Taxonomy and biogeography of African fruit bats (Mammalia, Megachiroptera). 4. The genus Rousettus Gray, 1821

48 Pages·1994·3.8 MB·English

by  Bergmans, Wim

Beaufortia
INSTITUTE FOR SYSTEMATICSANDPOPULATIONBIOLOGY
UNIVERSITYOF AMSTERDAM
Vol. 44, no.4 November 18, 1994
Taxonomyandbiogeographyof African fruit bats
(Mammalia, Megachiroptera).
4. ThegenusRousettus Gray, 1821
WimBergmans
InstituteforSystematics andPopulation Biology, University ofAmsterdam, P.O. Box 94766,
1090 GTAmsterdam, theNetherlands
Abstract
The concept ofthe genus Rousettus Gray, 1821 asestablished by Andersen (1912) is revised to accommodate R. madagas-
cariensisGrandidier, 1929, R. obliviosus Kock, 1978 and R. spinalatus Bergmans & Hill, 1980, and to reflect the following
mutations.Following Bergmans etal. (1988), thegenus Boneia,Jentink, 1879 istreated asasynonym ofRousettus, whichadds
to that genus the species B. bidensJentink, 1879. The subgenus Stenonycteris Andersen, 1912 is considered asynonym of
Rousettus, and thesubgenusLissonycterisAndersen, 1912 afullgenus, to bereviewedinthenextpartofthisseries.AllAfrican
Rousettus speciesarecharacterizedand their distributions, includingmany newrecords,and geographicalvariation areana-
lyzed. Following Eisentraut (1960), R. egyptiacus (E. Geoffroy-St. Hilaire, 1810)is dividedinto four,geographicallydisjunct,
subspecies: egyptiacus from southern Egypt northward to and alongthe Mediterranean coast,including Cyprus, into south-
ernTurkey; leachii(Smith, 1829)fromsouthern Sudan and Ethiopia throughEast Africa tothe Cape;unicolor(Gray, 1870)
fromnorthwestAngolatotheMountCamerounregionand from theretoSenegal; and the extralimital arabicusAnderson
& de Winton, 1902 fromthe southern halfofthe ArabianPeninsula tosouthern Iranand Pakistan(arabicusrecords from
East Ethiopia are thoughttobe based onsmall-skulled leachii). All foursubspecies areshown tovary geographically.The
differences between arabicus and R. leschenaultii(Desmarest, 1820), which bothinhabit Pakistan, arediscussed atlength.
Notwithstanding several recent publications, Rousettuslanosus O. Thomas, 1906 isconsidered very distinctfrom Rousettus
madagascariensis. Itsscatteredhighland distributionhas givenrise toappreciablegeographic variation. Threeroughdivisions
are describedbutfor thepresent nosubspecies are recognized: EastZaïre, adjoining Uganda and (probably) Rwanda;
(probably) West Ethiopia, South Sudan, EastUganda and Kenyan highlands; and NortheastTanzaniato Malawi.The
Malawirecords arethe first forthatcountry.Of R. madagascariensis, manynew specimens havecome tolight, and its diag-
nosis is adapted accordingly. Itsassociation, asasubspecies, with R. lanosusby somerecent authorsisbasedonamisinter-
pretation ofits original description and other literature, and rejected as untenable.Its differences fromR. obliviosus are
described in detailfor the firsttime. Somepreviously unrecorded specimens ofR.obliviosusarereported. Itissuggestedthat
sexualdimorphismmay beatleastpartlyresponsible forthe“geographic” variationnotedbyKock(1978).
79
INTRODUCTION Roblivio.sus Kock, 1978; R. spinalatus Bergmans &
Hill, 1980); and the synonymization with Rouset-
For a genera] introduction to the series of which tusofthegenusBoneia Jentink, 1879(with asingle
this paper forms the fourth part, the reader is species, B. bidensJentink, 1879) by Bergmans et
referredto the first part (Bergmans, 1988), which al., 1988, necessitate arevision of the concept of
also containsasection Material and Methods. Rousettus as formulated by Andersen(1912: 16-
23). (An analysis of the pre-Andersen develop-
mentalhistory ofthe concept ofRousettuswill not
TAXONOMICSECTION be attempted here. Andersen (1912), in his syn-
onymies and historical sections on genusand
Rousettus Gray, 1821 species, provided all the references and sum-
maries of the important ones. But the original
Rousettus Gray, 1821: 299(type species: Pteropus egyptiacus t. publications of Rousettus and some of its syn-
Geoffroy-St. Hilaire, 1810); Palmer, 1898: 112; onyms contain no descriptions and the descrip-
Matschie, 1899: viii, 65 (in part: excluding Pterocyon
tions of some other synonyms are very briefand
Peters, 1862, Myonycteris Matschie, 1899, and Cynonycteris
brachycephala Bocage, 1889); Andersen, 1907 b: 505; equivocal, while most include other taxa. The
Miller, 1907: 54(inpart: notMyonycteris Matschie, 1899); meant analysis would therefore focus on biblio-
Andersen, 1912: 16 (in part: excluding Lissonycteris graphical facts more than anything else and it
Andersen, 1912); Leche, 1921:40; Kulzer, 1979;Corbet seems much wiser to accept Andersen's descrip-
etal., 1991:40.
tion of the genus as the authoritative one, in
BoneiaJentink, 1879: 117(type species: Boneiabidens Jentink,
which all earlier accounts have been sufficiently
1879); Miller, 1907: 61; Andersen, 1912:55; Leche,
1921:40;Corbetetal., 1991:41. considered.) The present account of African
Stenonycteris Andersen, 1912: 23 (type species: Rousettus Megachiroptera is not the appropriate place for
lanosus O. Thomas, 1906);Leche, 1921:41; Kingdon, arevision of the entire genusRousettus as the fun-
1974: 117, 124. dament for a new genus concept. In the mean
time,availableknowledge ofthe species involved,
The genus name Rousettus Gray, 1821 has long including the five extralimitalones presently rec-
gone unnoticed. Several subsequent synonyms ognized, allows for the following amendmentsof
were based either on the type species, R. egyptia- Andersen's diagnosis anddescriptive notes:
cus (E. Geoffroy-St. Hilaire, 1810) or on very Generalshape ofskull as inEidolon, but brain-
close relatives or other species, even by Gray case deflection variable(the alveolarline project-
himself, until Rousettus was revived by Palmer, in ed backward passing through upper part of
1898. Both Miller(1907) and Andersen (1907b, occipital condyle or upper margin of foramen
1912) gave synonymies. Miller included Myonyc- magnum or level with medianpart of occipital
teris Matschie, 1899, proposed as a subgenus of ridge), and the tympanic without bony auditory
Rousettus to accommodate Cynonycteris torquata meatus; rostrum rather long (length greater than
Dobson, 1878 and C. angolensis Bocage, 1898. lachrymal width) and, in the larger species,
Andersen (1912) considered Myonycteris an inde- broad; front of orbit vertically above middle or
pendent genus, which is followedhere, but sepa- posterior halfof M1; premaxillaries separated or
rated C. angolensis from it and proposed a new in contact, in some taxa co-ossified in a minority
subgenus ofRousettus for this species, \.e. Lissonyc- of specimens. Incisors 2/2-2/2 but I 1 often
teris. Several laterauthorshave argued thatLisso- deciduousin bidens;lower incisors(when unworn)
nycteris should standas agenus, whichviewis also bifid except, probably, in bidens; cheek-teeth
held by the present author, andwill be elaborat- normally 5/6 (P1 deciduous in leschenaultii semi-
edin the nextpartofthis series. nudus Gray, 1870 and amplexicaudatus brachyotis
This differenceof opinion with Andersen; the (Dobson, 1877)); P1 sub-equal in bulk to an
addition, after 1912, ofseveral new species to the upperincisor; Mj shorter than M2 andM3 com-
genus (R. madagascariensis Grandidier, 1929; bined. Second digit clawed; membrane insertion
80
variable: from sides ofback or(in spinalatus) from introduced a different class of characters to dis-
the spinal line; a short but distinct tail; forearm tinguish Stenonycteris from Rousettus: a different
length 65-107mm.Palatalridge formula4+ 3 + wing posture when roosting, different foreclaw
1 (exceptionally 3 + 4+ 1),4 + 4+ 1 (sometimes use and locomotionwhenclambering about, and
4 + 4/3 + 1 or 4 + 3/4 + 1; foran explanation shape, posture and movements ofthe ear. He
of this notationsee Eisentraut, 1960), or 4 + 3 + therewithsubscribed to the viewsofLawrence et
1/2(sometimes 4 + 3 + 2/3)(in bidens). al. (1963) who used, amongothers, behavioural
A revision of the entire genus should also and locomotory characters to separate
include a reassessment of proposed subgeneric Lissonycteris from Rousettus. Kingdon considered
divisions. Removing Lissonycteris from Rousettus Stenonycteris also as a genuson itsown.
leaves us with the subgenera Rousettus and Steno- I fully agree that patterns of behaviour and
nycteris Andersen, 1912and, as suggested recently locomotion should be given full considerationin
by Corbet et al. (1991), Boneia. Stenonycteris was phylogenetic reconstructions. But whereas the
distinguished mainly on the basis ofits relatively separation ofLissonycteris appears to be justified
strong brain-case deflection and excessively nar- on other grounds (to be dealt with under that
row cheek-teeth.Leche(1921) thought thatSteno- genus in the next part of this series), there is as
nycteriswas only slightly less different fromtypical yet no solid basis for subgeneric divisionswithin
Rousettusthanwere BoneiaandLissonycteris, andall Rousettus. Only Andersen (1912) has compared
three would deserve generic rank. Describing a all Rousettus species (then known) - a prerequisite
series of Rousettus madagascariensis, Bergmans for the appreciation of possible meaningful sub-
(1977a) argued that this species is intermediate groupings. The known differences between
between typical Rousettus and Stenonycteris in all Boneiaand Rousettuswere thought to be ofa grad-
morphological characters except the narrowness ual nature (Bergmans et al., 1988), and as far as
ofits cheek-teeth whichwould place it in the lat- Stenonycteris is concerned, the apparent distinc-
ter, if one would wish to maintainthat. If I tions from Rousettus egyptiacus as observed by
understand him well, Kock (1978a), describing Kingdon (1974) relate to characters which afie as
Rousettus obliviosus, observed that a strong brain- yet of unknown state in the majority of other
case deflection is not necessarily a primitive Rousettus species. (The account of the genus
characterand thusofuncertainvalue inphyloge- Lissonycteris in the next part of this series will
netic assessments such as the recognition ofsub- include an analysis ofmany morphological char-
generic divisions. Bergmans (1977a) considered acters in Rousettus species as well.) To a lesser
brain-case deflection as a neotenic character, extent, this also holds for the faculty of acoustic
weakened but persisting in adults of madagas- orientation.Among Megachiroptera, this has
cariensis (and most strongly so inthe smallest indi- been found only in Rousettus. It has so far been
viduals). Rousettus bidens, with a fal of 94.3-103.5, established in the species egyptiacus, amplexicauda-
a gsl of43.8-46.3and weights of 150-194one of tus(E. GeofTroy-St. Hilaire, 1810) and leschenaultii
the largest species ofthe genus(see Bergmans et (Desmarest, 1820) (as seminudus Gray, 1870)
al., 1988), hasa strongly deflected brain-case but (Mohres et al., 1956; Novick, 1958a). Notwith-
is otherwise quite different from Stenonycteris, standing Kock's (1972) assertion ofthe opposite,
which indicates that adult brain-case deflection Kingdon (1974) made it plausible that R. lanosus
may indeed be an adaptation developed inde- Thomas, 1906 also orients acoustically. Payne et
pendantly in various strains of Rousettus. On al. (1985) implied this for R. spinalatus when they
grounds not statedbut most probably thesemor- wrote that this species lives in dark caves, while
phological considerations Corbet etal. (1991) Bergmans et al. (1988) reported on the cave-
synonymized Stenonycteris with Rousettus. (They dwelling habits of R. bidens and R. celebensis. R.
retainedBoneia as a subgenus for reasons stated madagascariensis also roosts in caves (this paper).
by Corbet et al. in 1992 and to be discussed in Kulzer (1979) said that other cave inhabiting
the next part of this series.) Kingdon (1974) fruit bat genera use partly lighted caves only.
81
Novick (1958a) found that Eonycteris Dobson, Cynonycterisaegyptiacus:WoodJones, 1917.
1873, as one of those genera, orients visually, Rousettus egyptiacus; Mendez, 1937:63; Corbet et al., 1992:
67.
andmay findits way through thedark by memo-
Rousettus aegyptiacusoccidentalis Eisentraut, 1960a: 231
ry or by random noise and echoes. Gould(1988) (type locality: Mueli).
suggested that they might use the clapping Roussetus aegyptiacus; Brosset, 1966c: 131.
sounds oftheirwings. Inthe diagnoses in the fol- Rousettus aegyptiacusssp.;Feiler, 1984:75.
lowing spe-cies accounts those differential char- (Furtherreferencesunder thesubspecies.)
acters have been selected which when used in
combination allow for the discrimination of the
African forms here considered. Some extralimi- Diagnosis: A medium-sized short-furred fruit
tal species are very distinct and cannot possibly bat, on average thelargest species ofthe genusin
be taken for any of the African ones, but some Africa, total fal range 82.1-106.3 (minimum in
others possess morphological similarities to cer- Africa 85.7); brain-case only very moderately
tain African species. Rousettus leschenaultii resem- deflected, alveolar line straight, if projected
bles R. egyptiacus and R. obliviosus; R. amplexicauda- back-ward passing near or through upperpartof
tus is not always easy to distinguish from occipital condyle; widths of P3, P4, M1, P3, P4
leschenaultiiand may thus also be confused with and Mj more than halftheir lengths; wing from
these. However, an extensive treatment of back of first toe or interspace between first and
extralimital species would not be in place here. secondtoe; dorsal side of tibiapractically naked,
Specimens ofknown origin need not present with very short and thinly spread hears only.
identification problems, except, perhaps, speci- Measurement ranges and ratios for subspecies
mens from Pakistan, where egyptiacus 'meets' combined:
leschenaultii. This matter will be discussed in the
account ofR. e. arabicus Ander-son& de Winton, fal cTcr 85.7 - 106.3,
1902. 99 82.1 - 101.7;
gsl eft? 38.8 - 46.6,
99 38.0 - 45.8;
Rousettus egyptiacus (É. Geoffroy - St. rl era 35.6 - 39.1% ofgsl,
Hilaire, 1810) 99 35.0 - 38.8% ofgsl;
C^C1 era 18.3 - 22.3% ofgsl,
PteropusEgyptiacus E. GeofFroy-St. Hilaire, 1810: 96(type 99 18.8 - 21.6% ofgsl;
locality: the greatPyramid, LowerEgypt - i.e. at Giza); M2-M2 era 27.9 - 32.1% ofgsl,
the specific epithethas laterbeencorrectedandchanged 99 28.1 - 32.9% ofgsl.
into ;aegyptiacus by its author (see foot-note in Andersen,
1912:29), butthis correctionis consideredanunjustified
emendationbyCorbetetal. 1992.
, Distribution:Figs. 1 and 2.
PteropusLeachiiSmith, 1829:433 (type locality: Cape Town).
Eleutherura unicolor Gray, 1870: 117(type locality: Gaboon).
RousettusarabicusAnderson & de Winton, 1902: 86, 88, 90
(typelocality: Aden). Remarks
Rousettusaegyptiacus; Andersen, 1912: 29;G.M.Allen, 1939a:
62; Rode, 1941: 80; Ellermanet al., 1953: 45; Kulzer,
Andersen (1912) distinguished three forms with-
1956; Mohres et al., 1956:2; Griffinet al., 1958; Bar-
Yosefetal., 1966;Kock, 1969: 16;S. Erkert, 1970;Pirlot, in what is presently considered Rousettus egypti-
1970;Pirlot etal., 1970;Lane etal., 1971;Norberg, 1972; acus: R.egyptiacus, R. leachiiand R. arabicus. His R.
Brown, 1973; Kingdon, 1974: 128; Suthers etal., 1980; egyptiacus included the typical subspecies and
K.D. Jiirgens et al., 1981; H.G. Erkert, 1982; Gray's Eleutheruraunicolor. In his diagnoses the
Kleinschmidt etal., 1982; Happold, 1984: 272; Dobat et
sexes are not separated. According to his key,
al., 1985: 297; Hickey etal., 1987;S. P.Thomas, 1987:
egyptiacus and leachii differ from arabicus in the
90; Bergmans etal., 1988: 11;Bernard, 1988; Norberg,
1989:205. greater length oftheir tibiae(40-45.5 against 37-
Roussettusaegyptiacus; Langetal., 1917:480,481. 39.5) and their rounded (against attenuated)
82
Fig. 1.DistributionofRousettusegyptiacus (É. Geoffroy-St. Hilaire, 1810): Westof38° Eandnorthof20°N,the nominate
subspecies; eastof 38° E in the Arabian Peninsula and eastward: R. e. arabicus Anderson & de Winton, 1902 (question-
markedlocalitiesindicate observationsof Rousettussp.” and arein needofconfirmation); the single Ethiopian record on
thismap probablyrepresentsR. e.leachii(Smith, 1829).Blackdots:squares fromwhich materialhas beenidentified by the
author. Open circles:records fromliterature,museumregisters, andcorrespondence.
eartips; egyptiacus differs from leachii in having a are averaging markedly larger than in leachii.
larger skull (43.6-46.7 against 40.5-43.8), aheav- Andersendifferentiatedarabicusalso mainly from
ier rostrum, a palatal ridge pattern of4 + 4 + 1 leachii, which heused as astandard insteadofthe
(against 4 + 3 + 1; with some variationin both). typical species: the skull in arabicus is smaller than
From his species accounts it further appears that, in leachii (38.7-41.8 against 40.5-43.8), the ros-
dueto amore developed frontalregion inegyptia- trum shorter(13-13.6 against 13.8-15.2) and the
cus the interorbitalwidth is larger than the pos- teeth are smaller (the molars narrower) than in
torbital width while in leachii the latter is almost leachii; the palatal ridge pattern in arabicus is
always larger than the interorbital width; the mostlyas inegyptiacus butsometimesas in leachii.
temporal ridges unite into a sagittal crest at a The first reviser after Andersenwas Eisentraut
more distal point in egyptiacus thanin leachii, and (1960a). He followed Ellerman et al. (1953) in
often remain separate in the latter; the coronoid ranking leachiias subspecies ofegyptiacus and con-
process of the mandibulum is higher and the sidered arabicusalso as a race of that species. He
angular portion stronger and more projecting in furtherproposed a new subspecies, occidentalis, for
egyptiacus thanin leachii;and the teeth in egyptiacus specimens from West Africa (from Senegal to
83
Gabon). He noted thatAndersen(1912) had syn- ridge pattern of4 + 3 + 1 withparts of an extra
onymized unicolorwithtypical egyptiacus on wrong ridge behind the sixth. R. e. occidentalis averages
grounds but did not use the name unicolorfor the larger than typical egyptiacus, has a fal of 86.5-
West African subspecies. Like Andersen, he did 102, an average skull size intermediatebetween
not separate the sexes inhis analysis. He took the egyptiacus and leachii; a rostrum also intermediate
typical subspecies as standard and his diagnoses between those two subspecies; rarely co-ossified
deviated fromAndersen's on somepoints: egyptia- premaxillaries; an interorbitalwidthmostiy as in
chasa fal ofu88.2-96.5; is largse-headed, with a leachii; smaller than postorbital widthin 14, equal
wide andhigh rostrum;co-ossified premaxillaries to it in 11, and larger thanit in 4 specimens; a
in aminority ofoldspecimens only (6 outof 22); strong sagittal crest in 5, a weak crest in 5, and
interorbital width usually larger than postorbital nocrest atall in 21 out of31 specimens; an angle
width (in 15 outof 18 specimens); adistinct sagit- between mandibular ramus and coronoid
tal crest in most old specimens (in 17 out of 22); process which is intermediatebe-tween egyptiacus
a relatively small angle between mandibular and leachii (129°-140°; mean 33.5°); dentition
ramus and coronoidprocess (121°-132°); strong- weaker than in egyptiacus, with narrower molars,
ly developed dentition, with broad molars; a more as in leachii; apalatal ridge pattern of4 + 3
palatal ridge pattern of4 + 3 + 1 with in most + 1 in 44 out of 61 specimens (with the fourth
specimens atleast the beginning of an additional divided in 5 specimens, which could thus be
ridge behind the sixth: 4 + 3/4 + 1 (or, when described as 3 + 4 + 1 - but the division is nar-
further developed, 4 + 4/3 + 1 and when com- row and difficult to interpret as indicative of a
plete 4 + 4 + 1). R. leachii has a fal of87.8-96.2, change, as the presentauthor hasobserved), 4 +
averaging slighdy smaller in body measurements 3/4 + 1 in 11,4 + 4/3+ 1 in 5, and4 + 4+1
than egyptiacus; a smaller head than egyptiacus, in 1.
with amore delicately built skull and aslendeier Eisentraut (1960a) observed that some speci-
rostrum (Eisentraut didnot state ifpremaxillaries mens from Gabon averaged somewhat larger
are co-ossified or not); a relatively small interor- than the types ofoccidentalis from Cameroun,
bital width (smaller thanpostorbital width in 5 while specimens from more western regions
out of 7 specimens); no sagittal crest, and sepa- (Senegal to Ivory Coast) were somewhat smaller.
rated temporal ridges in most specimens; a larger Koopman (1966) considered occidentalis a syno-
angle between mandibularramus and coronoid nym of unicolor, "the oldest name for the west
process (132°-142°) than egyptiacus; weaker denti- African subspecies", without referring to Eisen-
tion thanin egyptiacus, with narrower molars; a traut's observations. Bergmans (1979) studied
palatal ridge pattern of 4 + 3 + 1 in one speci- specimens from Congo which confirmed
men and 4 + 4 + 1 in another. R. arabicus aver- Eisentraut's findings; as the varation seemed
ages smaller in all measurements than the other modest and clinal heagreed with Koopman that
subspecies although there is considerable overlap all populations from Senegal to Cameroun and
in size. Eisentraut could study only two speci- from there to Angola (i.e. west of 16E) should be
mens and didnot give afullanalysis; no remarks assigned to thesame subspecies, unicolor.
were madeon premaxillaries, inter- and postor- In the following accounts I have maintained
bital widths, and sagittal crest. Mentionedare: A the four subspecies. Where necessary and possi-
relatively small angle betweenmandibularramus ble, theirdiagnoses are completed andamended.
and coronoidprocess (128°); strongly developed The differences between them are slight but
dentitionwithbroad and, in somecases, relative- obvious and indicate independent lines of devel-
ly long molars (Eisentraut pointed out here that opment inall four,whiletheirknowndistribution
Andersenerroneously described the dentitionof areas are disjunct.
arabicus as weaker and its molars as narrower (Note:Juste etal., 1993 have published two newsubspecies
than in leachii; in some cases the molars may be ofRousettus egyptiacus, from the islandsofPrincipe and Sao
Tomerespectively, intheZoological JournaloftheLinnean
slightly narrower than in egyptiacus); a palatal
Society.)
84
Rousettus egyptiacus egyptiacus assessed, 10-VII-1963 (SMF 22643/61);2 Cftf. 1 imm.<?, 3
(E. Geofiroy - St. Hilaire, 1810) 99, 1 imm. 9, ale., skulls ofadults, 7-1-1984, A. Walen
(ZMA 22.205/11). Cairo: 1 imm. <?, ale., no skull, 1888,
PteropusEgyptiacus E. Geofiroy- St. Hilaire, 1810: 96(type Stuhlmann(ZIZM 22062); 1 9, ale., 10-VII-1931(SMF
12035); 6 specimens, ale., E. Kulzer (SMF25778/79,
locality: thegreatPyramid, LowerEgypt - i.e.atGiza).
Rousettus aegyptiacus; Bonhote, 1909: 788; Andersen, 1912: 25861/64: captive bred); 3 specimens (BMNH 3.12.8.1, -
.3/.4); ale. material(BMNH); 1 specimen (USNM 312121).
29; G.M. Allen, 1939a: 62(in part), 1939b:234, 282;
Cairo region: 1 Cf, ale., 6-VI-1880, Mook (NMW 15203).
Ellerman et al., 1951: 92 (in part); Sanborn etal., 1955;
Damietta:ale. material(BMNH). El Mansuriya: 1 9, 1
Kulzer, 1958: 375, 1960: 240, 1961: 219; Kaisila, 1966;
imm. 9, ale., 28-V-1951, H. Hoogstraal (FMNH
Lehmann, 1966: 258; Storch, 1968;Atallah, 1970;
79163/164). El Walidiya: 3 cfcf, 2-IV-1957, H. Hoog-
Harrison, 1972:626; Madkour, 1976; Atallah, 1977:
straal/M. N.Kaiser (FMNH 87766/68). Gezira Island: 1
286; Kumerloeve, 1976a: 84, 1976b: 170; Kock etal.,
specimen (USNM 282381).(El) Giza: 1 d\ 1 9, ale., 27-V-
1979: 68; Wassifet al., 1984;Makin, 1989: 405, 1990;
1953,H. Hoogstraal (FMNH 85521/22); 4 specimens
Opstaele, 1990: 16; Churcher, 1991.
(BMNH 9.7.1.1/.4); ale. material (BMNH). Nr Luxor
Rousettus egyptiacus; Flower, 1932: 376; Setzer, 1952: 346;
(Kurna and Grand Hotel ruins): 8 Cfcf. 1 9, 1 specimen,
Lewisetal., 1962:474; Kuhn, 1968.
ale., 21/26-III-1938, H. Nelson(FMNH47772/81).
Rousettus aegyptiacus;; Dor, 1947: 50.
Mahallet el Kebir: ale. material, J. Anderson (BMNH).
Rousettus aegyptiacus aegyptiacus;Ellermanet al., 1953: 46 (in
Medinet el Fayum: 1 9, ale., skull, 1893, R. H. Brown
part); Eisentraut, 1960a: 222; Maser, 1966; Hayman et
al., 1971: 11; Gaisler et al., 1972: 4; Koopman, 1975: (ZMA 16.668). "Egypt": 1 Cf, 1829,via museum at
361; Eisentraut, 1976: 74; Spitzenberger, 1979: 440; Frankfurt (SMNS 3608); 1 Cf, 1851, vonMuller (SMNS
Qumsiyeh, 1980: 37, 1985: 13;Qumsiyeh etal., 1986: 426); asmallseries, 1-VI-1852,Th. vonHeuglin (NMW); 3
specimens,ale.,Letourneau (MNHNCG 1880-2039/40);6
140;Korineetal., 1990;Harrison etal., 1991: 24.
specimens (BMNH 39a/f); 1 9> ale., 3 skulls, 4 mounted
Rousettusaegyptiacusaegyptiacus; Hoogstraal, 1962: 145
specimens (RMNH); 2skulls, 2specimens (1 in ale.), 3 skins
Rousettus (Rousettus) aegyptiacus aegyptiacus; Anciaux deFa-
veaux, 1978a: 460 (inpart: the specimens from Egypt); (SMF892/93, 1314, 12232, 12444,-79, -83): 19, skin, von
Sack(ZMB348);3specimens,ale.(ZMB 10252/54).
Kock, 1978a: 206.
(Aswan, Burg el Arab, DeltaBarrage Gardens, Dumyat, El
Aiyat,ElFaiyum, ElKarnak, El Minya, ElTell el Kebir, El
Zamalik, Fuah, Ismailiya, Mataria, Matruh, Nigm, Port
Materialexamined Said,0,'asr,Qina,Thebe, WadielNatrun.)
ISRAEL.Herzliyya Cave,MountCarmel: 1 imm.9, 9-VI-
CYPRUS. Apsiou: 1 imm. C?, 1 9,ale., skulls, 30-111-1973, 1968,E.Shor (NMW 13535).Jaffa: 1 imm.9, 2 CfcT, ale.,
F. Spitzenberger (ZMA22.124/25). Konktia: 4 specimens, skulls, Aharoni (ZMB 53866/68).Jerusalem: I imm. C?,
ale., 21-IV-1913, G. Cecconi(ZMB). Somekm S ofLachi: ale.,5-II-1950,G. Haas(FMNH75788); 19, ale., skull, 17-
6 cfcf, 1 9, 2 imm. 99, 15/16-V-1985,J. van Wingerde X-1958 (SMF 20526). Mount Carmel: 1 specimen, ale.,
(ZMA 22.902/10). Larnaka: 1 specimen, G. Cecconi skull (SMF 18988).Ofer Cave: 16 adult, 3 imm., skulls,
(USNM 123303). Nicosia: 3 specimens (BMNH 3.12.4.3/ mostly incomplete, sex unknown, 22-X-1988, S.G. Sowler
.4, -.6). "Cyprus": 4 specimens (BMNH 79.10.16.5/.6, (fumigation victims; ZMA23.911/29).Tabgha: 1 skin, 4-1-
99.7.2.1/.2); 1 9, 1 imm. Cf, 2 specimens, ale., Rolle(ZMB 1912,E. Schmitz (ZMB). ? Tel Aviv: 2 99, 1951, E.
10248/51). Sochurek (NMW 8379, 17905)."Palestine": 1 C? (ZMB
(Akrotiri, Ayia Napa,nrAyioaEpiktitos, Bellapais,Episkopi 53869); 1 Cf, 1 imm.Cf, ale., via M.Dor (MNHNCG 1948-
Bay, Famagusta, between Klepini and Pentadactylos, 500, 1951-1).
Ktema, Lachi, Paralimni,Polis, Prastiou, betweenPyla and (Between Bertovia and Tel-Chai: in owl pellets, Dan,
Troulli, Trozina,YermasoyiaReservoir.) Hayonim cave: Pleistocene remains, Me'arath Hateamim
EGYPT. Abu Simbel(temple): 1 specimen,21-1-1963,L.D. nrHartuv, Rehobot,WadiKurn atAcre.)
Brongersma (RMNH 17668). Nr Assiut: ale. material JORDAN.Jericho: 1 skin (ZMB).
(BMNH). Cairo, Abbassia: 1 C?, 1975, K. Wassif(ZMA (Al-Mahhattah, Amman,ElHamma.)
22.162). Cairo, Citadel: 4Cfd1, 2 imm.Cfcf, 399, 1 imm.9, LEBANON, Antilyas: 1 specimen (BMNH 22.7.6.10); 1
1 imm., ale., 20-VIII/2-X-1952, H. Hoogstraal (FMNH imm. Cf, 19-III-1960,J. E.Stencel (FMNH99555). Beirut
80748/58). Cairo, Maade: ale. material(BMNH). Cairo, River Cave: 1 specimen (BMNH 61.392). Maam el Tien
Mohammed Ali Mosque: 1 Cf, 1 9, 3-V-1951, H. Hoog- Cave: 1 9, 2 imm. 99, 1 imm. Cf, ale., 1955,C.A. Reed
straal (FMNH 78591/92).Cairo, Sultan Hassan Mosque: 4 (FMNH 84501, -12/13, -16). Mount Lebanon: 1 specimen
dVJ,2imm.Cftf, 299, 1imm., 22-VI-1959,H. Hoogstraal (BMNH94.5.7.2).
(FMNH 90461/62, 12189,31390/95); 1 Cf, 1 9, H. (Amchite, Beirut, Beit Meri, Hazmieh, Jahmour,Junieh,
Hoogstraal (FMNH 89707/08); 5 cTcf, 14 99, ages not LebanonCave, Mogharet Saleh, Ras Beirut, Roman
85
aquaduct 2km E ofHazmieh, Saida,Tripoli.) W 99 112 (n = 1);
SYRIA.ZerkaRiver: I9,ale.,skull, 1886 (SMF 11913). rl eft? 36.8 - 37.9 ofgsl (n = 7),
TURKEY. "Turkey": subfossilremains (BMNH67.616).
(Alanya, Antakya, Antalya, Bedirgeregion, DermustluKoy, 99 35.0 - 37.6 ofgsl (n =6);
Giilnar, Harbiye, ? Iskenderun.) C'-C1 era 20.4 - 21.3 ofgsl (n =5),
99 20.6 - 21.5%ofgsl (n =4),
Diagnosis: Generally as for the species, but the ? 19.2%ofgsl (n = 1);
least specialized of the subspecies, with on aver- M2-M2 c?c? 28.2 - 30.2%ofgsl (n = 5),
age a large skull, broadrostrum and interorbital 99 29.3 - 30.8%ofgsl(n = 4),
region, a sagittal crest in most old specimens, a ? 29.8% ofgsl (n = 1).
small angle between mandibular ramus and
coronoidprocess, andstrong cheek-teeth. Setzer (1952) foundsmaller skulls (gsl male41.8,
The following measurement ranges and ratios 9 39.2) in specimens from Gezira Island, Cairo.
are from allover the subspecies' range(the speci- Spitzenberger (1979), in her excellentaccount of
mens of unknown sex are the series from Ofer a large series from Cyprus, found smaller mini-
Cave, Israel): mumdimensions in maleskulls (e.g. gsl 41.6, iow
7.9, C^-M2 15.6) and larger size rangesin99 (gsl
fal cfc? 87.1 - 101.4 (n = 37), 40.4-44.9; iow 7.5-9.0, C^M2 15.4- 16.9); she
99 86.0 - 100.3 (n =22); published a weight range for 66 males of 135-
tibia cfcf 41.4 - 45.6 (n = 8), 175. Gaisler et al. (1972) gave a weight range of
99 41.1 - 42.6 (n = 6); 125-170.5 in 8 cfcf, of89.5-151 in 3 99 and
ear cfcf 18.3 - 21.7 (n = 6), 101.5-162 in 3 pregnant 99 - all from Egypt.
99 20.0 - 21.4 (n = 4); Especially 9 specimens from Cyprus appear to
gsl cfcf 43.4 - 46.2 (n=12), average somewhat larger than those from Egypt
99 41.2 - 44.2 (n= 9), (fal range, including literature data, 89.0-100.3
? 42.0 - 42.7 (n = 2); against 84-97.0); from other areas the numbers
rl cfcf 16.3 - 17.0 (n = 9), ofavailable adult specimens are insufficient for
99 14.8 - 16.6 (n = 6); an assessment ofgeographical variation.
iow cfcf 8.5 - 9.9 (n = 9),
99 8.0 - 8.6 (n= 6), Distribution: Fig. 1.
? 7.5 - 8.7 (n = 16); Relatedspecies: In large parts of its distribution
pow cfcf 7.7 - 8.6 (n = 9), area Rousettus egyptiacus is the only representative
99 7.1 - 8.4 (n = 6), of its genus and not easily confused with other
? 7.2 - 8.4 (n = 16); species. It has no white facial markings or ear
zw cfcf 26.2 - 29.1 (n = 10), tufts, which distinguishes it from all epo-
99 24.9 - 27.0 (n = 6), mophorines. Its possession of an externally dis-
? 25.4 - 27.2 (n = 13); tinct tail is shared by three other genera only:
C'-M2 cfcf 17.0 - 18.2 (n = 9), Eidolon Rafinesque, 1815with larger dimensions
99 16.1 - 16.8 (n = 4), (fal 109.8-133.2, gsl 52.2-58.6), a partly straw-
? 15.9 (n = 1); coloured fur (against dull or dark but never
C,-M3 cfcf 17.2 - 19.7 (n = 7), bright in Rousettus egypticacus), separated pre-
99 17.2 - 18.2 (n = 4); maxillaries, and a bony auditory meatus;
M1 length cfcf 3.3 - 3.45(n = 7), Myonycteris Matschie, 1899with smaller species
99 3.1 - 3.45(n = 6), (fal 54.9-75.1, gsl 30.1-39.2), dense fur, a less
? 3.2 - 3.5 (n = 3); deflected brain-case, on average a relatively
M1 width cfcf 2.1 - 2.3 (n = 7), shorter rostrum, wings from second toe, a dorsal-
99 2.0 - 2.15 (n= 6), ly partly or wholly furred tibia, and a band of
? 2.1 - 2.3 (n = 3); thick hairs on necksides and foreneck in males;
86
Lissonycteris Andersen, 1912, wilh smaller body The owner did not want to sell it and two years
measurements (fal 67.8-88.0, gsl 36.6-46.4), long later both bat and shop had disappeared. Dr
and dense fur, a less deflected brain-case, squar- EJ.H. van Bree also observeda stuffed specimen
ish cheek-teeth, wings from second toe, a dorsally of the same species in a shop in Alanya in 1979
furred tibia, and a band of thick hairs on neck (pers. comm., VII-1979) but unfortunately the
sidesand foreneck in males. shop was closed. Kinzelbach (1986) mentioned
In parts ofEast Africa, Rousettus lanosus is sym- an observation of a "gigantic bat, obviously a
patic. This species has long fur, dorsally furred fruit bat" near Gulnar, in 1986. Dr R. Geldiay,
tibiae, somewhat smalleraverage size (fal 85.3- in aletter to Dr van Bree (27-VIII-1979), wrote
95.0, gsl 39.4-44.8), a strongly deflected brain- that the cultivationof fruits like apples, tanger-
case, narrow cheek-teeth and wings usually in- ines, oranges andbananas had become common
serted at thesecond toe. south ofthe Taurus Mountains, from Hatay
In Pakistan R. egyptiacus arabicus occurs together westwardto Finike (at 30°08'E). This has proba-
with R. leschenaultii. They have both been record- bly paved theroadforRousettus from the Antakya
ed in Karachi but are possibly largely allopatric. region, although immigration from Cyprus
R. leschenaultiiin thisregion is smaller, on average, should alsobe considered a possible source ofthe
andhas relatively shorter wings. The differences species near Alanya. (Dr V Calandra told the
between the two species have not yet been suffi- present author about stories referring to gigantic
ciently analysed, through lack of material, and bats which she heard in southern Sicily (pers.
thereader is referred to the section onR. egyptia- comm., 20-VIII-1987). Hufnagl (1972) wrote
cus arabicus, where underRemarks and in tables4 that "fruit bats have been found on some
and5available dataare given and discussed. Mediterraneanislands and one even in Mostar,
Yugoslavia." He gavenofurther details.)
Fitzinger (1869: 81) recorded that [Th.]
Remarks Kotschy discovered the species in Syria. Theori-
gin of his material (neither in the NMW, nor
T a x o no m y: There is a slight but distinct sex- located elsewhere yet) or observations is most
ual size dimorphism, with cfcf averaging larger probably Beirut, or somewhere near it, in pre-
than 99- Measurement ranges should therefore sent-day Lebanon(DrF. Spitzenberger, inlit., 17-
include either male or 99 values. The fal ranges X-1991). Seabra (1898a) recorded a specimen
(87.1-101.4 in C?Cf, 86.0-100.3 in 99) show an from Syria in the MLZA collection. Both
extension when compared with the range given Matschie (1899) and Andersen(1912) referred to
by Eisentraut (1960a). The dominantpalatal the latter record (and Andersen also to the for-
ridge pattern is 4+ 4+ 1 (5 outof 14specimens) mer), and Kock's contention that they did not
with its derivates 4 + 3/4 + 1 (5 specimens) and refer to material evidence is therefore incorrect
4 + 4/3 +1 (1 specimen); other patterns are 4 + (Kock, 1969). Kock himself published a record
3 + 1 (2 specimens) and 4 + 3/5 + 1 (1 speci- from Zerka River, Syria, based on specimen
men). Inthe latter specimen there are additional, SMF 11913 collected there in 1886 by a Dr
incomplete 6th and 8th ridges. As for the other Schumacher. Later, Kock etal. (1979) discovered
characters of this subspecies, Eisentraut's diagno- that this specimen is not from Syria but from
sis (1960a) isaptandsufficient. Jordan. Jordan has a Zerqua or Zarqa River
Distribution and geographical (Harrison et al., 1991 and Times Atlas, respec-
variation: After its discovery in theAntakya tively) and the species' occurrence there is per-
region in southern Turkey in the 1950s, R. e. fectly likely. Theproblem withthe Syrian records
egyptiacus has been recorded from two other is theirage. The name "Syria" covered a differ-
Turkish localities. Kinzelbach (1986) saw a ent region thenfrom thatwich it doesnow(com-
stuffed specimen in a shop in Alanya in 1975, pare Andersonet al., 1902: 87.) Nevertheless,
which had reportedly been shot near that town. Rousettus may still be discovered in the Mediter-
87
ranean section ofpresent-day Syria (but proba- aging in wild trees. In Lebanon, Lewis et al.
bly not at the Ouadi Zergane or Zarqua Suyu (1962) observed a preference for the coastal
River in that country, at 40° 22' E, 36° 55' N - plains and the Lebanon mountains, which they
my initialidentification ofZerkaRiver). thought to be almost certainly correlated with
According to Kinzelbach (1986) the species the availability of food. A largely similar pattern
would now also occur in Sinai, following thecul- appears to be found in the wholeof the East-
tivation ofFicusspecies. Mediterraneancoastal region, and may develop
Nader (1975) identified 17 specimens from in southern Turkey as well. In Egypt, the sub-
Tayif, Saudi Arabia, as R. e. egyptiacus. However, species is restricted to the Nile valley and delta
while the falranges inthis series (88-95 in6C?Cf, and some few oases. Several authors mention its
85-93 in 6 99) overlap with the lower values in consumption of Ficus species, mostly figs but in
the typical subspecies they perfectly fitthe ranges one instance also leaves (Anderson et al., 1902;
known for arabicus, and their skull dimensions(gsl Flower, 1932; Lewis et al., 1962; Atallah, 1977;
38.8-42.1 in 6 cfcT and 38.9-40.5 in 5 99) Kinzelbach, 1986), and Spitzenberger (1979)
demonstrate that they shouldbe referred to that recorded shoots of Morus alba and fruits of
subspecies. Ceratonia siliqua as part of the diet. As these tree
In Egypt, the typical subspecies appears to generanaturally occur in the African Mediterra-
reach its southern limit in Africa. Kulzer (1958) nean and/or West Asian regions, these observa-
observed the species atAswan and Maser (1966) tions probably relate to the subspecies' custom-
identified Aswan specimens as R. e. egyptiacus. ary natural food. A reconstruction of its natural,
Subsequently, Aswan has been considered its i.e. prehistorical distribution pattern should cer-
southernmost locality (Kock, 1969; Koopman, tainly start with one of the contemporaneous
1975). Aspecimen collectedon 21-1-1963 atAbu forestsandtheir composition.
Simbel (RMNH 17668) proves that the species
occurs, or has occurred, at least 200 km more to
the south. Old references to the species' occur- Rousettus egyptiacus leachii(Smith, 1829)
rence in "Nubia" (quoted by Andersen, 1912;
Kock, 1969), which were never supported by PteropusLeachiiSmith, 1829:433(typelocality: Cape)
? Rousettus aegyptiacus; Senna, 1905: 256;Andersen, 1912:30
material evidence, are becoming more plausible
(inpart: thespecimen fromErythrea).
with this record. Records from more southern Rousettusleachi; Andersen, 1912:25; Lonnberg, 1917: 47;
Sudaneselocalities are treatedunderR.e. leachii. Fitzsimons, 1919: 91; Loveridge, 1923: 692; Flower,
R. e. egyptiacus appears to be very flexible in its 1931: 160; G.M. Allen et al., 193: 45; Friant, 1951;
two mainecological requirements: caves for day- Zuckerman, 1953: 836; Eisentraut, 1958: 18; Kulzer,
1958: 377;Fain, 1959:4;Theodor, 1968: 321.
roosting and food trees. It has most probably
Rousettuslanosuskempi(not ofThomas, 1909);Granvik, 1924
occurred in (parts of) its present distribution area 9(atleastinpart:see text).
at least from Pleistocene times onwards, as a Rousettuscollaris; Cowles, 1936: 122.
Pleistocene record from Hayonim Cave, Israel Rousettusleachii;; G.M.Allen, 1939a:62; Moreauetal., 1940:
(Bar-Yosef et al., 1966), subfossil remains from 118; A. Roberts, 1951: 55; Lawrence etal., 1953: 17;
Turkey (BMNH) and the c. 4000 years old wall Kulzer, 1959: 15; Hoogstraal, 1960: 359; Lombard,
1962;Didier, 1965:341.
paintings at Beni Hasan, Egypt (G. M. Allen,
Rousettus (Rousettus) leachi; Schouten, 1944: 102;Leleup,
1939b, fig 2) suggest. Thespecies has been quick 1956: 75.
to exploit man-made cave-like habitats such as Rousettusaegyptiacus leachi;EUerman etal., 1953:46;Harrison,
chambers in pyramids and temples, mosques, 1959: 222; Ansell, 1960b: 8; Eisentraut, 1960a: 230;
and buildings of lesser stature. And it readily Kulzer, 1962b: 116;J. D.Jurgens, 1963;Hayman etal.,
exploits a variety of cultivated exotic fruits. 1966 (inpart: notthespecimens from Tamba-tambaand
Thysville); Koopman, 1966: 155;Ansell, 1967: 2, 28;
Almost all reports on the typical subspecies refer
Verschuren, 1967; Ansell, 1969: 5; Niort, 1970: 256;
to these two faculties and hardly ever to its Hayman etal., 1971: 11;Anciaux deFaveaux, 1972: 82;
occurrence in particular natural caves or its for- Dulic etal., 1973:232; Koopman, 1975:360;Anciaux de
88