1911
MATERIAL
INDONESIA: ZMA 24.953, 1 female, ale.,Jirlai village (ca.
MATERIAL 06°01'S 134°29'E),Kobroor Island, Aru Islands, IV-1993,
INDONESIA: ZMA 2047, 1 imm. female, skin, skull, Bogor leg. M. vander Wal, mistnetted at 19.00 pm, nearpapaya
(06°34'S 106°45'E),Java, 23-VI-1948,leg. T.vanBemmel; trees;ZMA24.958/59, 1 male, 1 female, ale., Baun Island
128