(NATURAL HISTORY)
,-;c?
The baculum in he Vespe ilioninae (Chi op e a:
Vespe ilionidae) wi h a sys ema ic e iew, a synopsis
o Pipis ellus and Ep esicus, and he desc ip ions o a
new genus and subgenus
J. E. Hill
Depa men o Zoology, B i ish Museum (Na u al His o y), C omwell Road, London SW7 5BD
D. L. Ha ison
Ha ison Zoological Museum, Bowe wood House, S Bo olph's Road, Se enoaks, Ken
TN133AL
Con en s
Synopsis
In oduc ion
Func ional and sys ema ic signi icance o he baculum
Na u e and scope o his s udy
Ma e ials and me hods
Au ho ship and esponsibili y
The baculum o Pipis ellus
The baculum o Ep esicus
The baculum in o he Vespe ilioninae
Sys ema ic conside a ions .....
Genus Pipis ellus Kaup, 1829
Genus Ep esicus Ra i nesque, 1 820
The s a us o he 'Nyc iceini'.
The classi ica ion o he Vespe ilioninae
Zoogeog aphical conside a ions
Conclusions
Addendum
Re e ences
Appendix 1 . Specimens examined
Tables 1-3
Figu es 1-22
Synopsis
Cu en classi ica ion o he Vespe ilioninae es s chie ly on a sui e o mainly adap i e cha ac e s, among
which acial sho ening h oughou he sub amily wi h consequen changes in he s uc u e, size, ela i e
posi ion and numbe o he incisi e and p emola ee h ea u es p ominen ly. Such cha ac e s may no
necessa ily e lec ela ionships o phyle ic di e si y, and some imes do no se e p ope ly o dis inguish he
gene a ha hey pu po o de ine, as in he dis inc ion o Pipis ellus and Ep esicus, whe e gene ic bounda ies
emain unclea . The sea ch o possibly less s ongly adap i e ea u es sugges ed he possibili y ha he
mo phology o he os penis o baculum migh p o e aluable in he s udy o he sys ema ics o hese gene a
and pe haps in he sub amily as a whole.
This pape e iews ea lie s udies o he baculum in he Chi op e a and hei ele ance o sys ema ics in he
O de , wi h an examina ion o i s g oss mo phology h oughou he Vespe ilioninae, especial a en ion being
gi en o species cu en ly alloca ed ei he o Pipis ellus o o Ep esicus. A synop ic e iew o he species
Bull. B . Mus. na . His . (Zool.) 52 (7): 225-305 Issued 30 July 1987
225
226 J. E. HILL & D. L. HARRISON
con en o hese gene a is p esen ed, wi h he ecogni ion and de ini ion o subgene a and included species
g oups: h ee such (pumilus, capensis and enuipinnis) cu en ly e e ed o Ep esicus on den al g ounds seem
ins ead mo e closely ela ed o Pipis ellus o which hey a e he e ans e ed. One subgenus o Pipis ellus is
desc ibed as new (p. 250).
The Vespe ilioninae as a whole display a wide ange o bacula a ia ion, which alls in o wo majo and
se e al mino g oups. This has sugges ed a e ision o he cu en classi ica ion o he sub amily, combining
bacula ea u es wi h hose con en ionally in use. Bacula mo phology p o ides a clea indica ion ha
he 'Nyc iceini' (o 'Nyc iceiini') is an a i icial g ouping and ha he genus Nyc iceius as p esen ly unde -
s ood is composi e. Cu en ly i is held o include wo species, he No h Ame ican hume alis and he A ican
schlie enii: hese a e he e hough o be su icien ly cha ac e ised o jus i y gene ic sepa a ion and a new
gene ic name is p oposed o schlie enii (p. 254).
A sugges ed classi ica ion o he sub amily is p esen ed, wi h a abula ed e iew o ea lie classi ica ions;
possible ela ionships be ween he cons i uen gene a a e discussed and he zoogeog aphy o he bacula ypes
wi hin he sub amily is examined.
In oduc ion
A penial bone is known o occu among mammalian O de s in he Insec i o a, Chi op e a,
P ima es, Roden ia and Ca ni o a. Va iously called he os penis, os p iapi o os glandis, i was i s
named he baculum by Thomas (191 5a), he co esponding s uc u e in he emale, he os cli o idis,
being la e called he baubellum by Sho idge (1934: 327, oo no e). The ea u es o he baculum
ha e been used ex ensi ely in a emp s o de e mine phyle ic ela ionships a a ious sys ema ic
le els (Pa e son & Thaele , 1982). Thomas (loc. ci .), o example, sugges ed ha he baculum
migh p o ide e idence aluable in he sub amilial classi ica ion o he Sciu idae and indeed
poin ed ou ha in his connec ion he e we e no bacula ea u es o suppo he associa ion o he
dwa squi els in a sepa a e sub amily, he Nannosciu inae. Mo e commonly, bacula ea u es
ha e been used o indica e o de e mine ela ionships wi hin gene a in he Sciu idae, among New
Wo ld oden s, and in he Mus elidae. Such cha ac e is ics ha e been employed in species desc ip-
ions, especially whe e bacula a ia ion is p onounced, and also o age de e mina ion. Nume ous
examples o hese uses o he baculum a e summa ised by Pa e son & Thaele (loc. ci .) while Bu
(1960) ga e an accoun o he ea lie o such s udies. Simila ea ly accoun s o he baculum in he
Chi op e a a e e iewed by Hamil on (1949).
The p esence o a baculum in some a leas o he Chi op e a has been long es ablished.
Dauben on (1760) desc ibed and illus a ed (in pa ) he baculum o Nyc alus noc ula and
Blain ille (1840) simila ly s udied he baculum o Rhinolophus e mmequinum, R. hipposide os,
Vespe ilio mu inus and again o Nyc alus noc ula, he la e au ho p o iding pe haps he i s
accu a e and qui e de ailed d awings o his s uc u e. La e wo ke s such as E colani (1868),
Robin (1881), Gilbe ( 1 892), Rau he ( 1 903), Ge ha d ( 1 905) and Chaine ( 1 926) p o ided u he
de ails o penial and bacula mo phology in he Chi op e a, Chaine in pa icula discussing and
illus a ing he baculum in se e al species and o some ex en summa ising ea lie wo k in he ield.
Howe e , none a emp ed o use he s uc u e o he baculum o sys ema ic pu poses.
The i s use o he baculum in chi op e an sys ema ics appea s o be by Thomas ( 9 5b) who
employed bacula cha ac e is ics in de ining he species o Nyc ophilus. This wo ke clea ly o e-
saw he alue o bacula ea u es in he de ini ion o some a leas o he species o ba s, beginning
om ha ime a collec ion o espe ilionid bacula a he B i ish Museum (Na u al His o y)
al hough subsequen ly making li le use o he ma e ial ha was accumula ing, excep in 1928
employing bacula cha ac e s o sepa a e Indo-Chinese species o Pipis ellus (Thomas, 19280, K).
Since hen he baculum has been u ilised in a a ie y o axonomic s udies o ba s, o example by
K u zsch (1959, 1962) and Lanza (1969) o examine i s alue in indica ing ela ionships in he
Megachi op e a, by Topal (19700) in de e mining he a ini ies o la, o by Helle & Volle h (1984)
as an indica o o ela ionship among he species o Pipis ellus and Ep esicus. The baculum o
Pleco us was ound aluable by Lanza (1960) in disc imina ing be ween P. au i us and P. wa di
( = P. aus iacus): he subsequen use o he baculum in dis inguishing hese species is summa ised
by Co be (1964). Genoways & Jones (1969) ound ha bacula ea u es dis inguished closely
VESPERTILIONINE SYSTEMATICS 227
ela ed species o No h Ame ican Myo is, LaVal (1973#) employing bacula cha ac e s o he
same pu pose among he Neo opical species o his genus.
The emphasis placed on bacula cha ac e s in chi op e an sys ema ics is pe haps bes illus a ed
by he numbe o s udies de o ed chie ly o bacula s uc u e, o en on a egional o aunal basis, as
o ins ance he wo k by Hamil on (1949) and K u zsch & Vaughan (1955) on No h Ame ican
species, by B own e al. (1971) on Neo opical ba s, by Topal (1958) on cen al Eu opean species,
by Bha naga (1967), Ag awal & Sinha (1973), Sinha (1976) and Khaju ia (1979, 1980, 1982) on
Indian ba s, o by Wassi & Madkou (1972) and Wassi , Madkou & Soliman (1984) on Egyp ian
ba s. Bacula a e some imes s udied in disc e e axonomic g oupings, as o example hose o New
Wo ld molossids by B own (1967) o o Malaysian Hipposide os by Zubaid & Da ison (in p ess).
Thus among he Chi op e a he baculum has been employed as a sou ce o axonomic ea u es a
se e al sys ema ic le els, bu p ima ily o indica e deg ees o ela ionship o o sepa a ion a he
speci ic and some imes he gene ic g ades, o especially o dis inguishing closely ela ed, o en
sympa ic species whose con en ional mo phological cha ac e s a e o he wise e y simila , as in
Myo is and Pleco us.
Func ional and sys ema ic signi icance o he baculum
Con lic ing hypo heses o bacula a ia ion we e e iewed in de ail by Pa e son & Thaele (1982).
These au ho s p oposed ha among oden s a leas he p obabili y was ha he baculum has a
p ecise ep oduc i e pu pose and unc ions p ima ily as a de ice con ibu ing o species isola ion.
Bacula di e ences among closely ela ed axa migh well hen ake an exagge a ed o m. As such,
he baculum would be he e o e a poo basis o sup a-speci ic classi ica ion, bu an excellen
s uc u e o species diagnosis. Thus hey would no conside a phyle ic basis o bacula a ia ion
o be app op ia e. They admi , howe e , ha while in some oden g oups he e a e pa e ns o
bacula mo phology ha do no ag ee wi h phyle ic di e gence as indica ed by o he mo pho-
logical ea u es, he e exis also g oss pa e ns o bacula a ia ion in o he g oups ha do in ac
con o m wi h accep ed phyle ic ela ionships. Indeed, hey ema ked ha he e can be li le doub
ha he baculum exhibi s phyle ic weigh and consequen ly may se e as a aluable axonomic
ool. Mo eo e , axa ha di e in ex e nal and c anial cha ac e s may ha e simila bacula, while
o he s ha a e simila in such ea u es may exhibi highly dis inc i e geni alia. Pa e son &
Thaele (loc. ci .) sugges ed ha al hough bacula mo phology e lec s phyle ic his o y on a g oss
scale, disco dance be ween pa e ns o bacula and phyle ic di e gence suppo s a unc ional
in e p e a ion o bacula a ia ion, especially a he species le el.
Simila ly, opinions a y as o he alue o bacula mo phology in chi op e an sys ema ics.
Hamil on ( 1 949) examined he baculum in No h Ame ican espe ilionids and concluded ha in
his amily he baculum was use ul in de ining ela ionships when conside ed wi h skull and o he
skele al cha ac e is ics. Thus he was able o sugges ha he close simila i y be ween he bacula o
Myo is (Fig. 19i, j) and Pizonyx (Fig. 19k) indica ed hei close ela ionship, and ha he dissimi-
la i y be ween he bacula o Pipis ellus sub la us (Fig. 2d) and P. Hespe us (Fig. 8d) sugges ed
gene ic o a leas subgene ic di e ence. This au ho also no ed ha in mos ins ances among
No h Ame ican espe ilionids he e we e ma ked gene ic di e ences in he baculum. He con-
side ed ha u he s udy was needed o de e mine he use ulness o he baculum in chi op e an
sys ema ics and ha wi h ime and su icien ma e ial he bone migh be u ilised in classi ica ion.
These conclusions we e ein o ced by K u zsch & Vaughan (1955) who examined he bacula o
u he No h Ame ican species. They ema ked ha in he case o hose ha a e closely ela ed
he baculum can se e as a c i e ion in judging ela ionship when o he clea cu dis inguishing
cha ac e s a e lacking. These au ho s ound bacula a ia ion in closely ela ed ba species o be
chie ly in shape, de ail o ou line, and g oss size: hei s udy led o he belie ha in a leas some
supe icially simila species well ma ked and consis en bacula di e ences ein o ced he mo e
sub le ex e nal and c anial dissimila i ies.
K u zsch (1959) accep ed he iew ha he baculum can p o ide addi ional e idence o classi i-
ca ion, o , in he absence o o he clea ly de ined cha ac e s, can se e as a c i e ion in judging
228 J. E. HILL & D. L. HARRISON
ela ionship. He added in ela ion o he P e opodidae ha by i ue o i s ela i e simplici y and
s uc u al s abili y he baculum migh well se e o help place en i ies o doub ul ela ionship in
hei na u al posi ion, al hough hey migh be o he wise mo phologically con adic o y. Among
p e opodids he ound ha in agene ic di e ences in he baculum in ol ed mino de ails o shape,
ou line and size. Gene a, howe e , migh be sepa a ed by mo e p o ound di e ences. His s udy
sugges ed ha al hough wi hin he genus well ma ked and consis en di e ences exis ed be ween
he bacula o indi idual species, he e was ne e heless a basic simila i y in pa e n h oughou he
genus, leading o he sugges ion ha ma ked a ian s om his mo phological s anda d in a single
genus migh p o ide g ounds o a eapp aisal o he a ini ies o he a ian . A u he s udy
(K u zsch, 1962) con i med hese opinions, especially in he b oad ag eemen o bacula a ia ion
in p e opodids wi h he axonomic a angemen o his amily by Ande sen (1912) and by Ta e
(1942 ). K u zsch concluded ha s ong in agene ic simila i ies exis among he bacula o p e o-
podids, bu ha ep esen a i e bacula o di e en gene a di e dis inc ly: al hough se ing well as
a sou ce o diagnos ic ea u es o he genus, he baculum does no seem o o e excep ional insigh
in o sup agene ic ela ionships. The la ge genus P e opus, howe e , o some ex en p o es o be
an excep ion, wi h a ious o i s many species demons a ing conside able a ia ion in bacula
s uc u e: on occasion di e ences be ween species equal hose be ween some megachi op e an
gene a.
Lanza ( 1 969) examined he baculum o P e opus in de ail and ound ha i s bacula mo phology
did no con o m o he classi ica ion p oposed by Ande sen (1912), a conclusion also eached by
Da is ( 1 947) who examined only i e species. Lanza ound ha in many cases bacula o an iden ical
size and shape could be ound among species belonging o he same g oup as well as o di e en
g oups; o ha he baculum could be ex emely di e en among o ms appa en ly o he wise e y
closely ela ed. Thus in his genus he ound he baculum o be o limi ed alue in phyle ic analysis.
Simila ly, LaVal (1973 ) ound ha wi h one excep ion he bacula o he a ious species o he
espe ilionid Rhogeessa a e no sha ply di e en ia ed om each o he : al hough in shape hey
show subs an ial geog aphic and indi idual a ia ion wi hin species hey seem ne e heless o
di e be ween species in a eas o sympa y o nea sympa y. Ha ison & B ownlow (1978) ound
ha indi idual a ia ion in he baculum o adul s o ano he espe ilionid, Sco ophilus, was such
ha i ende ed his s uc u e o li le o no alue in species diagnosis in his di icul genus.
Ma in (1978) discussed he adap i e alue o he baculum in ba s, ha ing ound a wide ange o
s uc u al a ia ion among se e al p e opodid species han was p e iously hough . He conside ed
ha he baculum may ha e a numbe o oles o a ying adap i e signi icance in suppo ing he
penis, as a s imula o y s uc u e, o in p e en ing u e h al closu e du ing he p essu es o copula-
ion. Al hough hese migh allow he baculum o main ain mo phological s abili y wi hin ce ain
axonomic uni s, his possible a iabili y o unc ion he hough ended o educe i s alue in
classi ica ion a he speci ic and subspeci ic le els.
Despi e hese possible limi a ions, many au ho s admi a leas he species-speci ici y o bacula
a ia ion among ba s, using he baculum o p o ide addi ional cha ac e s o sepa a e species ha
some imes o he wise closely esemble each o he . Some examples ha e been men ioned: o he s
include Wallin (1969) who d ew a en ion o bacula di e ences in Japanese Pipis ellus and who
used such di e ences o de ine wo species g oups in Vespe ilio, o Baag0e (1973) who u ilised
bacula cha ac e s in compa ing sibling species o Eu opean Myo is. Zubaid & Da ison (in p ess)
ound he baculum o be speci ically diagnos ic among Malaysian Hipposide os. In some gene a
au ho s ha e ou inely desc ibed and illus a ed he baculum o new species: o ins ance Sinha
(1969) in desc ibing Pipis ellus peguensis compa ed i s baculum wi h he bacula o he ela ed
species. Simila ly, McKean e al. (1978) desc ibed and illus a ed he baculum o 'Ep esicus'
sagi ula, compa ing i wi h he bacula o o he Aus alian 'Ep esicus', while Ki chene (1976)
employed he baculum o ' Ep esicus' douglaso um in he same way. Bacula cha ac e s some imes
o m an essen ial pa o e isiona y s udy, as by Ki chene e al. (1986) in de ining and keying he
Aus alo-Papuan ep esen a i es o Pipis ellus and Falsis ellus. The baculum has also ea u ed
in gene ic e ision, Hill (1966a) o example desc ibing and illus a ing ha o Phile o in he
cou se o such a s udy, o (1976) ha o he majo i y o he species o Hespe op enus.
Bacula a ia ion has also been employed o gene ic and subgene ic dis inc ion wi hin he
VESPERTILIONINE SYSTEMATICS 229
Vespe ilioninae. Wallin (1969) used penial cha ac e s in es ablishing Vespe ilio as a genus dis inc
om Ep esicus and in ecognising Hypsugo as a subgenus wi hin Pipis ellus, while Topal (1970a)
no ed ha bacula mo phology allied he abe an genus la mo e closely o Ep esicus han o
Pipis ellus wi h which i had been associa ed by some au ho s. Helle & Volle h (1984) summa -
ised published illus a ions o he bacula o Pipis ellus, Ep esicus and some o hei associa ed
gene a, d awing a en ion o hei axonomic implica ions. A a u he sys ema ic le el, Pine e al.
(1971) discussed he penial and bacula mo phology o An ozous and Baue us in ela ion o he
p esumed a ini ies o hese No h Ame ican gene a o he Aus alian and New Guinea gene a
Nyc ophilus and Pha o is wi h which hey ha e been associa ed in he sub amily Nyc ophilinae.
I is clea om he o egoing accoun ha he baculum is ega ded as a aluable sou ce o
diagnos ic in o ma ion by many s uden s o chi op e an sys ema ics. This seems especially ue in
he Vespe ilioninae, a sub amily in which diagnosis and de ini ion a bo h speci ic and gene ic
le els is some imes di icul i only he o hodox mo phological cha ac e s o ex e nal, c anial and
den al s uc u e a e o be elied upon.
Na u e and scope o his s udy
The basis o he cu en classi ica ion o he Vespe ilioninae was i s se ou in de ail by Mille
(1907), who ecognised a o al o hi y- wo gene a in he g oup, wi h diagnoses and sho desc ip-
i e accoun s. The classi ica ion o Mille was based chie ly on ex e nal, c anial and den al ea u es.
Ta e (1942a) e iewed he cha ac e s used o diagnosis in some de ail, di iding he sub amily
in o ou main ( ibal) agg ega ions, and a emp ing o quan i y he in e ela ionships o i s
many gene a. The majo ou lines o his classi ica ion ha e since been ollowed, some imes wi h
local modi ica ion as o ins ance by Koopman (19840, b, 1985) who subsumed he sub amily
Nyc ophilinae in o he Vespe ilioninae. Hill (1966) poin ed ou ha he sub amily comp ises a
complex o closely in e ela ed gene a sepa a ed in some ins ances by compa a i ely slende o
e en a he a bi a y dis inc ions, he pa e ns o ela ionship o en obscu ed by pa allelism o
con e gence.
The na owness o he o hodox dis inc ions ha de ine many o he cons i uen gene a o he
Vespe ilioninae has led o much axonomic combina ion and ecombina ion since Ta e w o e.
This si ua ion is exempli ied by he mo e ex eme a ian s o classi ica ion ha ha e been p o-
posed. Fo example, Kuzyakin( 1944, 1950, 1 965) included Pipis ellus and Ep esicus in Vespe ilio
while Simpson (1945) included Glisch opus, Sco ozous, Nyc alus and la in Pipis ellus and Rhinop-
e us, Hespe op enus, Tylonyc e is, Mime illus, Phile o , His io us and Laepho is in Ep esicus. A ye
mo e ex eme iewpoin was adop ed by Sokolo (1973) who conside ed ha Vespe ilio should
include no only Pipis ellus and Ep esicus as was hough by Kuzyakin, bu also all o he o he
abo e men ioned gene a excep Nyc alus. Ho acek & Hanak (19850, b) commen ed ha he
concep s o Kuzyakin and Sokolo (wi h he inclusion o Nyc alus} migh be p o isionally
accep ed, a leas un il ac ual p oo o pa aphylly in he g oup was o hcoming. Ne e heless,
hey conside ed his o be a e og ade solu ion since i exp esses no hing o he ac ual di e si y o
he g oup, p oposing ins ead ha he p oblema ic axa should be a anged in sepa a e gene a, hei
diagnoses hen making hei con en clea e hough na owe . Bo h Simpson and Sokolo also
included Sco oecus and Sco omanes in Nyc iceius as hen unde s ood, Baeodon in Rhogeessa,
Glauconyc e is in Chalinolobus and Dasyp e us in Lasiu us o p oduce a hea ily 'lumped' classi ica-
ion. The s a us o some such as Sco oecus, Dasyp e us and Idionyc e is has a ied om one au ho
o ano he o decades: in he Aus alian egion Sco eanax and Sco o epens ha e ecen ly achie ed
gene ic ank a e many yea s as nominal subgene a (Ki chene & Capu i, 1985) while la has once
again e e ed o Pipis ellus (Koopman 1984a, b, 1985) a e a b ie spell wi h Ep esicus. The
majo a ian s o espe ilionine classi ica ion a e summa ised in Table 1.
Many o he cha ac e s used o de ine axa and ela ionships among he Vespe ilioninae appea
s ongly adap i e and o equi ocal alue in gene ic and sup agene ic sys ema ics. Mos conce n
230 J. E. HILL & D. L. HARRISON
ea size and shape, agal s uc u e, he a chi ec u e o he skull, and he numbe and o ma ion o
he ee h. Zima & Ho acek (1985) poin ed ou ha he use o he mo phological cha ac e s
employed hi he o in he classi ica ion o he Vespe ilionidae as a whole migh no lead in a iably
o co ec axonomic conclusions, hei deg ee o di e en ia ion pe haps e lec ing he o ien a ion
and in ensi y o selec ion p essu e a he han ac ual phyle ic ela ionships. These au ho s indi-
ca ed an u gen need o new, su icien ly eliable and axonomically use ul c i e ia based on
ea u es ha did no possess a di ec adap i e signi icance, including among hese he mo phology
o he ep oduc i e o gans and he baculum.
Much weigh has been placed in he pas upon he p og essi e sho ening o he muzzle appa en
h oughou he Vespe ilioninae wi h concomi an educ ion and loss o he inciso s and p e-
mola s (Ta e, 1942a). In he incisi e den i ion he i s uppe oo h (i 1 ) 3 is absen , as in all ba s.
Reduc ion esul s in he emaining inne oo h (i 2 ) becoming peg-like and unicuspid, al hough
some imes qui e massi e, in a educ ion in size o he ou e oo h (i 3 ), i s displacemen inwa ds
o ou wa ds, o in i s e en ual obsolescence o loss. In he mandible, he i s (ij and second
(i 2 ) inciso ee h a e in a iably p esen , bu excep ionally he hi d (i 3 ) may be absen . The p ocess
also in ol es he educ ion and loss o he second uppe and lowe p emola s (pm}) and hen
o he an e io uppe p emola (pm 2 ): hus he p emola o mula anges om pm y o pm
5 i 5 he i s uppe and lowe p emola s (pm}) being p esumed o be hose ha a e absen om all
ba s. Se en di e en combina ions o inciso s and p emola s occu in he sub amily, i An ozous
and Baue us a e included. The ull complemen is usually aken as he p imi i e condi ion, he
educ ion and disappea ance o ee h as de i ed. These a e summa ised in Table 2, which gi es he
incisi e and p emola o mulae usually a ibu ed o each o he a ious gene a. Howe e , Myo is
occu in which pm 3 o pmj a e absen (Hill & Topal, 1973), hus in he i s ins ance p oducing
he o mula ypi ied by Lasionyc e is o Pleco us, in he second he o mula o Pipis ellus o
Nyc alus; pm 2 may be absen om Pipis ellus o gi e he a angemen o Ep esicus, o may be
p esen in Ep esicus o p oduce he o mula o Pipis ellus (Hill & Topal, loc. ci .); i 3 is a iable in
Sco ozous (o 45 examined, p esen in 34, absen om one side o he o he in 8, comple ely absen
om 3), when o ally absen o p oduce he incisi e o mula ha usually cha ac e ises Nyc iceius
and i s associa es (bu Thomas & W ough on ( 1 908) epo an example o 'Nyc iceius ' schlie enii
in which he le i 3 is p esen ); pm 2 is a iable in Sco oecus (Hill, 1974) and in Chalinolobus (Ryan,
1966; Koopman, 1971), and e y a ely may be p esen in 'Nyc iceius' schlie enii (Dobson, 1878;
Thomas, 1890).
Mos gene a o Vespe ilioninae can be de ined by o he ea u es besides hose o he incisi e and
p emola den i ion, al hough some imes only in di e ing combina ions. Thus al hough some
species exis ha combine he ex e nal ea u es o Myo is wi h he den al o mula o Pipis ellus o
he ex en ha ini ially hey (annec ans, idleyi) we e desc ibed in he la e genus, o he cha ac e s
such as he o m and s uc u e o he agus and he s uc u e o he inciso s enable hem o be
e e ed con iden ly o Myo is (Topal, 19706; Hill & Topal, 1973). Ano he ( osse i) was i s
desc ibed in Glisch opus, subsequen ly emo ed o Pipis ellus by Hill (1969) and inally (wi h
idleyi) o Myo is by Hill & Topal (loc. ci .). Howe e , he gene a Pipis ellus and Ep esicus do no
o e u he con en ional cha ac e s in his way and a e sepa a ed o he mos pa by he
p esence o pm 2 in he o me and i s absence in he la e . Wallin (1969) and Hill & Topal (loc. ci .)
discussed he a iabili y o his oo h in Pipis ellus and Ep esicus in de ail, he la e au ho s
concluding ha he p esence o absence o pm 2 can ha e no uni e sal alidi y in de ining he wo
gene a. Helle & Volle h (1984) also examined he ele ance o pm 2 in sepa a ing Pipis ellus and
Ep esicus and concluded ha i does no seem o be a eliable cha ac e is ic, a classi ica ion based
on i pe haps mis ep esen ing ue ela ionship. Ta e ( 942a) ecognised his di icul y bu adhe ed
o he con en ional p ac ice o sepa a ing he wo gene a by his ea u e, and indeed he majo i y o
au ho s ha e e ained he dis inc ion as a ma e o con enience, o en using he ex en o educ ion
and deg ee o displacemen o pm om he line o he oo h ow as a diagnos ic ea u e be ween he
species o Pipis ellus.
"Den al no a ion o Mille (1907)
VESPERTILIONINE SYSTEMATICS 23 1
Koopman (1975) has commen ed upon his p oblem. This au ho examined A ican species
alloca ed a iously o Pipis ellus and Ep esicus in an a emp o ind some o he cha ac e ha
would di ide his la ge assemblage in o wo majo g oups. He could ind none among he usual
sui e o mo phological ea u es. Al hough he ound no A ican Pipis ellus species ha closely
esembled he ho en o us, enuipinnis o lowe i g oups o Ep esicus and no A ican membe o he
la e genus ha was simila o hepipis ellus, Hespe us, sa ii o ueppellii g oups o Pipis ellus as
he de ined hem, he did ind a esemblance amoun ing in some cases o i ual iden i y (i pm 2 was
igno ed) be ween he kuhlii g oup o Pipis ellus and he Ep esicus capensis g oup. Expanding a
iew i s exp essed by Ta e (19420) in his accoun o Ep esicus, Koopman commen ed ha i is
p obable ha he an e io uppe p emola has been los mo e han once he e, and ha ue phyle ic
ela ionships un ac oss he 'gene ic' line. He hough ha i is e en possible ha in some cases a
'Pipis ellus species' and an 'Ep esicus species' a e ac ually conspeci ic, bu was o he opinion
ha he a ailable ma e ial was insu icien o es ablish his wi h ce ain y o any such pai a
he p esen ime. Al hough e aining Pipis ellus and Ep esicus as sepa a e gene a since he
belie ed ha he p oblem should be a acked on a cosmopoli an basis, he ema ked ha such an
a angemen is almos ce ainly w ong. Mo e ecen ly, Ho acek & Hanak (1985-1986) ha e
o e ed u he de ini ions o Pipis ellus, Hypsugo and Ep esicus.
Many a ying in e p e a ions can be placed upon ex e nal, c anial and den al mo phology o on
ka yological da a in he Vespe ilioninae. These ange om he ela ionship o one species o
ano he o sup agene ic ela ionships, e en o he iew ha Pipis ellus and Ep esicus may be
polyphyle ic. Menu (1984), o example, ema ked ha an exhaus i e odon ological s udy o
he Vespe ilioninae indica ed ha Pipis ellus includes species w ongly associa ed by eason
o hei iden ical den al o mula, bu which a e no ela ed. Williams & Ma es (1978) discussed he
ka yology o Ep esicus, which as cu en ly de ined hey hough seemed o be a composi e axon,
encompassing pe haps se e al phyle ic lines o pipis elloid species wi h educed numbe s o
p emola s. Helle & Volle h (1984) sugges ed ha Pipis ellus may be a e y he e ogeneous
assemblage and a e e iewing he ele ance o pm 2 in sepa a ing his genus om Ep esicus
conside ed he baculum o be a mo e eliable guide o he phylogeny o he species o Pipis ellus
and Ep esicus, using i s ea u es o supplemen hei indings om ka yological da a. Many yea s
be o e his Ta e (19420) ema ked ha i seemed p obable ha s udy o he baculum in he
Mic ochi op e a would yield aluable esul s, wi h he implica ion ha his migh ha e signi i-
cance in he classi ica ion o he Vespe ilionidae. Indeed, Ta e eco ds ha G. M. Allen had
ga he ed oge he a numbe o bacula ep esen ing many o he species o Pipis ellus which he
in ended o employ in e ising he genus. Mo eo e , ela i ely ew species o he nominal gene a
Pipis ellus and Ep esicus ha e so a been s udied: he imp ession gained om he li e a u e is
ha Pipis ellus as cu en ly unde s ood is digni ied chie ly by a long, slende sha ed baculum and
mos Ep esicus as i is p esen ly classi ied by a small, iangula s uc u e, which we ha e ound no
o be he case.
Ini ially ou in en ion was o examine he bacula o as many species o Pipis ellus as possible o
es ablish he pa e n o bacula a ia ion wi hin he genus, and o compa e i wi h he species
g oupings p oposed by Ta e (19420) and by Koopman (1973, 1975). As he wo k p og essed,
howe e , i became inc easingly appa en ha i s implica ions ex ended a beyond he limi s o
his nominal genus and ha i was necessa y in addi ion o s udy he bacula o Ep esicus so a as we
we e able, and o examine he bounda y be ween hese wo con en ional g oupings. Finally, o
place ou indings in p ope pe spec i e, we ha e su eyed he bacula o mos o he emaining
gene a o he Vespe ilioninae and ha e a emp ed o assess he gene ic signi icance o bacula
a ia ion in he sub amily. We ha e also examined he bacula o Nyc ophilus and Pha o is, bo h
usually e e ed o he closely ela ed sub amily Nyc ophilinae. This has been uni ed ecen ly wi h
he Vespe ilioninae (Koopman, 19840, b, 1985) and is hus ele an o ou s udy.
We ha e made no de ailed examina ion o he g oss mo phology o he chi op e an penis excep
inso a as i is e lec ed by bacula s uc u es. No ha e we a emp ed o s udy i s his omo phology.
These ea u es a e discussed by Smi h & Madkou (1980) in an e o o elucida e hei ele ance o
in e o dinal and in ao dinal phylogene ic ela ionships, and who e iew ea lie s udies o penial
mo phology.
232 J. E. HILL & D. L. HARRISON
Ma e ials and me hods
We ha e been able o examine bacula om he majo i y o species cu en ly lis ed in Pipis ellus
and om mos o hose p esen ly assigned o Ep esicus. In a ew ins ances we ha e elied
upon illus a ions and desc ip ions om he li e a u e. Simila ly, o he o he gene a o he
Vespe ilioninae ou s udy ma e ial has been d awn chie ly om specimens and o a much lesse
ex en om he published wo ks o o he s. The specimens ha we ha e examined a e lis ed in
Appendix 1 . Ou aim as a as gene a o he han Pipis ellus and Ep esicus a e conce ned has been
o p o ide illus a ions o ep esen a i e bacula, bu in hose ins ances whe e bacula s uc u e has
no be o e been s udied we ha e endea ou ed o examine as many species wi hin each genus as he
a ailable specimens pe mi ed. Clea ly, he ma e ial a ailable o us has been qui e inadequa e o
es ablish he ex en o indi idual a ia ion in any one species o subspecies. While only adul
specimens (wing epiphyses ully used) ha e been used, we ha e necessa ily had o accep ha o
he majo i y o species ou da a is limi ed. We ha e concen a ed he e o e on s udying and
compa ing he g oss mo phology (size, shape, g oss s uc u e) o he bacula ha we ha e examined
in an a emp o iden i y simila i ies, di e ences and gene al ends. The ine de ails pe haps mo e
aluable in species dis inc ion ha e ecei ed much less a en ion, al hough whe e i is known ha
species a e di icul o sepa a e by con en ional means a en ion has been d awn o bacula ea u es
ha may assis in iden i ica ion.
The specimens used in his s udy ha e been d awn almos en i ely om he collec ions o he
B i ish Museum (Na u al His o y), London (BM(NH)) and he Ha ison Zoological Museum,
Se enoaks, Ken (HZM). Apa om hese we ha e been able o examine one om he Na u -
his o isches Museum, Wien (NMW), by cou esy o D K. Baue , and one om he Ca negie
Museum o Na u al His o y, Pi sbu gh (CMNH), an anomalous specimen loaned o iden i ica-
ion by D D. A. Schli e , while D K. F. Koopman gene ously b ough o London an example o
Nyc iceius hume alis om he Ame ican Museum o Na u al His o y, New Yo k (AMNH) om
which a much needed baculum was ob ained. Specimens p epa ed many yea s ago a he B i ish
Museum (Na u al His o y) a e d y, some imes moun ed on ca d: he emainde ha e been p e-
pa ed in he cou se o his s udy. This has been accomplished by mace a ion o a sho pe iod in a
5% solu ion o po assium hyd oxide o which a small quan i y o aliza in ed has been added, a e
which he g osse mace a ed issue was emo ed by dissec ion, he specimen hen being clea ed and
s o ed in glyce in.
D awings ha e been p epa ed using ei he a s e eoscopic mic oscope wi h g a icule scale and
a ached came a lucida, o eehand using a simila ins umen . A ew we e d awn eehand using a
s e eo p ojec ion mic oscope wi h a elling mic ome e s age. The wide ange o size a ia ion
among espe ilionine bacula ( o example om a leng h o 1 mm o less o as much as 9 o 10 mm
in Pipis ellus) has necessi a ed he use o se e al scales o magni ica ion. So a as possible all
d awings on any one page o igu es a e a he same magni ica ion, wi h an app op ia e scale: o
acili a e compa ison he a ying magni ica ions used ollow an a i hme ic p og ession whe eby
each successi e la ge alue is wice i s p edecesso . I has no always been possible o con o m o
his a angemen , especially whe e d awings ha e been p epa ed om published illus a ions. As a
ule do sal (D) and igh la e al (RL) iews o each baculum a e p o ided: a ely h ough damage
he le la e al (LL) aspec is gi en. Occasionally whe e i is o especial in e es a hal en al (RVL
o LVL) d awing has been made, and in a ew ins ances whe e d awings ha e been aken om he
li e a u e i has been necessa y o gi e he en al (V) a he han he do sal aspec .
Au ho ship and esponsibili y
We ake join esponsibili y o he esul s and opinions pu o wa d and exp essed in his pape ,
and o he new names p oposed he ein.
The baculum o Pipis ellus
Fou bacula ypes ha e been iden i ied wi hin he nominal genus Pipis ellus. Wi h some excep-
ions, modi ica ion and combina ions, hese a e in b oad ag eemen wi h he g oupings o species
VESPERTILIONINE SYSTEMATICS 233
p oposed by Ta e (19420) and Koopman (1973, 1975). The classi ica ions o Ta e and Koopman
a e summa ised in Table 3.
(1) An elonga e s uc u e (Fig. la) wi h a slende sha and pai ed basal langes (e.g. Figs 2a-c,
3, 4, 5), he en al su ace o he p oximal pa o he sha ans e sely conca e, i s dis al pa
cylind ical o nea ly so; in p o ile he base in line wi h he sha o mo e o less de lec ed downwa d
a an angle o i ; he sha may be mo e o less s aigh , lexed o a iously cu ed in he e ical
plane, while he ip is gene ally bi id o o ked and may be di ec ed en ally o a g ea e o lesse
ex en .
Species agg ega ions in which his ype o baculum is ound include he ab amus, pipis ellus,
co omand a and ennis g oups o Ta e (19420); Koopman (1973) amalgama ed hese o o m a
pipis ellus g oup o which he added (1975) he A ican nanus and pe mix us. Howe e , nanus
(Fig. 6b) p o es o ha e a e y di e en baculum, as does imb ica us (Fig. 9a), included by Ta e in
he co omand a g oup and hus by Koopman ( 1 973) in he pipis ellus g oup. Pipis ellus babu (Fig.
4a), p o isionally placed by Ta e in he kuhlii g oup, also has he long, ela i ely s aigh baculum
cha ac e is ic o his pa o he di ision, as do endoi (Imaizumi, 1959) (Fig. 3b) and peguensis
(Sinha, 1969) (Fig. 15c), bo h desc ibed since Ta e w o e. The mo e ecen ly desc ibed wes alis
(Koopman, 1984c) (Fig. lOd), adamsi (Fig. lOc) and wa si (Fig. lOg) (Ki chene e al, 1986) also
belong wi h enuis (Fig. 9d) and i s allies in his g ouping. Taxa e e ed o he ceylonicus g oup by
bo h Ta e and Koopman (1973) p o e o ha e his bacula s uc u e, as do hose ha ha e been
examined o he ueppellii g oup (Figs 7e, , lOa, b) o Koopman (1975). Pipis ellus kuhlii and i s
associa es (Figs 5a-d, 6c) also belong in his di ision. In hese, howe e , he basal lobes o he
baculum a e sha ply angled o he sha in he e ical plane, and his bacula p o ile is e y
cha ac e is ic o kuhlii and i s ela i es. The sha is s aigh , wi hou lexion, and he ip is usually
bi id and no di ec ed en ally. Koopman (1975) included anchie ae (Fig. 6e) in he kuhlii g oup,
bu his p o es o ha e a e y di e en bacula con igu a ion.
A long-sha ed baculum o his ype occu s wi h li le modi ica ion in he majo i y o he
Aus alian species (Figs lla- , 12k) cu en ly e e ed o Ep esicus, in Nyc alus (Fig. lO ), in
Sco ozous (Fig. 16d) ( o which ueppellii and i s immedia e associa es ha e some imes been
e e ed) and in Sco oecus (Fig. 20a-e), in which he 'ho ns' o he bi id ip ex end in some ins ances
almos o o m a ing, a condi ion o eshadowed in Pipis ellus pa e culus (Fig. 3c). The Aus alian
Sco eanax (Fig. 16i) and Sco o epens (Figs 16g, h, 21e, also sha e his bacula ype: in Sco eanax
he 'ho ns' a he ip ha e become a ans e se ba , bu he species o Sco o epens e ain he bi id
o sligh ly bi id ip. A simila long-sha ed baculum bu wi h a simple ip occu s in he gene a
Hespe op enus (Fig. 21a-c, g) and Chalinolobus (Fig. 17a-e). The baculum o Glisch opus (Fig.
18a), al hough e y small, is also o his ype, wi h pai ed basal lobes, a slende sha , and bi id ip.
(2) A e y small s uc u e (Fig. Ib), consis ing o a b oad base wi h wo basal lobes (e.g. Figs 2d,
e, 9c, h), suppo ing a sho , e y sligh ly hollowed sha . This bacula ype is ound in sub la us
(Fig. 2d), ci cumda us (Fig. 2e), socie a is (Fig. 9c) and he mo e ecen ly desc ibed cup osus (Hill &
F ancis, 1984) (Fig. 9h).
(3) A ela i ely sho , s ou sha ed baculum (Fig. Ic), some imes wi h expanded base and ip
(e.g. Figs 6a, b, 7a, h, 8e, ), he base on occasion di ided in o pai ed lobes, some imes angled
e ically o he line o he sha , which is lu ed en ally a he han mos ly cylind ical; ip when
expanded ha ing i s an e io edge some imes di ided in o se e al i egula se a ions and on
occasion downwa dly di ec ed.
Such bacula a e ound in he sa ii g oup o Ta e (19420) and Koopman (1973, 1975) bu no in
made ensis (Fig. 5b) which was pu in o he sa ii g oup by bo h au ho s. I s baculum is howe e
qui e di e en and is like ha o kuhlii and i s associa es. Pipis ellus anchie ae (Fig. 6e), e e ed o
he kuhlii g oup by Koopman (1975) also belongs wi h sa ii, and he same bacula ype occu s in
nanus (Fig. 6b), alloca ed wi h pe mix us o he pipis ellus g oup by he same au ho . We ha e been
unable o examine he baculum o pe mix us bu ha o nanus and o helios (Fig. 6d) is o he ype
cha ac e is ic o his di ision, wi h i s basal pa qui e sha ply lexed o he sha al hough no
especially deep, and wi h an expanded, downwa dly di ec ed dis al pa . Pipis ellus eisen au i
240 J. E. HILL & D. L. HARRISON
dep ession; c anial p o ile almos s aigh om occipu o na es, sligh ly dep essed o e an e io
pa o o bi ; p emaxillae no sho ened; zygoma a slende , lacking any jugal eminence; in e den al
pala e longe han wide; maxilla y oo h ows pa allel o mos o hei leng h, an e io ly sligh ly
con e gen ; sho bony pos -pala e; sligh basial pi s; i 2 bicuspid, pos e io cusp 1/2-3/4 he heigh
o an e io cusp; i 3 abou he same in c own a ea o a li le la ge han i 2 , abou 1/2 o a li le mo e
i s heigh , wi h la ge cen al and smalle la e al accesso y cusps, lying pos e o-ex e nally o ha
oo h, sepa a ed om c 1 by a small dias ema; pm 2 la ge, un educed, i s c own a ea simila o ha
o i 2 o a li le less, sligh ly in uded bu sepa a ing c 1 and pm 4 ; i l _ 3 no much imb ica ed, i 3 abou
wice he bulk o i 1 _ 2 ; pm 2 no usually much educed, abou 1/2-3/4 o mo e he c own a ea o
pm 4 .
Included axa: aladdin, bac ianus, lac eus, na husii (Fig. 2b); medi e aneus, (?) pe mix us,
pipis ellus (Fig. 2a).
Among A ican Pipis ellus we ha e been unable o examine he baculum o pe mix us (Aellen,
1957) compa ed by i s desc ibe chie ly wi h na husii. I s den i ion, wi h bicuspid i 2 , he pos e io
cusp 2/3 he heigh o he an e io cusp, i 3 wi h la e al accesso y cusps, i s main cusp equal in heigh
o he pos e io cusp o i 2 , la ge, sligh ly in uded pm 2 which is abou as big as i 3 , and un educed
pm 2 , i s c own a ea abou 3/4-4/5 he c own a ea o pm 4 sugges s ha i should be e e ed o he
pipis ellus subg oup. Koopman (1975) e e ed i o he pipis ellus g oup.
(a) (ii) ja anicus (ab amus) subg oup. B aincase sligh ly globula , ele a ed pos e io ly; pos -
o bi al egion wide; sup ao bi al egion dis inc ly b oadened o p oduce ab up ly incu ing la e al
ma gins o he an e io pa o he pos o bi al a ea; os um b oad, do sally la ened, wi h no
mo e han an indica ion o a median os al dep ession; c anial p o ile almos s aigh om
occipu o na es, sligh ly la ened o e he occipu and a li le dep essed o e he an e io pa o
he o bi s; p emaxillae no sho ened; zygoma a slende bu no weak, lacking any jugal eminence;
in e den al pala e only li le longe han wide; pala e s ongly domed wi h b oad an e io ema gi-
na ion; maxilla y oo h ows mo e o less pa allel, sca cely con e gen an e io ly; sho bony
pos -pala e; shallow basial pi s; i 2 well de eloped, bicuspid, pos e io cusp some imes small,
usually abou 3/4 heigh o an e io cusp; i 3 simila in size o i 2 o sligh ly la ge , abou as high as i s
pos e io cusp, wi h la ge cen al and smalle la e al accesso y cusps, lying pos e o-ex e nally o
ha oo h, sepa a ed om c 1 by a na ow dias ema; pm 2 li le educed, equal o o a he less han
i 3 in c own a ea, in ecess be ween c 1 and pm 4 which app oach bu do no ouch; ij_ 3 sca cely
imb ica ed, i 3 as a ule simila in size o i 2 , bo h a li le mo e massi e han i^ pm 2 abou 1/2-3/4 he
size o pm 4 , e y sligh ly in uded om oo h ow.
Included axa: ab amus (Fig. 3a), akokomuli, babu (Fig. 4a), bancanus, camo ae (Fig. 15d),
endoi (Fig. 3b), i e i us, ja anicus (Fig. lOe), meyeni, peguensis (Fig. 15c), pa e culus (Fig. 3c),
pumiloides.
Cu en ea men s o Asian Pipis ellus usually include ab amus in P. ja anicus ( ala i ius ,
Lau ie & Hill, 1954) as a alid subspecies. The e appea o be ew con en ional ea u es ha clea ly
sepa a e ja anicus om ab amus bu hei bacula di e qui e sha ply in he high deg ee o e ical
lexion o he sha e iden in he la e . This di e ence was used by Thomas (1928a) who examined
Indo-Chinese Pipis ellus and di e en ia ed ab amus om ap o , ja anicus (as ala i ius) and
co omand a by i ue o he double cu a u e o i s baculum, he o he s being s aigh . Van Peenen
e al. (1969) eco ded co omand a, ja anicus and mimus om Vie nam bu he baculum ha hey
illus a e o ja anicus is clea ly ha o ab amus. This bacula di e ence sugges s ha ja anicus and
ab amus should be conside ed speci ically dis inc e en al hough he e seem o be ew c anial and
den al cha ac e s o sepa a e hem. The b aincase in ja anicus is sligh ly mo e in la ed han in
ab amus and i s os um na owe , he pala e is usually a li le wide in ela ion o i s leng h and is
sligh ly mo e exca a ed and domed, while pm 2 is a li le less educed and less in uded, ending
a he mo e o sepa a e c 1 and pm 4 . Bo h occu in Vie nam (Thomas, 1928a; specimens lis ed
below). I seems likely ha bancanus and camo ae, which has an un lexed baculum, a e mo e
closely ela ed o ja anicus han o ab amus.
Soo a & Cha u edi (1980) ema ked ha Thomas (191 5c) had poin ed ou ha he baculum o
ab amus is doubly cu ed and ha in pa e culus i is s aigh , bu hey s a ed u he ha ma e ial o
VESPERTILIONINE SYSTEMATICS 241
pa e culus in he collec ions o he Zoological Su ey o India e ealed ha i s baculum is doubly
cu ed. Howe e , specimens in he collec ions o he B i ish Museum (Na u al His o y) e e ed o
pa e culus (some he o iginal ma e ial seen by Thomas) ha e ela i ely s aigh bacula when
compa ed wi h he sinuous baculum o ab amus. We ha e ound his sinuous baculum o be
cha ac e is ic o ab amus, o which pe haps he specimens seen by Soo a & Cha u edi should be
e e ed.
The e y elonga e baculum o pa e culus, wi h i s s ongly bi id ip, he 'ho ns' o which a e
de lec ed en ally and ex end o some ex en o o m a ing (Thomas, 191 5c) is eminiscen o he
baculum o Sco oecus. A e y long baculum is also ound in endoi, bu in his species he ip is
less s ongly bi id and he 'ho ns' a e de lec ed do sally. Bo h, howe e , a e clea ly e e able o
Pipis ellus on c anial and den al cha ac e s, Sco oecus being dis inguished especially by a massi e
unicuspid i 2 , he loss o i 3 , a g oo ed c 1 , and usually by he absence o pm 2 .
(a) (iii) co omand a subg oup. Small, wi h small, ounded b aincase, ele a ed pos e io ly and
sligh ly so on ally; pos o bi al egion wide; os um sho , ela i ely na ow; no median os al
dep ession; c anial p o ile s aigh o nea ly so om occipu o ip o os um; p emaxillae excep-
ionally sho ; zygoma a slende , wi hou jugal p ojec ion; in e den al pala e abou as long o a
li le longe han wide; sho bony pos -pala e: no basial dep essions; i 2 usually bicuspid, pos e io
cusp some imes e y small o a ely absen , when p esen abou 1/2 o a li le mo e he heigh o he
an e io cusp; i 3 equal o g ea e han i 2 in c own a ea, eaching o ip o i s pos e io cusp, wi h
la ge p incipal cusp and smalle la e al accesso y cusps, lying pos e o-ex e nally o he inne
oo h; pm 2 no much educed, nea ly as g ea o as g ea in c own a ea as i 3 , wi h well de eloped,
sligh ly inwa dly di ec ed poin ed cusp, in ecess be ween c 1 and pm 4 ; ^ _ 3 no much imb ica ed, i 3
a li le la ge han i : _ 2 ; pm 2 abou 1/2 c own a ea and heigh o pm 4 , sligh ly ex uded.
Included axa: adamsi (Fig. lOc), a ghanus, angula us, collinus (Fig. 4b), co omand a (Fig. 7c),
glaucillus, mimus (Fig. 7g), mu ayi (Fig. 4c), ni idus, papuanus (Fig. 2c), poncele i (Fig. 4d),
po ensis, p incipulus, sewelanus, s u deei; possibly subulidens which may howe e ep esen
ja anicus; enuis (Fig. 9d), ama us (Fig. 7b), wa si (Fig. lOg), wes alis (Fig. lOd).
(a) (i ) ceylonicus subg oup. La ge, wi h a he sho , b oad b aincase; wide pos o bi al egion;
some deg ee o sup ao bi al expansion; os um b oad, a he long; weak, di use median os al
dep ession; c anial p o ile sligh ly con ex, aised o e he on al egion; p emaxillae no mal, no
sho ened; zygoma a mode a e, wi hou jugal eminence o p ocess, in e den al pala e longe han
wide; maxilla y oo h ows pa allel; sho bony pos -pala e; sligh basial pi s; i 2 la ge and massi e,
bicuspid o almos unicuspid, wi h mode a e o small pos e io cusp abou 2/3 heigh o an e io
cusp; i 3 massi e, as la ge o la ge han i 2 , ex ending o o a li le beyond pos e io cusp o ha
oo h, wi h la ge p incipal cusp and smalle la e al accesso y cusps, lying pos e o-la e ally o i 2 ,
na owly sepa a ed om c 1 ; pm 2 la ge, nea ly as g ea o g ea e in c own a ea han i 3 , usually
illing he ecess be ween c 1 and pm 4 in o which i is in uded, hese almos in con ac labially; i : _ 3
sligh ly imb ica ed, i 3 a li le la ge han i 1 - 2 ' pm 2 almos as la ge in c own a ea as pm 4 , e y
sligh ly ex uded om he oo h ow.
Included axa: bo neanus, ceylonicus (Fig. 7d), ch yso h ix, indicus, (?) minahassae, ap o (Fig.
3d), shano um, subcanus.
An accoun o minahassae is gi en by Ta e ( 942a) who e e ed i o a minahassae g oup o
which i was he sole membe . The skull o he holo ype has ne e been desc ibed and Ta e's
ema ks a e based on a e e ed specimen in he Ame ican Museum o Na u al His o y, New Yo k
(AMNH 102359). I has a sho , high b aincase wi h udimen s o a sagi al c es , p ominen
sup ao bi al ube cles and slende zygoma a; i 2 is long, wi h well de eloped pos e io cusp, c 1
slende , lacking an accesso y cusp, pm 2 only sligh ly in uded, i s c own a ea g ea e han ha o i 3 ,
and i^-j sca cely imb ica ed. These ea u es sugges ha i his specimen ep esen s minahassae
he axon should be alloca ed o Pipis ellus (Pipis ellus) and p o isionally we place i in he
ceylonicus subg oup o he pipis ellus g oup, bu clea ly hese decisions can only be specula i e.
(b) ueppell i g oup
Baculum as in pipis ellus g oup; b aincase high, b oadened, ounded and globose; pos o bi al
242 J. E. HILL & D. L. HARRISON
egion wide; sup ao bi al egion sligh ly expanded; os um sho ; wi h shallow, ill-de ined median
dep ession; c anial p o ile almos s aigh , a li le aised o e on al egion, a li le dep essed o e
os um; p emaxillae no sho ened; zygoma a slende , wi hou jugal p ojec ion; in e den al pala e
a li le longe han wide; maxilla y oo h ows sligh ly con e gen ; sho bony pos -pala e; no basial
pi s, ins ead a shallow dep ession; i 2 s ongly bicuspid, pos e io cusp abou 3/4 heigh o an e io
cusp; i 3 usually e y small o minu e, i s c own a ea less han 1/2 ha o i 2 , i s ip some imes ba ely
ising abo e he cingulum o he inne oo h, on occasion (e.g. nanulus) la ge , equal o o sligh ly
exceeding i 2 in c own a ea, abou 1 /2 o a li le mo e he heigh o ha oo h; i 3 lying subla e ally o
i 2 , sepa a ed om c 1 by a wide dias ema; pm 2 no usually g ea ly educed, i s c own a ea simila o
ha o i 2 , wi h s ong cusp, sepa a ing c 1 and pm 4 , occasionally (c assulus) much educed, simila
in size o i 3 in i s much educed condi ion, o (c assulus, nanulus) ecessed be ween hese ee h; i _ 3
li le imb ica ed, i 3 sligh ly he la ges as a ule; pm 2 abou 3/4 o mo e as la ge in c own a ea as
pm 4 and abou 3/4 i s heigh , a ely (coxi, c assulus) mo e educed, abou 1/2 c own a ea and
heigh o pm 4 .
Included axa: P obably coxi', c assulus (Fig. le), uscipes, leucomelas, nanulus (Fig. l ),pulche
(Fig. lOa), ueppellii (Fig. 0b),senegalensis, e nayi.
Vansonia Robe s, 1946 is a ailable should u he sepa a ion o he ueppellii g oup be hough
jus i ied: an ea lie name, Alobus Pe e s, 1867 is p eoccupied.
(c) kuhlii g oup
Baculum o mode a e leng h wi h na ow cylind ical sha and pai ed basal lobes as in pipis ellus
and ueppellii g oups bu basal lobes s ongly angled o line o sha in e ical plane; b aincase low
bu no la ened, ounded, only sligh ly elonga e; pos o bi al egion wide; sup ao bi al egion no
widened o swollen; os um long, unwidened, wi h e y sligh median la ening; c anial p o ile
almos s aigh om occipu o na es, sligh ly aised o e on al egion, sligh ly dep essed o e
on o o bi s; p emaxillae sligh ly sho ened; zygoma a slende , weak, wi hou jugal eminence;
in e den al pala e longe han wide; maxilla y oo h ows almos pa allel; sho bony pos -pala e;
small, na ow basial pi s; i 2 usually unicuspid, a bes only sligh ly bicuspid; i 3 small, i s c own a ea
1/2 o less ha o i 2 , i s ip ex ending only sligh ly beyond he cingulum o ha oo h, o which i lies
la e ally o subla e ally, sepa a ed om c 1 by a mode a e o na ow dias ema; pm 2 small, simila in
c own a ea o i 3 , in uded o lie in ecess be ween c 1 and pm 4 , hese mo e o less in con ac ; i _ 3
mode a ely imb ica ed, i 3 sligh ly he la ges ; pm 2 educed, abou 1/2 o less he c own a ea and
heigh o pm 4 .
Included axa: P obably ae o; dese i(aegyp ius, Qumsiyeh, 1985) (Fig. 5c), usca us, ikwanius;
p obably inexspec a us; kuhlii (Fig. 5a), made ensis (Fig. 5b), ma ensis, us icus (Figs 5d, 6c).
We ha e been unable o examine he baculum o inexspec a us (Aellen, 1959) bu his axon was
placed in he kuhlii g oup by Koopman (1975) who also e e ed made ensis o he sa ii g oup.
Howe e , an example o made ensis in he collec ions o he B i ish Museum (Na u al His o y) has
a baculum clea ly o he kuhlii ype.
Romicia G ay, 1838 is a ailable o he kuhlii g oup should his be hough wo hy o u he
sepa a ion.
Subgenus Pipis ellus ( Vespadelus)
Baculum usually wi h long cylind ical o en ally sligh ly lu ed sha , pai ed basal lobes and a
blun ip; sha sho e and wide in sagi ula; basal lobes some imes lexed o line o sha in
e ical plane; b aincase sligh ly b oadened, la ened and elonga ed; pos o bi al egion wide;
sup ao bi al egion sligh ly b oadened; os um sho bu no g ea ly widened; shallow median
os al dep ession; c anial p o ile almos s aigh om occipu o na es, a li le dep essed o e
os um; p emaxillae no sho ened; zygoma a slende , wi hou jugal p ocess; in e den al pala e a
li le longe han wide; maxilla y oo h ows sligh ly con e gen an e io ly; sho bony pos -pala e;
no basial pi s: i 2 bicuspid, pos e io cusp almos as high as an e io cusp; i 3 much educed, i s
c own a ea 1 /2 o less ha o i 2 , i s ip ba ely ex ending beyond he cingulum o ha oo h, o which
i 3 lies pos e o-la e ally, sepa a ed om c 1 by a na ow dias ema; pm 2 almos in a iably absen ,
VESPERTILIONINE SYSTEMATICS 243
when p esen a small spicule in ecess be ween c 1 and pm 4 : i { _ 3 mode a ely imb ica ed, i 3 sligh ly
he la ges , pm 2 g ea ly educed, in c own a ea abou 1/2 o mo e usually less he c own a ea o
pm 4 , and 1/2 i s heigh .
Included axa: cau inus (Fig. lib), douglaso um (Fig. 1 ld),pumilus (Figs 1 la, 12k), egulus (Fig.
1 le), sagi ula (Fig. 1 1 0, ul u nus (Fig. 1 Ic).
Fo me ly e e ed o Ep esicus, he ans e o hese axa o Pipis ellus was i s sugges ed by
Helle & Volle h (1984), pu ely on bacula g ounds. The bacula , c anial and den al ea u es o his
g oup sugges ha i ep esen s P. (Pipis ellus) in Aus alia, he ew membe s o his subgenus
(adamsi, wes alis and pe haps ja anicus) ha also occu he e being possibly sligh ly less di e en-
ia ed by i ue o hei ela i ely sligh ly less sho ened skulls and hei e en ion o pm 2 . The
pipis ellus g oup o he subgenus ex ends widely h ough he islands o Indo- Aus alia o New
Guinea, he Solomon Islands and Aus alia, chie ly as he co omand a subg oup, o which adamsi
and wes alis belong. Theja anicus subg oup eaches a leas o Ja a and Sulawesi and may ex end
o Aus alia (Hill, 1983) bu he Aus alian eco d o ja anicus is based on wo old examples and
has ne e been con i med. Possibly he sligh ly di e en ia ed pumilus and i allies esul om a
u he pe haps ea lie in asion o Aus alia. Bacula di e ences in his subgenus (Figs 11, 12k)
sugges ha i may consis o wo g oups: i has been possible o examine only pumilus.
The e has been hi he o a wide geog aphical hia us in he Indo-Aus alian dis ibu ion o
Ep esicus as o me ly unde s ood. Beyond hese Aus alian o ms, no o he axon a ibu ed o
his nominal genus has been epo ed u he eas in Indo-Aus alia han sou he n Thailand, o he
han an uncon i med eco d om Sa awak o Ep esicus sp. (Pi lo , 1968) which p o ided no
de ails.
Subgenus Pipis ellus (Pe imyo is)
Baculum e y small, Y-shaped, wi h pai ed basal lobes and e y sho sha ; b aincase sligh ly
elonga e, ounded, almos globose; pos o bi al egion wide; sup ao bi al egion sligh ly b oad-
ened; os um long, ele a ed, sligh ly widened; shallow median on al dep ession; a e y sligh
la e al dep ession on each side jus an e io o he o bi al im; c anial p o ile sinuous, aised
o e on al egion, a li le dep essed o e on o o bi s; p emaxillae no sho ened; zygoma a
mode a e, a sligh jugal eminence; in e den al pala e longe han wide wi h wide an e io pala al
ema gina ion; maxilla y oo h ows con e gen an e io ly; e y sho bony pos -pala e; sligh
basial dep essions; i 2 bicuspid wi h well de eloped pos e io cusp abou 3/4 heigh o an e io cusp;
i 3 massi e, i s c own a ea exceeding ha o i 2 , in heigh eaching o exceeding he heigh o an e io
cusp o ha oo h, wi h la ge p incipal cusp and smalle la e al accesso y cusps, lying pos e o-
ex e nally o inne oo h, sepa a ed om c 1 by a wide dias ema; pm 2 la ge, i s c own a ea equal o
ha o i 3 , in oo h ow, some imes sepa a ed om pm 4 by a sligh dias ema; i 2 " 3 and pm 2 almos
iden ical o hose o P. na husii; i^_ 3 no imb ica ed, i 3 only sligh ly bulkie han i : _ 2 ; pm 2 no
g ea ly educed o comp essed in oo h ow, i s c own a ea abou 1/2 o mo e ha o pm 4 , abou
1/2-3/4 i s heigh ; agus myo ine, abou 1/2 heigh o ea , ape ing o blun poin .
Included axon: sub la us (Fig. 2d).
Menu (1984) p oposed he genus Pe imyo is o P. sub la us, chie ly on accoun o he ea u es o
he canine and pos -canine den i ion in which he belie ed his species o app oach Myo is. How-
e e , Hill & Topal (1973) in discussing Myo is osse i and M. idleyi which also combine he agal
ea u es o Myo is wi h he Pipis ellus den i ion (pm| absen ) no ed ha in Myo is i 2 is sho and
b oad, i s pos e io cusp wide basally han he an e io cusp, while in Pipis ellus his oo h is
linea , o en na owe pos e io ly han an e io ly. Also, in Myo is he p incipal cusp o i 3 is equal
o o exceeds ha o i 2 in heigh and he oo h is o en hooked o p oduce a canini o m appe ance
while in Pipis ellus i is lowe and is no hooked. In Myo is i 3 is usually much la ge han i l _ 2 bu
in Pipis ellus he e is as a ule no such g ea dis inc ion in size. The incisi e den i ion o sub la us
co esponds closely wi h ha o Pipis ellus.
The baculum o sub la us is o a ype no ound in Myo is. Menu (1984) s a ed on he basis o
published igu es ha he baculum app oached ha o ce ain Myo is and mo e pa icula ly ha o
Pleco us au i us. We ind no signi ican esemblance o he mo phologically a he s able, saddle-
244 J. E. HILL & D. L. HARRISON
like baculum o Myo is and al hough he e a e some simila i ies wi h he bacula o Pleco us au i us
(Fig. 19g) and P. ene i ae (Fig. 14d), ha o P. aus iacus (Fig. 19h) is nea e in s uc u e o he
myo ine baculum. The bacula ype ound in sub la us occu s in a simila o m in Pipis ellus
ci cumda us (Fig. 2e), P. socie a is (Fig. 9c) and P. cup osus (Fig. 9h). The e a e conside able
di e ences, howe e , be ween sub la us and ci cumda us and i s allies, no leas in he deg ee o
educ ion o pm 2 , his oo h in hese h ee species being e y small o absen .
The unsho ened os um and he den al ea u es o sub la us sugges ha i is nea es o P.
(Pipis ellus), which i appea s o ep esen in No h Ame ica. We ind hespe us, he o he No h
Ame ican species o Pipis ellus, o belong on bacula and den al g ounds o P. (Hypsugo). Thus
we do no suppo Menu's iew (p. 410, oo no e) ha Pipis ellus is limi ed o he Old Wo ld and
ha he lines leading o his genus did no en e he No h Ame ican con inen . The ma ked
di e ences be ween sub la us and hespe us indica e wo qui e di e en pipis elline g oups, as
Hamil on (1949) ema ked in ela ion o hei bacula, bu bacula and den al e idence sugges s
alliance o es ablished Old Wo ld g oupings, he baculum o sub la us being pe haps a educed
o m o he sha ed s uc u e ound in P. (Pipis ellus) , ha o hespe us a modi ica ion o he ype
ound in P. (Hypsugo) .
Subgenus Pipis ellus (Hypsugo)
Baculum usually sho , s ou , some imes wi h expanded base and ip; base a ely bilobed, some-
imes do sally ele a ed; sha gene ally la ened do so- en ally, some imes wide, i s unde side
ans e sely conca e o lu ed; ip en ally hollowed as an ex ension o en al lu ing o sha ,
when expanded an e io ly sub-squa e o sligh ly ounded, i s an e io edge some imes i egula ly
se a ed, ip some imes downwa dly di ec ed, i s la e al ma gins on occasion o ming wo b oadly
based, en ally di ec ed p ojec ions; pm 2 gene ally much educed, small, minu e, o a ely absen .
Wallin ( 1 969) conside ed Hypsugo a alid subgenus wi hin Pipis ellus bu included only P. sa ii:
Ho acek & Hanak (1985#, b) added cado nae and pul e a us and sugges ed he ele a ion o
Hypsugo o gene ic ank, subsequen ly (1985-1986) widening i s possible con en and conside ing
i gene ically dis inc .
(a) sa ii g oup
Pos o bi al egion, sup ao bi al egion and os um no g ea ly widened; sup ao bi al ube cles i
p esen small and unde eloped.
(a) (i) pul e a us subg oup. B aincase elonga e, in la ed; pos o bi al egion wide, sup ao bi al
a ea no b oadened; os um long, no widened; shallow on al dep ession; no median os al
dep ession; b oad, shallow la e al dep essions abo e an e io pa o o bi ; c anial p o ile
somewha sinuous, dep essed o e on o o bi s; p emaxillae no sho ened; zygoma a obus ,
wi h e y sligh jugal eminence; in e den al pala e longe han wide; maxilla y oo h ows almos
pa allel; mode a e bony pos -pala e; no basial pi s; i 2 bicuspid, pos e io cusp abou 3/4 heigh o
an e io cusp; i 3 la ge, wide, i s c own a ea equal o o sligh ly exceeding ha o i 2 , eaching o ip
o he pos e io cusp o ha oo h, wi h mode a e la e al accesso y cusps, lying pos e o-ex e nally
o he inne oo h, sepa a ed om c 1 by a mode a e dias ema; pm 2 abou equal o nea ly equal o
i 3 in c own a ea, in ecess be ween c 1 and pm 4 which a e closely app oxima ed; ^.3 sligh ly
imb ica ed, i 3 a li le he la ges ; pm 2 a li le less han 1/2 he c own a ea o pm 4 , 1/2-3/4 i s heigh .
Included axon: pul e a us (Fig. 8c).
(a) (ii) nanus subg oup. B aincase ele a ed, sligh ly in la ed, mo e o less globose bu a li le
elonga ed; pos o bi al egion wide; sup ao bi al a ea sligh ly widened wi h small sup ao bi al
swellings; os um no especially sho ened o b oadened; shallow median os al dep ession;
sligh la e al dep essions jus an e io o sup ao bi al egion; c anial p o ile sinuous, s ongly
dep essed and conca e o e os um; p emaxillae no sho ened; zygoma a slende , lacking jugal
p ojec ion; in e den al pala e longe han wide; maxilla y oo h ows sligh ly con e gen ; sho
bony pos -pala e; no basial pi s; i 2 unicuspid o wi h small pos e io cusp ex ending o abou 3/4 o
VESPERTILIONINE SYSTEMATICS 245
i s heigh ; i 3 wide, i s c own a ea sligh ly exceeding ha o i 2 , abou 1/2-3/4 he heigh o ha oo h,
ex ending almos o he ip o i s pos e io cusp, wi h sligh la e al cusps, lying pos e o-ex e nally
o he inne oo h, sepa a ed om c 1 by a wide dias ema; pm 2 abou 1/2-2/3 o a li le mo e he
c own a ea o i 3 , in uded in o ecess be ween c 1 and pm 4 , hese some imes in con ac o nea ly so;
i : _ 3 no o only e y sligh ly imb ica ed, i 3 sligh ly he la ges ; pm 2 abou 1/2 he c own a ea and
heigh o pm 4 .
Included axa: a abicus (Fig. 7a), culex, Helios (Fig. 6d); p obably musciculus; nanus (Fig. 6b),
s amp lii.
Cu en lis ings (i.e. Hayman & Hill, 1971; Koopman, 1975) uni e Helios wi h P. nanus as a
synonym o possibly as a alid subspecies. Howe e , he bacula ea u es o his pale o m sugges
ha i may ep esen a species dis inc om nanus wi h which i may be sympa ic in no he n and
eas e n Kenya and in he Sudan. No baculum has been a ailable o musciculus, which was placed
in a Hespe us g oup by Koopman (loc. ci .). Al hough i s incisi e and p emola den i ion ag ees
wi h he nanus subg oup i s placemen he e emains specula i e.
(a) (in) sa ii subg oup. B aincase a he low and la , elonga e a he han globose; pos o bi al
egion no especially widened; sup ao bi al egion unwidened o only sligh ly widened; os um o
mode a e leng h; a shallow median os al dep ession; usually sligh la e al os al dep essions jus
an e io o sup ao bi al and an e io o bi al im; c anial p o ile s aigh o sligh ly conca e;
p emaxillae no sho ened; zygoma a obus , o en wi h sligh jugal p ocess; in e den al pala e a
li le longe han wide; maxilla y oo h ows mo e o less pa allel; sho bony pos -pala e; shallow
o no basial pi s; i 2 unicuspid o wi h pos e io cusp, when p esen abou 3/4 heigh o an e io
cusp; i 3 simila o o exceeding i 2 in c own a ea, abou 1/2-3/4 he heigh o i 2 , wi h s ong
cen al cusp lanked by smalle la e al accesso y cusps, lying pos e o-ex e nally o mo e la e ally
(anchie ae) o he inne oo h, sepa a ed om c 1 by a s ong dias ema; pm 2 much educed, minu e
o absen , when p esen c own a ea less han 1/2 ha o i 3 , in ecess be ween c 1 and pm 4 , hese as a
ule in con ac ; ^ _ 3 sligh ly o mo e s ongly imb ica ed, simila in size o i 3 sligh ly he bulkies ;
pm 2 educed, abou 1/2 o less in c own a ea han pm 4 and abou 2/3 i s heigh .
Included axa: anchie ae (Fig. 6e); p obably a iel', p obably aus enianus; bodenheime i (Fig. 9 ),
caucasicus, da wini, mau us, sa ii (Fig. 6a).
We ha e been unable o examine he baculum o a iel. The baculum o a small Pipis ellus om
he Na u his o isches Museum, Wien ( om Sayala, Uppe Egyp ) en a i ely iden i ied as a iel is
illus a ed by Gaisle e al. (1972) bu is e iden ly o he kuhlii ype. Qumsiyeh (1985) employs he
desc ip ion o his baculum in his accoun o a iel. Howe e , D K. Baue in o ms us (in li .) ha
he specimen (NHW 10351) o which he baculum is igu ed by Gaisle e al. (loc. ci .) is no
e e able o a iel bu is ins ead a small dese i, an iden i ica ion clea ly suppo ed by i s bacula
s uc u e. Mo eo e , D Baue has loaned h ee simila ly small specimens, one male, he o he s
emale (NHW 27501-3) (leng h o o ea m 29-2, 28-9, 28-2; condylobasal leng h 11-0, 10-5. 10-9;
c-m 3 4-0, 3-8, 3-9) appa en ly om Uppe Egyp , ha also ep esen dese i: a baculum om his
sample is again exac ly o he kuhlii ype. The c anial (na ow b aincase, unexpanded os um,
sho b oad na ial and an e io pala al ema gina ions, na ow basioccipi al) and den al (long i 3 ,
minu e pm 2 ) ea u es o a iel clea ly indica e ha i belongs wi h sa ii, o which g oup Koopman
(1975) e e ed i .
syn ype o Ep esicusbicolo (Bocage, 1889)(BM(NH) 89.5. 1.3) (Fig. 9e) p o es o be iden ical
c anially, den ally and in bacula mo phology wi h Pipis ellus anchie ae (Seab a, 1900) ( ide
in a, p. 249). Howe e , he poin needs con i ma ion o o he wise by examina ion o he o he
syn ype in he Museu Nacional de Lisboa. I should be no ed ha bicolo is he p io name
(Honackie aL, 1982).
The ela ionship be ween he pul e a us , nanus and sa ii subg oups is illus a ed by a abicus and
bodenheime i, he bacula o which a e compa ed di ec ly by Ha ison (1982). The baculum o
a abicus (Fig. 7a) app oaches ha o anchie ae (Fig. 6e) ye c anially and den ally his species is
nea e o nanus (Fig. 6b), while ha o bodenheime i (Fig. 9 ) is like he baculum o pul e a us (Fig.
8c) bu c anially and den ally he species is close o sa ii (Fig. 6a). These combina ions o ea u es
link he h ee subg oups.
246 J. E. HILL & D. L. HARRISON
(a) (i ) Hespe us subg oup. Baculum a lu ed s uc u e, much like ha o pul e a us o bodenhei-
me i. B aincase low bu b oad, elonga ed; pos o bi al egion wide; sup ao bi al a ea sligh ly
widened; os um sho , no g ea ly b oadened; a shallow median on al dep ession; sligh la e al
os al dep essions jus abo e an eo bi al o amina; c anial p o ile almos s aigh , sligh ly
dep essed abo e an e io oo o zygoma a; p emaxillae no sho ened; zygoma a slende , a li le
widened an e io ly, lacking any jugal eminence; in e den al pala e abou as wide as long; maxilla y
oo h ows con e gen ; sho bony pos -pala e; no basial pi s; cochlea bullae in la ed wi h na ow
basioccipi al; i 2 unicuspid; i 3 sligh ly g ea e in c own han i 2 bu abou 1/2 i s heigh , wi h li le
ace o la e al accesso y cusps, lying pos e o-ex e nally, sepa a ed om c 1 by mode a e o small
dias ema; pm 2 small o minu e, a bes abou 1/2 o less in c own a ea han i 3 , in ecess be ween c 1
and pm 4 which a e closely app oxima ed; ij_ 3 sca cely o no imb ica ed, simila in size; pm 2
educed, abou 1/2 he c own a ea o pm 4 , a li le less han 1/2 i s heigh .
Included axon: Hespe us (Fig. 8d).
Ho acek & Hanak (1985a, b, 1985-1986) ha e indica ed ha hey in end o p opose gene ic
s a us o Hespe us and indeed ha e sugges ed ha i be e e ed o Pa as ellus which hey o e as a
new name. I is howe e a nomen nudum in hese publica ions. The e seem good g ounds o
conside ing Hespe us he No h Ame ican ep esen a i e o P. (Hypsugo) o which i s bacula ,
c anial and den al ea u es ally i . Like bodenheime i (Fig. 9 ) i s baculum app oaches ha o
pul e a us (Fig. 8c) bu c anially and den ally i is nea e o sa ii&nd i s immedia e allies. Koopman
(1975) e e ed Hespe us o a Hespe us g oup in which he also included he A ican musciculus, he e
p o isionally alloca ed o he nanus subg oup.
(a) ( ) eisen au i subg oup. B aincase b oad, ele a ed and globula ; in la ed on ally; pos -
o bi al egion wide; sup ao bi al egion b oadened, wi h small sup ao bi al ube cles; os um
sho , deep, wide and massi e; sligh median os al dep ession; c anial p o ile s aigh o sligh ly
con ex; p emaxillae no sho ened; zygoma a s ong, lacking any jugal p ojec ion; in e den al
pala e e y sligh ly longe han wide; maxilla y oo h ows almos pa allel; sho bony pos -pala e;
sligh basial pi s usually p esen ; i 2 long, na ow, bicuspid, pos e io cups abou 3/4 heigh o
an e io cusp; i 3 sho , wide, simila o o sligh ly g ea e in c own a ea han i 2 , abou 1/2 o a li le
mo e i s heigh , wi h la ge cen al cusp and smalle la e al accesso y cusps, lying la e ally and
sligh ly pos e io ly o he inne oo h, sepa a ed om c 1 by a mode a e dias ema; pm 2 small, abou
he same in c own a ea as i 3 , sandwiched in o ecess be ween c 1 and pm 4 , hese almos in con ac ;
i _ 3 sligh ly imb ica ed, i 2 _ 3 simila in size, bo h la ge han i ; pm 2 abou 1/2 c own a ea and
heigh o pm 4 .
Included axon: eisen au i (Fig. 9g).
Koopman (1975) places eisen au i in a ueppellii g oup, no doub on accoun o i s ele a ed,
in la ed b aincase and i s bicuspid i 2 , bu i s bacula ea u es do no associa e i wi h his species
and i s immedia e allies. I s baculum is e y simila o ha o imb ica us (Fig. 9a) o mac o is
(Fig. 9b).
(a) ( i) imb ica us subg oup. B aincase in la ed, globula , aised pos e io ly; pos o bi al egion
Wide; sup ao bi al a ea sligh ly widened wi h e y small sup ao bi al ube cles; os um sho ,
no especially b oadened; no median os al dep ession; c anial p o ile almos s aigh ; sligh ly
conca e abo e sup ao bi al egion; p emaxillae no sho ened; zygoma a mode a e o s ong,
some imes wi h a ace o a jugal eminence; in e den al pala e abou as wide as long, no domed;
maxilla y oo h ows almos pa allel; e y sho bony pos -pala e; well de eloped basial pi s; i 2
bicuspid, pos e io cusp abou 3/4 heigh o an e io cusp; i 3 simila in c own a ea o i 2 , abou 1/2
i s heigh , wi h la ge cen al cusp and smalle la e al accesso y cusps, lying la e ally o he inne
oo h, sepa a ed om c 1 by a na ow dias ema; pm 2 g ea ly educed, 1/4 o less he c own a ea o
i 3 , in ecess be ween c 1 and pm 4 , hese in con ac ; i l _ 3 sca cely imb ica ed, i 2 _ 3 o simila size, a
li le la ge han i ; pm 2 abou 1/2 he c own a ea and heigh o pm 4 .
Included axa: cu a us, imb ica us (Fig. 9a), mac o is (Fig. 9b), o de manni.
(a) ( ii) lophu us subg oup. B aincase in la ed, ounded, sligh ly elonga e, aised pos e io ly;
pos o bi al egion wide; sup ao bi al a ea li le widened; a bes only a ace o sup ao bi al
VESPERTILIONINE SYSTEMATICS 247
ube cles; os um mode a e in leng h, longe han in imb ica us subg oup, no b oadened; no
median os al dep ession; c anial p o ile almos s aigh , sligh ly dep essed o conca e abo e
sup ao bi al egion; zygoma a s ong wi h dis inc jugal eminence; in e den al pala e a li le longe
han wide; maxilla y oo h ows sligh ly con e gen ; mode a e bony pos -pala e; deep basial pi s;
inciso and p emola den i ion closely simila o ha o imb ica us subg oup bu i 3 lying mo e
pos e o-la e ally o i 2 , and pm 2 some imes (lophu us) sligh ly la ge , abou 1/2 c own a e o i 3 .
Included axa: cado nae, ki chene i (Fig. 8e), lophu us (Fig. 80-
The baculum o ki chene i is unusual in he p esence dis ally o wo an e io do so-la e al,
pos e io ly di ec ed p ocesses, wi h en ally a mo e o less ape ed median gu e . As in lophu us,
he ip is di ec ed sligh ly en ally.
(b) s enop e us g oup
B aincase la ge, ounded and globula ; pos o bi al egion e y wide; sup ao bi al egion much
widened o include well de eloped sup ao bi al ube cles; os um sho , wide; shallow median
os al dep ession an e io o on al egion; c anial p o ile sligh ly con ex, ele a ed o e on al
a ea; p emaxillae no sho ened; zygoma a a he weak, lacking jugal p ocess bu usually wi h
small descending p ocess ex e nal o m 3 ; pala e sho and b oad, he in e den al pala e as wide as
long; maxilla y oo h ows pa allel o nea ly so; sho bony pos -pala e; shallow basial pi s; i 2
small, bicuspid, pos e io cusp 1/2-3/4 heigh o an e io cusp; i 3 a li le smalle in c own a ea han
i 2 , i s ip eaching almos o ip o he pos e io cusp o ha oo h, wi h la ge cen al cusp and
smalle la e al accesso y cusps, lying pos e o-la e ally o he inne oo h, only na owly sepa a ed
om c 1 o almos in con ac wi h i ; c 1 wi h dis inc , well de ined pos e io accesso y cusp; pm 2
small o minu e, abou equal in c own a ea o a li le la ge han i 3 (s enop e us) o abou 1/3-1/4
he c own a ea o his oo h (jo ei, an honyi), in ecess be ween c 1 and pm 4 , which ouch; i _ 3 no
much imb ica ed, all o simila size; c own a ea o pm 2 sligh ly exceeding ha o pm 4 , pm 2 simila
in heigh o he second oo h (s enop e us), o c own a ea o pm 2 abou 1/2 ha o pm 4 , pm 2 almos
as high as ha oo h (jo ei, an honyi).
Included axa: an honyi, jo ei, s enop e us (Fig. 7h).
The baculum o s enop e us is unusual, al hough o he sa ii ype: i has a na ow lobed base,
hollowed sha , and expanded ip he la e al ma gins o which p ojec en ally as wo b oadly
based 'ho ns'. The s enop e us g oup as he e unde s ood is he jo ei g oup o Ta e ( 1 942a) and (in
pa ) o Koopman (1973). Bo h jo ei and s enop e us ha e been e e ed in he pas o Nyc alus
bu as men ioned abo e (p. 234) he baculum o s enop e us has no esemblance o he long-sha ed
baculum o ha genus (Fig. 100- Ta e (19420) e e ed bo h o Pipis ellus wi h he commen ha
he g oup app oached O ien al membe s o he sa ii g oup, and migh a a la e ime be acco ded
gene ic ank.
Subgenus Pipis ellus (Falsis ellus)
Baculum a b oad, p oximally widened and en ally deeply lu ed s uc u e wi h no dis al expan-
sion; b aincase elonga e; pos o bi al egion wide; sup ao bi al a ea no expanded; os um long,
no b oadened; zygoma a mode a e o s ong; and pala e a he na ow, he in e den al pala e
longe han wide.
Pipis ellus (Falsis ellus) appea s o be ela ed o P. (Hypsugo) o which i may be he eas e n
ep esen a i e. I is app oached in bacula mo phology by some o he la e subgenus such as
imb ica us (Fig. 9a), mac o is (Fig. 9b), ki chene i (Fig. 8e) and lophu us (Fig. 8 ), and indeed he
baculum in P. (Falsis ellus) appea s o be an ex eme a ian o he b oad, en ally lu ed
s uc u e o many o P. (Hypsugo).
Ki chene e al. (1986) aised Falsis ellus o gene ic ank bu did no include a inis andpe e si,
con ining hei compa isons o he Aus alasian Pipis ellus (i.e. adamsi, angula us, collinus,
papuanus, wa si and wes alis) he e e e ed o he co omand a subg oup o P. (Pipis ellus). These
au ho s d ew a en ion o i s la ge size; o i s small i 3 which is an e io ly displaced and swi elled o
o a ed ou wa ds o lie alongside i 2 , i s conca i y acing ou wa ds (a ea u e which may ha e
in luenced I edale & T ough on, 1934 in placing i in Glisch opus); and o i s combina ion o
248 J. E. HILL & D. L. HARRISON
unicuspid i 2 , iny pm 2 and p onounced occipi al c es , which asTa e (19420) no ed gi es he ea o
he skull a 'helme ed' appea ance. Excep ing he la ge size and he p esence o a s ong occipi al
c es , hese ea u es occu elsewhe e in he a ious g oups o Pipis ellus: he ex en o he occipi al
c es may be a unc ion o he la ge size o he skull.
(a) a inis g oup
B aincase a he na ow, mas oid wid h ma kedly less han zygoma ic wid h; pos o bi al egion
wide; sligh , a he poo ly de eloped sup ao bi al idges; e y shallow median os al dep ession
jus an e io o on al egion; do so-la e al ma gin o os um mo e o less s aigh om pos -
o bi al cons ic ion o an e io edge o o bi ; c anial p o ile almos s aigh , sligh ly ele a ed
on ally, dep essed o e sup ao bi al egion; p emaxillae no sho ened; zygoma a mode a e o
obus wi h jugal eminence; maxilla y oo h ows sligh ly con e gen ; mode a e bony pos -pala e;
no basial pi s; i 2 s ongly bicuspid, pos e io cusp 3/4 o mo e he heigh o he an e io cusp; i 3
la ge in c own a ea han i 2 , i s heigh abou equal o he heigh o he pos e io cusp o ha oo h,
wi h s ong cen al cusp and smalle la e al accesso y cusps, lying pos e o-la e ally o he inne
oo h, sepa a ed om c 1 by a mode a e dias ema; pm 2 almos as g ea in c own a ea as i 3 (a inis) o
abou 3/4 i s c own a ea (pe e si), in ecess be ween c 1 and pm 4 ; i^ _ 3 mode a ely imb ica ed, i 3 he
la ges ; pm 2 abou 1/2 he c own a ea o pm 4 and abou 2/3-3/4 i s heigh .
Included axa: a inis (Fig. 8a), (?) mo dax, pe e si (Fig. 8b).
I has no been possible o examine mo dax. Indian eco ds o his species appea o be based on
specimens in he collec ions o he B i ish Museum (Na u al His o y) en a i ely labelled as such.
These, howe e , ag ee closely wi h he desc ip ion o P. a inis by Dobson (1871) and wi h he
accoun o a specimen e e ed o his species om Likiang, Yunnan by Ta e (19420). I co ec ly
alloca ed, mo dax (Pe e s, 1867) is he ea lies name in he g oup.
(b) asmaniensis g oup
La ge and dis inc i e; b aincase high, wi h well de eloped sagi al c es ; pos o bi al egion wide
and s ong; no median os al dep ession; c anial p o ile s aigh ; p emaxillae sligh ly sho ened;
zygoma a s ong wi h sligh jugal p ocess and small in e io p ocess; maxilla y oo h ows nea ly
pa allel; sho bony pos -pala e; sligh basial dep essions; i 2 la ge, unicuspid; i 3 small, i s c own
a ea abou 1 /4 ha o i 2 , ba ely ex ending abo e he cingulum o ha oo h o which i lies la e ally,
i s hollowed ace ou wa dly di ec ed, sepa a ed om c 1 by a mode a e dias ema: pm 2 e y small,
abou 1 /3 he c own a ea o i 3 , in ecess be ween c 1 and pm 4 , which ouch; i l _ 3 much imb ica ed, i 3
wice he bulk o i l - 2 pm 2 much educed, abou 1/4 he c own a ea o pm 4 and abou 1/2 i s
heigh .
Included axa: mackenziei (Fig. lOh), asmaniensis (Fig. 8g)
Subgenus Pipis ellus (Neo omicia)
Baculum wi h dis inc pai ed basal lobes, slende cylind ical sha and a iously expanded ip;
b aincase b oad, some imes sligh ly elonga e, a he la ened; pos o bi al egion wide; sup a-
o bi al a ea unwidened o only sligh ly b oadened; os um mode a e o sligh ly leng hened;
c anial p o ile s aigh ; p emaxillae sho ened; zygoma a mode a e, no jugal p ocess; maxilla y
oo h ows only sligh ly con e gen ; sho bony pos -pala e; no basial pi s; i 2 unicuspid o wi h
small pos e io cusp ex ending o abou 3/4 i s heigh ; i 3 smalle han i 2 , i s c own a ea 3/4-1/2 o
less ha o he inne oo h, abou 1/2 i s heigh , wi h usually a la ge cen al cusp and sligh la e al
accesso y cusps, he inne cusp as a ule e y small, he oo h an e io ly displaced o lie alongside o
almos alongside i 2 , sepa a ed om c 1 by a mode a e o wide dias ema; pm 2 almos in a iably
absen , when p esen e y small, in ecess be ween c 1 and pm 4 , i l _ 3 sligh ly o mode a ely imb i-
ca ed, o simila size o wi h i 2 _ 3 a li le he la ge ; pm 2 educed, i s c own a ea 1/2 o less ha o
pm 4 and i s heigh 1/2-3/4 o he heigh o ha oo h.
This subgenus is wholly A ican and hi he o i s membe s ha e been e e ed o Ep esicus,
al hough he e is ka yological e idence ( ide in a) sugges ing ha one a leas should be mo ed o
Pipis ellus. I inco po a es he Ep esicus capensis and E. enuipinnis g oups o Koopman (1975).
VESPERTILIONINE SYSTEMATICS 249
These can be ecognised eadily by he s uc u e o he baculum, capensis and i s allies (Fig. 12a-d,
-i) ha ing he dis al pa o he baculum spa ula e and en ally de lec ed, enuipinnis and i s
associa es (Fig. 12e, j) ha ing he baculum modi ied dis ally in o a lobed, almos e ical pla e-like
s uc u e.
Published ka yological da a e e s only o capensis, al hough s udies o o he membe s o he
subgenus a e in p og ess (Rau enbach & Schli e , 1985a, b). Pe e son & Nago sen (1975) ound
ha capensis has a diploid numbe o 32 and a undamen al numbe o 50: Williams & Ma es
(1978) discussed he possible composi e na u e o Ep esicus as sugges ed by Koopman (1975) and
poin ed ou ha he species i ed ka yologically wi hin he a ia ion exhibi ed by Pipis ellus. This
genus has a diploid numbe a ying om 26 o 44, and undamen al numbe s om 44 o 60, hese
indings appa en ly suppo ing Koopman's obse a ions. These au ho s ema ked, howe e , ha
he ka yo ype o capensis is mo e simila o ha o Pipis ellus nanus (2N = 36, FN = 50) han o P.
kuhlii (2N = 44, FN = 50), Koopman ha ing hough capensis nea e o he kuhlii g oup han o he
pipis ellus g oup in which he placed nanus. Williams & Ma es (loc. ci .) also ound, in con as ,
ha small Ep esicus om he New Wo ld (diminu us, u inalis) ha e he ypical 'ep esicoid' ka yo-
ype (2N = 50, FN = 48-50), and added ha he ka yo ypic di e ences be ween Ep esicus (sensu
s ic o) and Pipis ellus migh p o e mo e use ul o sepa a ing hese gene a han o he s uc u al
ea u es.
Ou s udy o he bacula o A ican 'Ep esicus ' con i ms hese obse a ions and indica es he
isola ion o capensis, enuipinnis and hei ela i es om Ep esicus sensu s ic o ( ide in a): Helle
& Volle h (1984) also ans e ed capensis o Pipis ellus, en i ely on accoun o i s published
ka yology. I is in e es ing o no e also ha he baculum o P. nanus indica es ha his species
should be e e ed o P. (Hypsugo) a he han o P. (Pipis ellus) whe e Koopman (1975) e ec-
i ely alloca ed i . The bacula mo phology o capensis, enuipinnis and hei allies sugges s
s ongly ha hese o me g oupings o Ep esicus a e mos closely allied o P. (Hypsugo) as he
ka yological simila i y o capensis o P. nanus indica es. The an e io uppe p emola (pm 2 ) is e y
small, es igial o absen in P. sa ii and is e y small in mos o he membe s o P. (Hypsugo): e y
a ely i is p esen in capensis (Wallin, 1969; Hill & Topal, 1973). On he same poin , we ha e been
able o examine a specimen (MJS 2846) om Somalia, in he Ca negie Museum o Na u al
His o y, which has a small pm 2 on bo h sides o he jaw, leading o i s e s while iden i ica ion as
Pipis ellus dese i. The baculum, howe e , is cha ac e is ically ha o 'Ep esicus' somalicus, which
in ac he specimen ep esen s.
Koopman (1975) sugges ed ha Vespe us bicolo Bocage, 1889 ( = Ep esicus bicolo ) and
Pipis ellus anchie ae (Seab a, 1900), bo h om Angola, may be conspeci ic, ha ing examined
syn ypes o bo h a he B i ish Museum (Na u al His o y). This au ho hough ha bicolo migh
be a o m o 'Ep esicus ' enuipinnis as Hayman & Hill (1971) sugges ed, o ha i migh be based on
a specimen o Pipis ellus anchie ae wi h missing an e io uppe p emola s. Bocage (loc. ci .) says
'pas de ace de la p emie e p emolai e a la machoi e supe ieu e'. Fu he s udy o he syn ype
(BM(NH) 89.5. 1 .3) in London shows i o ha e a small pm 2 in a ecess be ween c 1 and pm 4 on each
side: c anially i ag ees exac ly wi h he syn ype o anchie ae (9 BM(NH) 6. 1 .3. 1) and i s baculum is
exac ly as in ha species. Cu iously, Bocage s a es ha bo h o iginal specimens o bicolo a e
emale. The specimen in London is qui e clea ly lis ed as a 'Co- ype' by Thomas in he ele an
accession egis e .
(a) capensis g oup
Tip o baculum la ened, de lec ed en ally, some imes a small sub-apical do sal p ojec ion;
b aincase la ened, sligh ly elonga e; os um no especially b oadened; pala e long, na ow,
in e den al pala e longe han wide; i 3 1/2 o less he c own a ea o i 2 .
Included axa: capensis (Fig. 12b, g); p obably b unneus, ga ambae, g andidie i; guineensis
(Fig. 12c), ma oka (Fig. 12a), melcko um (Fig. 12 ); minu us (?) (Fig. 12i); p obably ec i agus;
somalicus (Fig. 12h); p obably ansoni; zuluensis (Fig. 12d).
The baculum o b unneus sensu s ic o has no been examined. Tha (Fig. 14b) o a Nige ian
specimen (BM(NH) 48.702) collec ed by I.T. Sande son and hi he o e e ed o his species is
e y simila o ha o endalli (Fig. 12e), wi h which his example ag ees in c anial and en al
256 J. E. HILL & D. L. HARRISON
Dobson he hough o ha e bu one pai o uppe inciso s. T ouessa (1897) ini ially ollowed
Dobs~>n (1878) in alloca ing schlie enii o Sco ozous as a subgenus o Vespe ugo, bu la e (1904)
changed his opinion o conside Sco ozous a subgenus o Sco ophilus. Mille (1907) e e ed
schlie enii o Sco einus, al hough in ac he species does no display he educ ion o m| ha he
conside ed diagnos ic o his genus and which occu s in he Aus alian species (bals oni, g eyii,
now inco po a ed in o Sco o epens) ha he alloca ed o i . Mille 's iew was adop ed by Thomas &
W ough on (1908) and in di e ing ways by many subsequen au ho s. Howe e , Allen (1911)
when desc ibing a icanus e e ed i o he hi he o Ame ican genus Nyc iceius, commen ing on i s
simila i y o N. hume alis and Hollis e (1918) ema ked ha Old Wo ld ba s usually placed in he
genus Sco einus did no seem o di e gene ically om he Ame ican species o Nyc iceius, o which
he also e e ed a icanus. Since hen a icanus has been elega ed o subspeci ic s a us o synonymy
in schlie enii (B aes up, 1935; Allen, 1939; Aellen, 1952). B aes up (loc. ci .) also employed
Nyc iceius o schlie enii in p e e ence o Sco einus, and poin ed ou ha i s las uppe mola was
no educed in he way ha Mille (1907) had desc ibed o ha genus. This au ho d ew a en ion
o he a ini y hus es ablished be ween he E hiopian and Ame ican aunas, bu did no exclude he
possibili y o con e gen e olu ion om di e en Pipis ellus- ike o ms. Ta e (1942a) main ained
Nyc iceius and Sco einus as dis inc gene a bu Simpson (1945) uni ed hem, a lead ollowed by
many mode n au ho s who ha e conside ed Sco einus a subgenus o Nyc iceius. Thus Elle man &
Mo ison-Sco (1951) and Elle man e al. (1953) e e ed schlie eni o Sco einus as a subgenus
o Nyc iceius, while Lau ie & Hill (1954) lis ed he Aus alian species be o e hen alloca ed o
Sco einus in Sco eanax and Sco o epens as u he alid subgene a o Nyc iceius. On he o he
hand, Rose ea (1965) conside ed Nyc iceius and Sco einus synonymous. Koopman (1965)
e e ed schlie enii o Sco einus as a subgenus o Nyc iceius bu la e (in li . in Hayman & Hill,
1971) e ised his opinion o alloca e i o Nyc iceius (Nyc iceius), since hen (1978) ein o cing his
iew.
The classi ica ion o he Vespe ilioninae
Ea lie classi ica ions o he Vespe ilioninae (Mille , 1907; Ta e, 19420) ely hea ily on he pa e n
o educ ion o he inciso and p emola ee h, chie ly on he p esence o absence o he ou e uppe
inciso (i 3 ), o one o bo h o he i s (pm 2 ) o second (pm 3 ) uppe p emola s, and on he p esence
o absence o he second (pm 3 ) o he lowe p emola s, as Ta e's 'phyle ic' diag ams (loc. ci .)
indica e. These den al ea u es ha e been discussed in mo e de ail abo e (p. 230): hey e lec
he deg ee o sho ening ha o ms an e iden end wi hin he sub amily. When combined wi h
he ela i e size o one o mo e o hese ee h and he posi ion o he ele an oo h o ee h in he
oo h ow such ac o s o m an impo an elemen in gene ic iden i ica ion and diagnosis (c .
Mille , loc. ci .). The many di e en combina ions o incisi e and p emola o mula in he sub-
amily (Table 2), he e anescence in some gene a o some o he ee h in ol ed, he ex eme
endencies owa ds educ ion seen in some such as Pipis ellus, and he a ie y o posi ions wi hin
he oo h ow adop ed by i 3 and pm 2 in pa icula ein o ce he conclusion ha such ea u es e lec
a uni e sal end ha may ha e occu ed mo e han once wi hin he g oup and which as a esul
may no p o ide a o ally eliable ya ds ick by which ela ionship may be judged.
In addi ion o hese den al ea u es, Ta e (1942a) e iewed a numbe o o he cha ac e s used in
he classi ica ion o he sub amily. These include he p esence o absence o accesso y canine cusps;
he o m and shape o he b aincase and os um; he deg ee o educ ion o he zygoma a; he
s uc u e o he pala e, i s an e io ema gina ion and accesso y an e io and pos e io spines; he
p esence o absence o basial pi s; enla gemen o he ea s and hei associa ed bony s uc u es;
he p esence o absence o adhesi e pads on he humb o oo ; and he na u e o o he mino
s uc u es such as he calca . These ea u es, howe e , seem o g ea e alue in he dis inc ion o
species and species g oups, ha is, o in agene ic classi ica ion, o o he diagnosis o indi idual
gene a.
The alue o such cha ac e s has been discussed a some leng h by Zima & Ho acek (1985)
who poin ed ou ha he e a e g ounds o hinking ha some o he adi ional mo phological
VESPERTILIONINE SYSTEMATICS 257
cha ac e s may no p o ide unequi ocally eliable c i e ia o he es ablishmen o a classi ica ion
based on p esumed phyle ic ela ionship, and ha hei axonomic signi icance may be limi ed.
They also ema ked ha such cha ac e s may e lec pa allelism o con e gence, o esul om
selec ion p essu e a he han ela ionship. These easons led hem o sugges ha he baculum
migh p o ide one o se e al al e na i e sou ces o eliable, axonomically use ul c i e ia based on
cha ac e s ha do no ha e a di ec adap i e signi icance.
The s uc u e o he baculum in he Vespe ilioninae sugges s some modi ica ions o ibal
classi ica ion wi hin he sub amily, al hough clea ly o he mo phological cha ac e s need o be
gi en equi alen o g ea e weigh . P o isionally, he e o e, we o e an a angemen o he Vespe -
ilioninae in which bacula mo phology is used in associa ion wi h he adi ional diagnos ic
ea u es o sugges possible ela ionship. This classi ica ion is p esen ed in Table 1 .
The e appea o be wo majo bacula ypes in he Vespe ilioninae, each wi h nume ous a ia-
ions as migh be expec ed in such a la ge and di e se sub amily. A classi ica ion ha includes a
majo conside a ion o bacula mo phology shows signi ican esemblances o ea lie a ange-
men s based on adi ional and con en ional mo phological ea u es. Howe e , he e a e some
wide di e gences, as o example he seemingly a i icial na u e o he 'Nyc iceini' o he associ-
a ions o he a ious gene a o big-ea ed ba s. Ta e (19420) commen ed upon he la e and
poin ed ou ha e y la ge ea s and hei associa ed audi o y specialisa ions in he skull occu ed
independen ly in h ee sec ions o he sub amily: indeed, i An ozous and Baue us a e included,
hese ea u es occu ou imes in he g oup. In pa icula , bo h Mille (1907) and Ta e (loc. ci .)
associa ed Laepho is wi h His io us on c anial and den al mo phology bu i s bacula s uc u e
shows a clea a ini y wi h Pipis ellus (Neo omicid) as he e ecognised. O onyc e is, ano he
big-ea ed ba , was allied by Ta e (loc. ci .) o he 'Nyc iceini' bu p o es o ha e a baculum much
mo e like hose o he pleco ine gene a.
One majo bacula ype is 'saddle-like' o 'slippe -like' and is exempli ied by Myo is and
Pizonyx. Thei bacula a e e y simila , emphasising he close ela ionship ha is gene ally accep ed
be ween hese gene a. The baculum o Lasionyc e is is somewha di e en in he p esence o a
leng hened sha . Howe e , in compa ison wi h he long-sha ed bacula ound in he Pipis ellini
he baculum o Lasionyc e is is ela i ely sho , and i e ains indica ions o he mo e cha ac e is ic
myo ine ype in i s up aised p oximal and dis al po ions. The occasional p esence o a la ened
do sal p ominence on i s base also ecalls he condi ion ound in Idionyc e is. The genus, al hough
ha ing some specialised ea u es, is allied i mly o Myo is by Mille (1907) and Ta e ( 942a). I has
sligh ly hooked uppe inciso s, i 3 wi h a sligh ly canini o m p o ile as in Myo is', pm 2 is in he line o
he oo h ow; m 3 is un educed; pm 2 _ 3 a e exac ly as in Myo is, much smalle han pm 4 , wi h pm 3
no emo ed om he line o he o he ee h. Al hough pm 3 has been los , his appea s o be a
specialisa ion; as Ta e (loc. ci .) poin ed ou , pm 2 _ 3 s ill ag ee closely wi h hose o he less
specialised species o Myo is no only in ela ion o each o he bu also in hei p opo ional size
ela i e o pm 4 . Al hough associa ed wi h Myo is, his genus is conside ably specialised in o he
ways (Mille , loc. ci .) and i s bacula s uc u e may well e lec his di e gence. I s baculum migh
be ega ded as de i ed om he mo e ypical myo ine s uc u e.
Bacula a iously eminiscen o he saddle shaped s uc u e ound in Myo is occu in a numbe
o o he gene a. Such bacula cha ac e ise Pleco us (including Co yho hinus), Idionyc e is,
Ba bas ella, Rhogeessa, Baeodon, Nyc iceius, O onyc e is, Lasiu us, Dasyp e us, An ozous and
Baue us, and possibly may be ound in Eude ma. Ta e (19420) pos ula ed he g ouping 'Pleco ini'
o Pleco us, (Co yno hinus), Idionyc e is and Eude ma, allying i o he Myo ini bu no employing
he e m in a o mal axonomic o sys ema ic sense. Bacula mo phology hus lends suppo o his
hypo hesis ha he pleco ine gene a should be associa ed wi h Myo is. Also, he baculum o
Ba bas ella sugges s ha i oo belongs he e: Mille (1907) pos ula ed such a ela ionship, despi e
se e al mo phological di e ences. Rhogeessa, Baeodon, Nyc iceius and O onyc e is also seem
allied o his g ouping. Ta e (19420) e e ed hese gene a o he 'Nyc iceini' wi h Sco oecus,
Sco omanes and Sco ophilus on accoun o hei incisi e and p emola den i ion. Howe e , he
bacula o Rhogeessa, Baeodon, Nyc iceius and O onyc e is a e a ian s o he saddle-like ype; ha
o Sco oecus is like ha o Pipis ellus (Pipis ellus), and he bacula o Sco omanes and Sco ophilus
a e b oadly simila o hose o Ep esicus and i s allies. Lasiu us, Dasyp e us, An ozous and Baue us
258 J. E. HILL & D. L. HARRISON
ha e u he a ian s o his bacula ype, bu a e qui e dis inc i e on o he mo phological
g ounds.
The bacula o An ozous and Baue us a e no a all like ha o O onyc e is, wi h which hese
gene a ha e been en a i ely associa ed (Pine e al., 1971), no do hei bacula ha e any signi ican
esemblance o hose o Nyc ophilus o Pha o is, hus suppo ing he iew (Koopman, 19846,
1985; B eed & Inns, 1985) ha hese No h Ame ican gene a should no be associa ed wi h
he Aus alian Nyc ophilus and Pha o is in he sub amily Nyc ophilinae. Bacula mo phology
sugges s ins ead an associa ion wi h hose gene a ha ha e he myo ine ype o baculum, o which
he bacula o An ozous and Baue us ha e many esemblances. The bacula o Nyc ophilus and
Pha o is (Fig. 22a-h) a e consis en ly homogeneous and di e in many ways om hose o he
gene a usually e e ed o he Vespe ilioninae. Fo he p esen we would place hese wo gene a in
a sepa a e sub amily, he Nyc ophilinae, a he han me ge hem in o he Vespe ilioninae as is
done by Koopman (1984a, 19846, 1985).
A u he basically iangula and la ened a ian o he saddle-like baculum cha ac e ises he
gene a Ep esicus, Vespe ilio (i he pseudobaculum is igno ed), la and His io us. Mille (1907)
ema ked ha he skull o Vespe ilio showed a s ong likeness o ha o Lasionyc e is bu ha
he o me was in all espec s a ypical Ep esicus. Vespe ilio and Lasionyc e is a e sepa a ed
by ma ked den al and bacula di e ences: he bacula mo phology o Vespe ilio allies i wi h
Ep esicus as Mille sugges ed. I is pe haps no un easonable o specula e ha Lasionyc e is which
has a s ongly myo ine den i ion has di e ged among he Myo ini in he same way as Vespe ilio
has di e ged among he Vespe ilionini, he la e genus suppo ing a long penis ei he by a
cen ally si ua ed baculum o pe haps mo e e ec i ely by he de elopmen o a ca ilaginous
pseudobaculum, his unc ion in Lasionyc e is by a sho sha . The gene a Tylonyc e is and
Mime illus also belong he e. The A ican Glauconyc e is has been associa ed (Ryan, 1966;
Koopman, 1971) wi h he Aus alian Chalinolobus bu hei bacula di e widely. Al hough s uc-
u ally a iable wi hin he genus, he bacula o Glauconyc e is a e mo e like he espe ilionine
o ep esicine ype: hose o Chalinolobus a e long-sha ed and like he bacula o Pipis ellus
(Pipis ellus). Finally, he baculum o Sco omanes appea s o be a de i a i e o he saddle-like ype,
leading o he dis inc i e baculum o Sco ophilus.
The genus Pipis ellus seems o s and mo e o less a he cen e o he second majo g ouping.
I has b oadly wo di isions in bacula e ms, one cha ac e ised by a long baculum wi h well
de eloped basal lobes and a ela i ely long, mos ly cylind ical sha , i s ip o en bi id o wi h
simila elabo a ion. The second di ision includes hose species in which he basal lobes a e some-
imes small o obsole e and which ha e a sho e , la e , en ally lu ed sha , i s ip some imes
elabo a ed in o a spa ula e o pla ele -like s uc u e.
These g oupings ha e been used in his s udy o suppo subgene ic di ision o his la ge genus.
The i s di ision includes Pipis ellus (Pipis ellus}, P. (Vespadelus), P. (Pe imyo is} and P.
(A ielulus}. Reduc ion and loss o pm 2 occu s in P. (A ielulus) and he oo h is almos in a i-
ably absen in P. (Vespadelus}. The second di ision con ains P. (Hypsugo) in which pm 2 may be
e y small o absen , P. (Neo omicia) om which i is again almos in a iably absen , and P.
(Falsis ellus}. Al hough p ima ily Old Wo ld in dis ibu ion, bo h o hese di isions a e ep e-
sen ed in he New Wo ld, each by a single species. The Aus alian P. ( Vespadelus} seems on bacula
ea u es o ep esen P. (Pipis ellus); he wholly A ican P. (Neo omicia) is appa en ly simila ly
ela ed o P. (Hypsugo), o which P. (Falsis ellus} appea s o be an eas e n ep esen a i e.
The majo i y o he gene a he e alloca ed o he Pipis ellini show s ong bacula a ini ies o
Pipis ellus (Pipis ellus): some such as Glisch opus and Sco ozous ha e been conside ed congene ic
wi h Pipis ellus in he pas . Besides Glisch opus and Sco ozous hese include Nyc iceinops,
Sco eanax, Sco o epens, Sco oecus, Nyc alus, Hespe op enus and Chalinolobus, all wi h long-
sha ed bacula. O he emainde , Laepho is in bacula s uc u e is simila o P. (Neo omicia}, while
Phile o has a baculum ha appea s o be an elabo a ion o he bacula s uc u e ound in some o
P. (Hypsugo). Ta e (1942a) pos ula ed a ela ionship be ween Phile o , Tylonyc e is and pe haps
Mime illus bu he bacula o he i s wo a e o ally dissimila and he s uc u e is appa en ly
absen om Mime illus: i is e y small in Tylonyc e is. Hill (1966a) d ew a en ion o he unusual
geni alia o Phile o and ollowing Ta e's (loc. ci .) sugges ion o a ini y wi h Pipis ellus jo ei and
VESPERTILIONINE SYSTEMATICS 259
i s associa es allied Phile o wi h his g oup. Un o una ely, excep ing o he abe an species
s enop e us he bacula o he s enop e us subg oup (including P.jo ei) o his p esen s udy emain
unknown.
Bacula mo phology sugges s ha he con en ional iew ha Ep esicus and i s immedia e allies
de i e om o a e closely ela ed o Pipis ellus can be ques ioned. C anially and den ally he e a e
many simila i ies be ween 'Ep esicus ' as o me ly denned and Pipis ellus and as Koopman (1975)
has poin ed ou , he loss o pm 2 enables a species o c oss he bounda y be ween he wo gene a as
hen unde s ood, a p ocess which in his iew migh ha e occu ed mo e han once. Ou conclusions
do no challenge his opinion: hose 'Ep esicus' species in which pm 2 has been ound occasionally
o occu p o e on bacula g ounds o be close o Pipis ellus han o Ep esicus as we unde s and i ,
while Pipis ellus as o me ly de ined has long been known o include some species om which on
occasion his 'diagnos ic' oo h is absen . Clea ly, ou indings suppo Koopman's (loc. ci .)
opinion ha his p ocess may ha e occu ed se e al imes and indeed may be occu ing in some
species, bu all belong o he one genus, Pipis ellus.
As we unde s and i s composi ion, Ep esicus is now a mo e es ic ed genus in which he i-
angula , la ened baculum is basically close in s uc u e o he saddle-like g ouping han o he
long-sha ed g oup, al hough some Ep esicus do indeed ha e bacula ha sugges he beginnings
o basal lobula ion o o a e y sho sha . We sugges he e o e ha in bacula e ms he
Vespe ilionini o which we e e Ep esicus may ep esen a ansi ional s age be ween he saddle-
like baculum and he p edominan ly basally lobed and long-sha ed ype. Tylonyc e is and
Glauconyc e is also show his endency.
Den al educ ion p oceeds h oughou bo h o he majo bacula g oups. In he g ouping wi h
b oadly myo ine o saddle-like bacula he den i ion a ies in numbe o ee h om a o al o 38
(Myo is, Pizonyx) h ough 36 (Lasionyc e is, Pleco us and allies), 34 (Ba bas ella, Ep esicus and
allies), 32 (Lasiu us), 30 (Dasyp e us, Rhogeessa, Baeodon, Nyc iceius, O onyc e is, Sco omanes,
Sco ophilus) o 28 (An ozous, Baue us}. In he second o he wo majo bacula g oups, den al
educ ion a ies om Eudiscopus wi h a o al o 36 ee h (i s associa ion he e is p esumed)
h ough 34 (Pipis ellus, Glisch opus, Sco ozous, Nyc alus, Chalinolobus), 32 (Laepho is, Phile o ,
Hespe op enus) o 30 (Nyc iceinops, Sco eanax, Sco o epens, Sco oecus). Thus his end occu s
concu en ly in he wo majo g oupings, aking he same o m in each by inc easing he size and
bulk o i 2 , he educ ion, ansposi ion and loss o i 3 , and he p og essi e educ ion, ansposi ion
and loss o pm 2 , pm 3 and pm 2 .
Zoogeog aphical conside a ions
The saddle-shaped o slippe -like baculum cha ac e is ic o he Myo ini, Pleco ini, Lasiu ini and
An ozoini as he e unde s ood is cosmopoli an in bu one genus, Myo is. I occu s in one Hola c ic
genus, Pleco us, in one Palaea c ic genus, Ba bas ella, i sel p obably closely ela ed o Pleco us,
and in one o he Old Wo ld genus, O onyc e is, ha occu s in sou hwes e n Asia and no he n
A ica. O he wise his bacula ype is limi ed o he New Wo ld. Lasionyc e is, exclusi ely No h
Ame ican, has a baculum appa en ly de i ed om his ype, as does Nyc iceius, also No h
Ame ican, al hough in his genus he baculum is conside ably modi ied o he ex en ha Hamil on
(1949) commen ed upon i s unique cha ac e among he gene a ha he had examined. Thus
al hough he saddle-shaped baculum o i s de i a i es is ep esen ed abou equally in numbe o
species in he Old and New Wo lds, gene a wi h bacula o his ype p edomina e in he la e , i s
ex ension in o he Old Wo ld being p ima ily h ough he many species o Myo is, wi h a lesse
con ibu ion om Pleco us, Ba bas ella, and O onyc e is.
A u he a ie y o his bacula ype is ound in he Vespe ilionini, ha is, in Ep esicus and i s
close ela i es. In hese, he baculum is less s ongly saddle-shaped o slippe -like, la e , and o en
mo e iangula in ou line. This bacula ype is p ima ily Old Wo ld in numbe s o gene a and
species, only Ep esicus among Old Wo ld gene a ex ending o he New Wo ld whe e he e is a
closely ela ed genus, His io us. In he Old Wo ld, Vespe ilio is also closely ela ed o Ep esicus.
Ano he Old Wo ld genus, la, is a gian ep esen a i e o his same bacula ype. The sou heas e n
260 J. E. HILL & D. L. HARRISON
Asian Tylonyc e is and he A ican Glauconyc e is ha e bacula ha a e modi ied a ian s o his
ype: Mime illus, in which no baculum has been ound, also appea s o belong he e. Two u he
Old Wo ld gene a, Sco omanes and Sco ophilus, also ha e bacula ha a e simila in many espec s
o he espe ilionine ype.
The sha ed o long-sha ed bacula ype is con ined almos exclusi ely o he Old Wo ld, and is
ep esen ed in he New Wo ld by no mo e han wo species o Pipis ellus in he Nea c ic egion,
one o hese wi h a highly modi ied baculum. This bacula ype is es ic ed o he Pipis ellini and
wi hin ha g ouping o hose gene a ha o he mos pa can be shown on o he g ounds o
clus e a ound Pipis ellus. Indeed, some such as Sco ozous, Glisch opus, Sco eanax, Sco o epens
and pe haps e en Nyc alus migh on bacula g ounds be ega ded as subgene a o his widesp ead
genus. In a educed o m his bacula ype appea s in wo o he subgene a o Pipis ellus, P.
(Pe imyo is) and P. (A ielulus). Widesp ead in he Palaea c ic egion and in sou heas e n Asia,
his bacula ype is ep esen ed in Aus alia by i e dis inc g oupings: Pipis ellus (Pipis ellus},
P. ( Vespadelus), Sco eanax, Sco o epens, and Chalinolobus. This ype o baculum also occu s in
A ica among Pipis ellus kuhlii and i s associa es, which migh in ac be conside ed o wa an
ecogni ion as a u he subgenus o Pipis ellus.
A u he a ian o he sha ed bacula ype is ound in Pipis ellus (Hypsugo) and P. (Falsis el-
lus). In hese he sha is sho e and is en ally lu ed, o en wi h expansion o he ip. Pipis ellus
(Hypsugo) is con ined chie ly o Asia and A ica, whe e in he la e egion i appea s o be closely
associa ed wi h P. (Neo omicia) in which pm 2 is gene ally los . Thus as in Aus alia whe e P.
(Vespadelus) in which pm 2 is also gene ally absen appea s o de i e om P. (Pipis ellus), so in
A ica P. (Neo omicia) is appa en ly simila ly ela ed o P. (Hypsugo). O he wo No h Ame ican
pipis elles, P. sub la us has a educed o m o he P. (Pipis ellus) baculum, he sha e y sho
and s ubby: his species has a myo ine agus and has been conside ed (Menu, 1984) o ha e a
myo ine den i ion. Howe e , on he balance o ea u es i appea s o be clea ly e e able o
Pipis ellus and indeed o be c anially and den ally close o P. (Pipis ellus), which appa en ly i
ep esen s in No h Ame ica. The e do no appea o be su icien g ounds o jus i y i s gene ic
sepa a ion om Pipis ellus as has been ecen ly e ec ed (Menu, loc. ci .), al hough subgene ic
ecogni ion wi hin ha genus seems app op ia e. The second No h Ame ican species o Pipis el-
lus, P. Hespe us, should e iden ly be e e ed o P. (Hypsugo) wi h which i has close bacula and
den al simila i ies, al hough ecen ly gene ic sepa a ion (Ho acek & Hanak, 19850, b, 1985-1986)
has been p oposed o i . Finally, P. (Falsis ellus) is es ic ed o sou heas e n Asia, Aus alasia
and Tasmania: he deeply en ally lu ed baculum o his subgenus, lacking basal and dis al
modi ica ion bu massi e and subs an ial appea s o be an ex eme o he P. (Hypsugo) ype:
possibly P. (Falsis ellus) ep esen s P. (Hypsugo) which seems o be linked o i by se e al o i s
Asian species.
One co olla y o he emo al o he A ican capensis and enuipinnis g oups o 'Ep esicus' o
Pipis ellus, and o he simila ans e o he Aus alian species o me ly e e ed o 'Ep esicus' is
ha in he Old Wo ld Ep esicus now becomes p ima ily Palaea c ic, wi h ou lie s, pe haps all
closely connec ed o E. se o inus, in A ica while in he New Wo ld i ex ends o e bo h No h and
Sou h Ame ica. In sou heas e n Asia he genus becomes es ic ed o no u he eas han sou he n
Thailand, he o me eno mous hia us in i s dis ibu ion be ween his pa o sou he n Asia and
Aus alia ha ing been emo ed.
Conclusions
(1) The cu en classi ica ion o he Vespe ilioninae is based chie ly on adap i e cha ac e s wi h
conside able emphasis on acial sho ening and concomi an den al educ ion and loss. Se e al
au ho s ha e d awn a en ion o he de iciencies and dange s o any classi ica ion ha elies hea ily
on such ea u es. A e iew o bacula mo phology wi hin he sub amily sugges s ha his s uc u e
p o ides indica ions o ela ionship ha in many espec s suppo he exis ing classi ica ion bu
which also indica e se e al changes o he cu en a angemen . In pa icula , bacula mo phology
sugges s a numbe o majo and mino changes in he sys ema ics o he nominal gene a Pipis ellus
VESPERTILIONINE SYSTEMATICS 261
and Ep esicus, sepa a ed hi he o only by den al o mula, i sel subjec o a ia ion in bo h 'gene a'
as hey a e cu en ly unde s ood.
(2) The p esence o absence o he an e io uppe p emola (pm 2 ) in Pipis ellus and Ep esicus,
used o me ly as hei p incipal diagnos ic cha ac e , has li le axonomic signi icance. The oo h is
a iable in Pipis ellus as he e unde s ood, being educed o los in h ee o i s subgene a, and is
absen om Ep esicus as we en isage i .
(3) Bacula mo phology in Pipis ellus and Ep esicus p o ides g oupings ha la gely ag ee in
species con en wi h hose p oposed by ea lie au ho s such as Ta e (19420) and Koopman (1973,
1975) al hough in basing hei s udies on 'con en ional' mo phological cha ac e s nei he con-
side ed hese gene a in hei en i e y. The bacula mo phology o 'Ep esicus' as i is cu en ly
unde s ood p o ides a clea indica ion ha as such i is no a na u al g oup, bu ha h ee species
agg ega ions, he Aus alian pumilus g oup and he A ican capensis and enuipinnis g oups,
should be ans e ed o Pipis ellus.
(4) I has been possible o ecognise and de ine subgene a o he majo species g oups in bo h
Pipis ellus and Ep esicus and o sugges possible ela ionships be ween hem. One subgenus is
desc ibed as new as Pipis ellus (A ielulus) o P. ci cumda us and i s allies.
(5) The e appea o be clea links be ween ce ain o he pipis elline subgene a: Pipis ellus
( Vespadelus} in Aus alia seems o ep esen P. (Pipis ellus) in bacula e ms while P. (Hypsugo) is
appa en ly ep esen ed in Indo-Aus alia by P. (Falsis ellus) and is ela ed o he A ican P.
(Neo omicid). Al hough he ea u es o he wo Nea c ic species o Pipis ellus ha e been hough
o jus i y hei ecogni ion in sepa a e, indi idual gene a we conside ha he cha ac e s o one
(sub la us) me i no mo e han subgene ic s a us as he sole species o P. (Pe imyo is), which i sel
pe haps ep esen s P. (Pipis ellus), while he o he (Hespe us) is pe haps mo e app op ia ely
e e ed o P. (Hypsugo).
(6) The examina ion o bacula in Pipis ellus has sugges ed ha some axa hi he o anked as
subspecies, o example ab amus, pa e culus o helios, migh in ac be dis inc species.
(7) As we now unde s and he species con en o Pipis ellus and Ep esicus he o me emains
p ima ily an Old Wo ld genus whe e i is widesp ead and di e se in he opics and sub opics,
ex ending in o he empe a e zones and jus o No h Ame ica. In con as , ou concep o
Ep esicus limi s his genus o he New Wo ld and in he Old Wo ld p ima ily o he Palaea c ic,
wi h ou lying ep esen a i es in A ica. I does no ex end signi ican ly in o Aus alasia.
(8) Bacula mo phology sugges s he in o mal ecogni ion o wo majo g oupings wi hin he
sub amily Vespe ilioninae. The i s includes he Myo ini, Pleco ini and Lasiu ini; An ozous and
Baeu us, which in bacula e ms ha e no ela ion o Nyc ophilus and Pha o is ( he Nyc ophilinae);
he Sco ophilini o include Sco omanes and Sco ophilus; and inally he Vespe ilionini, he e
educed in con en o include Ep esicus and i s close ela i es His io us, la and Vespe ilio, wi h
Tylonyc e is, Mime illus and Glauconyc e is.
(9) The second g ouping consis s o Pipis ellus and hose gene a which clus e ound i . All wi h
he possible excep ion o Phile o appea o ela e qui e closely in bacula e ms o one o o he o
he subgene a ha we ecognise in Pipis ellus, p incipally o P. (Pipis ellus). Laepho is, o me ly
conside ed ela ed o His io us, is ins ead in bacula e ms closely associa ed wi h P. (Neo omicid).
The bacula o Chalinolobus and Glauconyc e is a e widely dissimila a hough hese gene a ha e
been closely allied in he pas ; Chalinolobus is o he pipis elline ype while he baculum o
Glauconyc e is appa en ly associa es i mo e app op ia ely wi h Ep esicus and i s allies.
(10) Bacula mo phology p o ides clea indica ions ha he 'Nyc iceini' o Ta e ( 942a) and
Koopman (1984, 1985) is no a na u al g oup, i s cons i uen membe s despi e c anial and den al
simila i ies ha ing widely di e en bacula. Thus Rhogeessa, Baeodon, Nyc iceius sensu s ic o, and
O onyc e is ha e been he e allied o he pleco ine ba s on bacula g ounds, while Sco eanax,
Sco o epens and Sco oecus a e qui e clea ly associa es in bacula e ms o Pipis ellus. 'Nyc iceius',
a one ime hough o include he Aus alian Sco eanax and Sco o epens as well as i s No h
262 J. E. HILL & D. L. HARRISON
Ame ican ype species hume alis and he A ican schlie enii, has ecen ly been es ic ed only o he
Ame ican and A ican o ms. These p o e o ha e widely di e en bacula; hume alis has been
associa ed wi h he pleco ine ba s on his accoun , while gene ic s a us has been acco ded o
schlie enii wi h he p oposal o a new gene ic name, Nyc iceinops.
(11) The wo b oad bacula ypes ha we disce n in he sub amily Vespe ilioninae ha e de ini e
geog aphical pa e ns: he saddle-like baculum and i s a ian s ha cha ac e ise he i s g oup
no ed abo e is p ima ily New Wo ld and Palaea c ic, ex ending less signi ican ly in o he Old
Wo ld opics o Aus alasia, while he sha ed baculum o he second g oup is chie ly con ined o
he Old Wo ld.
Addendum
A phene ic analysis o he ela ionships o selec ed espe ilionine species (chie ly hose cu en ly
e e ed o Pipis ellus and Ep esicus) by Ho acek & Hanak (1985-1986) appea ed while his pape
was in p ess. These au ho s p o ided de ini ions o Pipis ellus, Hypsugo (which hey conside ed
gene ically alid) and Ep esicus, based on he mo phology o he penis and baculum, he uppe
mola s, he basisphenoid pi s, he pel ic gi dle, and he ibia, ail and epiblema.
Ho acek & Hanak sugges ed ha he classi ica ion o pipis elloid ba s migh be cla i ied by he
ecogni ion o addi ional subgene a o gene a o hose species o species g oups ha do no
con o m p ecisely wi h hose ha hey included wi hin hese h ee gene ic g oupings. To some
ex en such ecogni ion is p o ided in se e al ins ances by he classi ica ion he e p oposed and
al hough some majo di e ences exis be ween he in o mal assessmen s and species g oups o
Ho acek & Hanak and he o mal a angemen pu o wa d in his pape he e is ne e heless
a b oad measu e o ag eemen . Ho acek & Hanak did no a emp any classi ica ion o he
Vespe ilioninae as a whole, bu 'Nyc iceius' schlie enii, he e conside ed o ep esen a dis inc
monospeci ic genus (Nyc iceinops gen. no .) was hough by hese au ho s o be e e able ei he o
Ep esicus ( Rhynep esicus), o possibly o jus i y he es ablishmen o a new subgenus wi hin
Ep esicus.
Re e ences
Acloque, A. 1899. Faune de F ance, con enan la desc ip ion des especes indigenes disposees en ableaux
analy iques e illus e de igu es ep esen an les ypes ca ac e is iques des gen es. I. Mammi e es, Oiseaux,
Poissons, Rep iles, Ba aciens, P o ocho des. Pa is: Lib ai ie J.-B. Ballie e e Fils.
Aellen, V. 1952. Con ibu ion a 1'e ude des Chi op e es du Came oun. Memoi es de la Socie e Neucha eloise
des Sciences Na u elles, Neucha el 8: 1-121.
- 1957. Les chi op e es a icains du Musee Zoologique de S asbou g. Re ue Suisse de Zoologie, Gene e
64:189-214.
1959. Chi op e es nou eaux d'A ique. A chi es des Sciences, Gene e, 12: 217-235.
Ag awal, V. C. & Sinha, Y. P. 1973. S udies on he bacula o some O ien al ba s. Ana omische Anzeige , Jena
133: 180- 192, 4 igs.
Allen, G. M. 1908. No es on Chi op e a. Bulle in o he Museum o Compa a i e Zoology, Ha a d 52: 25-62.
- 1911. Ba s om B i ish Eas A ica. Bulle in o he Museum o Compa a i e Zoology, Ha a d 54:
321-331.
- 1914. Mammals om he Blue Nile Valley. Bulle in o he Museum o Compa a i e Zoology, Ha a d 58:
305-357. 3 igs
1939. A checklis o A ican mammals. Bulle in o he Museum o Compa a i e Zoology, Ha a d 83:
1-763.
Allen, H. 1891. Change o name o a genus o ba s. P oceedings o he Philadelphia Academy o Na u al
Sciences 43: 466.
Ande son, K. 1912. Ca alogue o he Chi op e a in he collec ion o he B i ish Museum. 2nd ed. Vol. I:
Megachi op e a. London: T us ees o B i ish Museum.
A allah, S. I. 1970. Ba s o he genus Myo is (Family Vespe ilionidae) om Lebanon. Occasional Pape s
Uni e si y o Connec icu , S o s, (Biological Sciences Se ies) 1: 205-212, 2 igs, 1 ab.
VESPERTILIONINE SYSTEMATICS 263
Baagee, H. J. 1973. Taxonomy o wo sibling species o ba s in Scandina ia, Myo is mys acinus and Myo is
b and i (Chi op e a). Videnskabelige Meddelelse a Dansk Na u his o isk Fo ening i Kjobenha n 136:
191-216, 17 igs, 2 abs.
Bha naga , K. P. 1967. Bacula o some Indian Megachi op e a. Jou nal o Mammalogy, Bal imo e 48:
494-497, 3 i gs .
Bianchi, V. 1917. No es p elimina ies su les chau e-sou is ou Chi op e es de la Russie. Annuai e du Musee
Zoologique de I'Academie Impe iale des Sciences, S Pe e sbu g (Lening ad) (1916) 21: LXXIII-LXXXII.
(In Russian).
Blain ille, H. M. D. de 1840. Os eog aphie ou desc ip ion iconog aphique compa ee du squele e e du sys eme
den ai e des Mammi e es, ecen s e ossiles pou se i de base a la zoologie e a la geologie. Pa is: J. B.
Ballie e e Fils.
Bly h, E. 1840. The Mammalia, Bi ds and Rep iles. In Cu ie , G. [L.C.F.D.], The Animal Kingdom, a anged
a e i s o ganisa ion, o ming a na u al his o y o animals, and an in oduc ion o compa a i e ana omy.
London: W. S. O & Co.
Bocage, J. V. B. du 1889. Chi op e es a icains nou eaux, a es ou peu connus. Jo nal de Sciencias
Ma hema icas, Physicas e Na u aes, Lisboa (2), 1: 1-7, 2 igs.
Bonapa e, C. L. J. L. 1832-1841. Iconog aphia della Fauna I alica pe le qua o classi degli Animali
Ve eb a i. Roma: Tipog a ia Sal iucci.
B aes up, F. W. 1935. Repo on mammals collec ed by M Ha y Madsen du ing P o esso O. Olu sen's
Expedi ion o F ench Sudan and Nige ia in he yea s 1927-28. Vidensk Meddelelse a Dansk Na u -
his o isk Fo ening i Kjobenha n 99: 73-130.
B eed, W. G. & Inns, R. W. 1985. Va ia ion in spe m mo phology o Aus alian Vespe ilionidae and i s
possible phylogene ic signi icance. Mammalia, Pa is 49: 105-108, 2 pis.
B own, R. E. 1967. Bacula o some New Wo ld molossid ba s. Mammalia, Pa is 31: 645-667, 51 igs.
, Genoways, H. H. & Jones, J. K. J . 1971. Bacula o some Neo opical ba s. Mammalia, Pa is 35:
456-464, 1 ig.
Bu , W. H. 1960. Bacula o No h Ame ican mammals. Miscellaneous Publica ions, Museum o Zoology,
Uni e si y o Michigan, Ann A bo No. 113: 1-75, 25 pis.
Chaine, J. 1926. L'os penien, e ude desc ip i e e compa a i e. Ac es de la Socie e Linneenne de Bo deaux 78:
5-195, 133 igs.
Chasen, F. N. 1940. A Handlis o Malaysian mammals. Bulle in o he Ra les Museum, Singapo e No. 15;
i-xx, 1-209, map.
Chu chill, S. K., Hall, L. S. & Helman, P. M. 1984. Obse a ions on long-ea ed ba s (Vespe ilionidae:
Nyc ophilus) om no he n Aus alia. Aus alian Mammalogy 7: 17-28, 6 igs, 3 abs.
Co be , G. B. 1964. The g ey long-ea ed ba Pleco us aus iacus in England and he Channel Islands.
P oceedings o he Zoological Socie y o London 143: 51 1-515, 2 abs.
& Hill, J. E. 1980. A wo ld lis o mammalian species. 1s ed. London/I haca: B i ish Museum (Na u al
His o y)/Coms ock Publishing Associa es, Co nell Uni e si y P ess.
1986. A wo ld lis o mammalian species. 2nd ed. New Yo k/London: Fac s on File/B i ish Museum
(Na u al His o y).
Dauben on, E. L. 1 760. His oi e na u elle gene a e e pa iculie e a ec la desc ip ion du Cabine du Roi. Tome 8.
Pa is: Imp ime ie Royale.
Da is, D. D. 1947. The bacula o some ui ba s (P e opus). Fieldiana: Zoology, Chicago 31: 125-131, 2 igs.
Da is, W. H. & Rippy, C. L. 1968. Dis ibu ion o Myo is luci ugus and Myo is aus o ipa ius in he sou h-
eas e n Uni ed S a es. Jou nal o Mammalogy, Bal imo e 49: 1 13-1 17, 2 igs.
De Blase, A. F. 1980. The ba s o I an: sys ema ics, dis ibu ion, ecology. Fieldiana: Zoology, Chicago N.S.
No. 4 (Pub. 1307): i-x ii, 1^24, 180 igs.
Dobson, G. E. 1 87 1 . No es on nine new species o Indian and Indo-Chinese Vespe ilionidae, wi h ema ks on
he synonymy and classi ica ion o some o he species o he same amily. P oceedings o he Asia ic Socie y
o Bengal, Calcu a 210-215.
- 1876. Monog aph o he Asia ic Chi op e a and Ca alogue o he species o ba s in he collec ion o he
Indian Museum, Calcu a. London: T us ees o Indian Museum.
1878. Ca alogue o he Chi op e a in he collec ion o he B i ish Museum. London: T us ees o B i ish
Museum.
I He man, J. R. & Mo ison-Sco , T. C. S. 1951. Checklis o Palaea c ic and Indian mammals 1758-1946.
London: T us ees o B i ish Museum (Na u al His o y).
, & I layman, R. W. 1953. Sou he n A ican mammals 1758-1951: a ^classi ica ion. London:
T us ees o B i ish Museum (Na u al His o y).
264 J. E. HILL & D. L. HARRISON
E colani, G. B. 1868. Dei Tessu i e degli o gan! e e ili. Memo ie dell 'Accademia delle Scienze dell'Ins i u o di
Bologna (2), 8: 281-362, 10 pis.
Fai on, J. 1980. Deux nou elles especes de Chei op e es pou la auna du Massi de 1'Ai (Nige ); O onyc e is
hemp ichi Pe e s, 1 859 e Pipis ellus nanus (Pe e s 1 852). Bulle in de I'lns i u Royal des Sciences Na u elles
de Belgique, B uxelles 52 (17): 1-7, 2 igs, 2 pho os, 1 ab.
Fische , J. B. 1829. Synopsis mammalium. (Wi h addenda e emendanda, index, co igenda). S u ga diae.
Fi zinge , L. J. 1870. K i ische Du chsich de O dnung de Fla e hie e ode Hand liigle (Chi op e a).
Familie de Flede mause (Vespe iliones). V. Ab heilung. Si zungbe ich e de [ Kaise lichen J Akademie de
Wissenscha en. Ma hema isch-Na u wissenscha liche Classe, Wien 62: 353-438.
Gaisle , J., Madkou , G. & Pelikan,. J. 1972. On he ba s o Egyp . P i odo edne p dce iis a u Ceskoslo enske
Akademie Ved B ne (Ac a Scien ia um Na u alium Akademiae Scien a ium Bohemoslo aceae B no, N.S. 6
(8): 1-40, 5 igs, 4 pis, 20 abs.
Genoways, H. H. & Jones, J. K., J . 1969. Taxonomic s a us o ce ain long-ea ed ba s (genus Myo is) om
he sou hwes e n Uni ed S a es and Mexico. The Sou hwes e n Na u alis , Dallas 14: 1-13, 5 igs, 1 ab.
Ge ha d , U. 1905. Mo phologische und biologische S udien iibe die Kopula ionso gane de Sauge ie e.
Jenaische Zei sch i u Na u wissenscha , Jena 39: 43-1 18, 1 pi.
Gilbe , T. 1892. Das Os p iapi de Sauge hie e. Mo phologisches Jah buch, Leipzig 18: 805-831, 1 pi.
G ay, J. E. 1838. A e ision o he gene a o ba s (Vespe ilionidae) and he desc ip ion o some new gene a
and species. Magazine o Zoology and Bo any, Edinbu gh & Dublin 2: 483-505.
- 1866. Synopsis o he gene a o Vespe ilionidae and Noc ilionidae. Annals and Magazine o Na u al
His o y, including Zoology, Bo any and Geology, London (3), 17: 89-93.
Hamil on, W. J. 1949. The bacula o some No h Ame ican espe ilionid ba s. Jou nal o Mammalogy,
Bal imo e 30: 97-1 02, 1 pi.
Hanak, V. 1965. Zu sys ema ik de Ba lede maus Myo is mys acinus Kuhl, 18 19 und iibe das Vo kommen
on Myo is ikonniko i Ogne , 1912 in Eu opa. Ves nik Ceskoslo enske Spolecnos i Zoologicke, P aha 29:
353-367, 9 igs, 3 abs.
1970. No es on he dis ibu ion and sys ema ics o Myo is mys acinus Kuhl, 1819. In P oceedings o he
Second In e na ional Ba Resea ch Con e ence, Ams e dam, 1970. Bijd agen o de Die kunde , An s e dam
& Leiden 40 (1): 40-44, 8 igs, 1 ab.
1971. Myo is b and ii (E e smann, 1845) (Vespe ilionidae, Chi op e a) in de Tschechoslo akei.
Ves nik Ceskoslo enske Spolecnos i Zoologicke, P aha 35: 175-185, 7 igs, 1 ab.
Ha ison, D. L. 1960. A new species o pipis elle ba (Chi op e a: Pipis ellus) om sou h Is ael. Du ban
Museum No i a es 5(19: 261-267, 2 igs, 2 pis.
- 1982. Obse a ions on some a e A abian Pipis ellus (Chi op e a: Vespe ilionidae) wi h special
e e ence o he ex e nal male geni alia. Banne Zoologische Bei dge 33: 187-190, 2 igs.
& B ownlow, I. P. 1978. A compa a i e s udy o he baculum in ba s o he genus Sco ophilus
(Chi op e a: Vespe ilionidae) Mammalia, Pa is 42: 123-130, 7 igs.
H ay man. R. W. & Hill, J. E. 1 97 1 . 2. Chi op e a. In Mees e , W. & Se ze , H. W. [Eds] The mammals o A ica.
An iden i ica ion manual. Washing on: Smi hsonian P ess.
Helle , K-G. & Volle h, M. 1984. Taxonomic posi ion o 'Pipis ellus socie a is' Hill, 1972 and he ka yo-
logical cha ac e is ics o he genus Ep esicus (Chi op e a: Vespe ilionidae). Zei sch i u Zoologische
Sys ema ik und E olu ions o schung, F ank u am Main 22 (1): 65-77, 5 igs, 1 ab.
Heuglin, M. T. on 1877. Reise in No dos -A ica. Schilde ungen aus dem Gebie e de Beni Ame undHabab,
nebs zoologischen Skizzen undeinem Fuh e u Jagd eisende. B aunschweig: Geo ge Wes e mann.
Hill, J. E. 1966a. A e iew o he genus Phile o (Chi op e a: Vespe ilionidae). Bulle in o he B i ish Museum
(Na u al His o y) , (Zoology), London 14: 375-387, 3 igs.
- 19666. The s a us o Pipis ellus egulus Thomas (Chi op e a: Vespe ilionidae) Mammalia, Pa is 30:
302-307.
- 1969. The gene ic s a us o Glisch opus osse i Oey, 1951 (Chi op e a: Vespe ilionidae). Mammalia,
Pa is 33: 133-1 39.
- 1971. The s a us o Vespe ilio b achyp e us Temminck, 1840 (Chi op e a: Vespe ilionidae).
Zoologische Mededelingen, Leiden 45: 139-146.
1972. The Gunong Benom Expedi ion 1967. 4. New eco ds o Malayan ba s, wi h axonomic no es and
he desc ip ion o a new Pipis ellus. Bulle in o he B i ish Museum (Na u al His o y), (Zoology), London
23: 2 1-42, 3 abs.
1974. A e iew o Sco oecus Thomas, 1901 (Chi op e a: Vespe ilionidae). Bulle in o he B i ish Museum
(Na u al His o y), (Zoology), London 27: 167-188, 4 igs, 1 ab.
1976. Ba s e e ed o Hespe op enus Pe e s, 1869 (Chi op e a: Vespe ilionidae) wi h he desc ip ion o
a new subgenus. Bulle in o he B i ish Museum (Na u al His o y) , (Zoology), London 30: 1-28, 5 igs, 4 pis.
VESPERTILIONINE SYSTEMATICS 265
- 1983. Ba s (Mammalia: Chi op e a) om Indo-Aus alia. Bulle in o he B i ish Museum (Na u al
His o y), (Zoology), London 45: 103-208, 14 abs.
& F ancis, C. M. 1984. New ba s (Mammalia: Chi op e a) and new eco ds o ba s om Bo neo and
Malaya. Bulle in o he B i ish Museum (Na u al His o y), (Zoology), London 47: 305-329, 2 igs, 4 abs.
& Topal, G. 1973. The a ini ies o Pipis ellus idleyi Thomas, 1898 and Glisch opus osse i Oey, 1951
(Chi op e a: Vespe ilionidae). Bulle in o he B i ish Museum (Na u al His o y), (Zoology), London 24:
447^56.
Hollis e , N. 1918. Eas A ican mammals in he Uni ed S a es Na ional Museum. Pa I. Insec i o a,
Chi op e a and Ca ni o a. Bulle in o he Uni ed S a es Na ional Museum, Washing on No. 99: 1-194, 3
igs, 55 pis.
Honacki, J. H., Kinn an, K. E. & Koeppl, J. W. [Eds] 1982. Mammal species o he wo ld: a axonomic and
geog aphic e e ence. Law ence, Kansas: Allen P ess/Associa ion o Sys ema ic Collec ions.
Ho acek, I. & Hanak, V. 1985a. Gene ic s a us o Pipis ellus sa ii (Bonapa e, 1837) and ema ks on
sys ema ics o he genus Pipis ellus. Abs ac s Se en h In e na ional Ba Resea ch Con e ence Thi d
Eu opean Ba Resea ch Symposium. Join Mee ing. Abe deen: Uni e si y o Abe deen. [Unpagina ed].
[Augus ].
, - 19856. Gene ic s a us o Pipis ellus sa ii (Bonapa e, 1837) and ema ks on sys ema ics o he
genus Pipis ellus. Ba Resea ch News, Po sdam, New Yo k 26: 62. [No embe ].
1985-1986. Gene ic s a us o Pipis ellus sa ii and commen s on classi ica ion o he genus
Pipis ellus (Chi op e a, Vespe ilionidae). Myo is, Bonn 23-24: 1 1-16, 4 igs.
Ibanez, C. & Fe nandez, R. 1986. Sys ema ic s a us o he long-ea ed ba Pleco us ene i ae Ba e -Hamil on,
1907 (Chi op e a; Vespe ilionidae). Sduge ie kundliche Mi eilungen, Miinchen (1985), 32: 143-149, 2 igs,
2 abs.
& Val e de, J. A. 1985. Taxonomic s a us o Ep esicus pla yops (Thomas, 1901) (Chi op e a, Vespe i-
lionidae). Zei sch i u Sduge ie kunde, Hambu g & Be lin 50: 241-242.
Imaizumi, Y. 1959. A new ba o he " Pipis ellus ja anicus" g oup om Japan. Bulle in o he Na ional Science
Museum, Tokyo No. 45, (N.S. 4): 363-371, 6 igs, 1 pi, 3 abs.
I edale, T. & T ough on, E. le G. 1934. A check-lis o he mammals eco ded om Aus alia. Memoi o he
Aus alian Museum, Sydney No. 6: i-xi, 1-122.
Kaup, J. J. 1829. Skizzi e En wicklungs-Geschich e und na ii liches Sys em de Eu opdischen Thei wel . E s e
Theil, welche die Vogelsauge hie e und Vogel, nebs Andeu ung de En s ehung de le z e en aus Amphibien
en hdl . Da ms ad & Leipzig: Commision die Ca l Wilhelm Les e.
Khaju ia, H. 1979. S udies on he ba s (Chi op e a: Mammalia) o M.P., India. P . I. (Families P e opidae,
Rhinopoma idae and Embalonu idae [sic]). (Taxonomical and ecological s udies on ba s o Jabalpu
Dis ic Madhya P adesh, India. Pa I. Families P e opidae, Rhinopoma idae and Embalonu idae [sic]).
Reco ds o he Zoological Su ey o India Miscellaneous Publica ions Occasional Pape s, Delhi No. 13: 1-59,
6 igs, 7 pis, 8 abs.
- 1980. Taxonomical and ecological s udies on ba s o Jabalpu Dis ic Madhya P adesh, India. P . II.
(Families Megade ma idae, Rhinolophidae and Vespe ilionidae). Reco ds o he Zoological Su ey o
India Miscellaneous Publica ions Occasional Pape s, Delhi No. 19: 1-73, 8 pis, 8 abs.
1982. Ex e nal geni alia and bacula o some cen al Indian Mic ochi op e a. Sduge ie kundliche
Mi eilungen, S u ga 30: 287-295, 5 igs.
Ki chene , D. J. 1976. Ep esicus douglasi, a new espe ilionid ba om Kimbe ley, Wes e n Aus alia.
Reco ds o he Wes e n Aus alian Museum, Pe h 4: 295-301, 1 ig., 2 abs.
& Capu i, N. 1985. Sys ema ic e ision o Aus alian Sco eanax and Sco o epens (Chi op e a: Vespe i-
lionidae) wi h ema ks on ela ionships o o he Nyc iceiini. Reco ds o he Wes e n Aus alian Museum,
Pe h 12: 85-146, 19 igs, 3 abs.
& Jones, B. 1986. Re ision o Aus alo-Papuan Pipis ellus and Falsis ellus (Mic ochi op e a:
Vespe ilionidae). Reco ds o he Wes e n Aus alian Museum, Pe h, 12: 435-495, 20 igs, 3 abs, appendix.
Kolena i, F. A. 1856. Eu opa's Chi op e n. 1. Synopsis de Eu opaischen Chi op e a. Allgemeine deu sche
Na u his o ische Zei ung (New Folge), D esden & Leipzig 2: 121-133.
[F. A.] 1858. Eine neue os e eichische Flede maus. Si zungbe ich e de [Kaise lichen] Akademie de
Wissenscha en. Ma hema isch-Na u wissenscha lichen Classe, Wien 29: 250^256.
Koopman, K. F. 1965. S a us o o ms desc ibed o eco ded by J. A. Allen in The Ame ican Museum Congo
Expedi ion Collec ion o Ba s'. Ame ican Museum No i a es, New Yo k No. 2219: 1-34.
- 1971. Taxonomic no es on Chalinolobus and Glauconyc e is (Chi op e a, Vespe ilionidae). Ame ican
Museum No i a es, New Yo k No. 2451: 1-10, 1 ig.
1973. Sys ema ics o Indo-Aus alian pipis elles. Pe iodicum Biologo um, Zag eb 75: 1 13-1 16, 3 igs.
272 J. E. HILL & D. L. HARRISON
BM(NH) 27.12.1.37 Ta n Dao, Tonkin, Vie nam, 3000 (No. 41 1 o Thomas, 1928a).
BM(NH) 28.7. 1 .20 Phu Qui, Annam, Vie nam, 100 (No. 866 o Thomas, 19286, who iden i ied he specimen
as P. co omand a ama us, bu wi h a longe baculum han hose p e iously examined).
BM(NH) 83.76 Silau Silau T ail, Moun Kinabalu, Sabah, Bo neo. (Fig. lOe)
Pipis ellus pa e culus
BM(NH) 14.7.8.62 Pyaunggaung, N Shan S a es, Bu ma, 2794 .
BM(NH) 14.7.19.241 Kyauk Myaung, I awaddy, W Bu ma.
BM(NH) 14.7.19.242 Moun Popa, Uppe Bu ma (Holo ype). (Fig. 3c)
BM(NH) 14.7.19.240 Mandalay, Bu ma.
Pipis ellus angula us
BM(NH) 67.2125 Schoolmas e 's House, Nuhu, Guadalcanal I, Solomon Is (poncele i). (Fig. 4d)
Pipis ellus collinus
BM(NH) 50.983 Baiyanka, Pu a i-Ramu Di ide, SE Bisma ck Range, Papua New Guinea. (Fig. 4b)
Pipis ellus co omand a
BM(NH) 32. 1 1 . 1 .7 Nam Tamai, Uppe Bu ma.
BM(NH) 50.478 Ningma, Uppe Bu ma.
BM(NH) 76.1263 Sumka Uma, Uppe Bu ma.
HZM 1.7317, HZM 2.73 18 Nea Mi zapu , India. (Fig. 7c, HZM 2.7318)
HZM 4.7320 Dala pu , nea Mi zapu , India.
BM(NH) 4.6.8.1 Annam, Vie nam ( ama us). (Fig. 7b)
BM(NH) 27.12.1.40 Bac-kan, Tonkin, Vie nam ( ama us) (O iginal No. 444, seen by Thomas, 1928a).
Pipis ellus mimus
BM(NH) 98.5.5.20 Dangs, Bombay, India.
HZM 1 .10456 Vikas Vidyalaya, nea Ranchi, Biha , India. (Fig. 7g)
Pipis ellus mu ayi
BM(NH) 99.8.6.34 Ch is mas I, Indian Ocean (Holo ype). (Fig. 4c)
BM(NH) 9.1.16.7 Flying Fish Co e, Ch is mas I, Indian Ocean.
Pipis ellus papuanus
BM(NH) 22.2.2.3 F ed ik Hend ik I, I ian Jaya. (Fig. 2c)
BM(NH) 34.1.14.8 Kokoda, Papua New Guinea.
Pipis ellus enuis
BM(NH) 85.912 Coas o Sabah, Bo neo (ni idus). (Fig. 9d)
Pipis ellus ceylonicus
BM(NH) 95.6.12.1 Pundibiya, India.
BM(NH) 2.4.2.8 As oli, Belgoum, India. (Fig. 7d)
BM(NH) 1 1.4.5.5 Lanje, Konkan, India.
BM(NH) 13.9.8.102 Guje a , India.
BM(NH) 9.1.4.73 Mangalo e, Malaba Coas , India (Holo ype indicus)
BM(NH) 4.6.8.7-8 Tonkin, Vie nam ( ap o ). (Fig. 3d, BM(NH) 4.6.8.7 Holo ype).
Pipis ellus c assulus
BM(NH) 4.2.8.1 E ulen, Came oun (Holo ype). (Fig. 7e)
Pipis ellus nanulus
BM(NH) 4.2.8.8 E ulen, Came oun (Holo ype). (Fig. I )
BM(NH) 79.508 Sou h Nimba, Libe ia.
Pipis ellus ueppellii
BM(NH) 68.12.22.3 Zanziba (Holo ype pulche ). (Fig. lOa)
BM(NH) 99.9.9.20 Egyp .
BM(NH) - - Uganda. (Fig. lOb)
HZM 3.3170 Kabompo Boma, Zambia.
HZM 7. 12109 Suez, Egyp .
Pipis ellus dese i
BM(NH) 79.987 Hogga Pla eau, Alge ia. (Fig. 5c)
NMW 27503 (?) Uppe Egyp .
VESPERTILIONINE SYSTEMATICS 273
Pipis ellus kuhlii
BM(NH) 92.9.9.25 Uppe Egyp .
BM(NH) A gos oli, Cephaloni, G eece. (Fig. 5a)
BM(NH) 63.335 Sangha, Malya Khola, E Nepal.
HZM 5. 1 1607 Ho e o, nea Volos, G eece.
HZM 11.1016 Rapallo, N I aly.
HZM 138.4563 Yal bu Hillal, Ba inah, Oman.
HZM 154.4619 Saham, Ba inah, Oman.
HZM 203.7232 Dig Dagga, Ras al Khaima, Uni ed A ab Republic.
HZM 218.7402 Benghazi, Libya.
HZM 227.91 10 Kapsowa , Ma akwa , Kenya.
Pipis ellus made ensis
BM(NH) 86.528 Madei a. (Fig. 5b)
Pipis ellus us icus
BM(NH) 35.9.1.108 Oka ango-Oma ako Junc ion, G oo on ein Dis ic , Namibia.
BM(NH) 79.1731 Oli Ri e , Bo gu G.R., Nige ia. (Fig. 6c)
HZM 4.3285 Sen inel Ranch, Ri e Limpopo, Zimbabwe. (Fig. 5d)
Pipis ellus ( Vespadelus}
Pipis ellus pumilus
BM(NH) 70.1093 E Boni hon Range, C Aus alia 2342'S, 12902'E, 1400 .
BM(NH) 71.1497 Wes wood, nea Rockhamp on, Queensland, Aus alia. (Fig. 12k)
Pipis ellus (Pe imyo is)
Pipis ellus sub la us
HZM 1.2422 Big Wyando e Ca e, C aw o d Coun y, Indiana, USA. (Fig. 2d)
Pipis ellus (Hypsugo)
Pipis ellus anchie ae
BM(NH) 69.1248 Ngoma, Zambia.
BM(NH) 70.2632 Balo ale, Zambia. (Fig. 6e)
BM(NH) 89.5.1.5 Caconda, Angola (Syn ype o Vespe us bicolo Bocage, 1889). (Fig. 9e)
Pipis ellus bodenheime i
HZM 3.3786 Jazi a al Abid, Aden, Sou h Yemen.
HZM 5.8279 Bin Gedi, Is ael. (Fig. 90
Pipis ellus sa ii
BM(NH) 31.11.11.13, BM(NH) 66.4644 E slope o Moun Olympus, G eece.
BM(NH) 61.395 Ainab, Lebanon. (Fig. 6a)
Pipis ellus a abicus
HZM 4.10060 Wadi Sah an, Oman.
HZM 5.1 1625 Wadi Fidah, Dank/Ib i, Oman. (Fig. 7a)
Pipis ellus helios
BM(NH) 39.133 N Guaso Nyi o, Kenya. (Fig. 6d)
BM(NH) 69.207 Kanga e , S Tu kana, Kenya.
HZM 2.4086 A che 's Pos , No he n F on ie Dis ic , Kenya.
Pipis ellus nanus
BM(NH) 49.484 Kon au , Gambia.
HZM 3.2778 Soko o, N Nige ia.
HZM 3.4026, HZM 4.4027 Nea Mon o ia, Libe ia.
HZM 83.4387 Ha oni-Lusi u Beacon 74, Zimbabwe.
HZM 107.3212 Kabompo Boma, Zambia.
HZM 146.5161, HZM 147.5162 Rondo, Lindi, Tanzania.
HZM 165.5321 Liwale, Tanzania.
HZM 200.6581 Ka onga, Malawi.
HZM 258.1 1469 Kunyale S eam, Mwinilunga Dis ic , Zambia.
274 J. E. HILL & D. L. HARRISON
HZM 260.12175 Lam o, I o y Coas .
HZM 26 1 . 1 2 1 76 I o y Coas .
HZM 263.12451, HZM 264.12452 Kamuani A ea, Machakos Dis ic , Kenya. (Fig. 6b, HZM 263.12451)
Pipis ellus pul e a us
BM(NH) 79.702 Nea Nicholson Goa Bungalows, Hong Kong I.
BM(NH) 79.903 Peace Mansion, Tai Hang Road, New Te i o ies, Hong Kong (Fig. 8c)
Pipis ellus Hespe us
BM(NH) 98.3.1.8 Sie a Laguna, Baja Cali o nia, Mexico.
BM(NH) 29.1 1.7.10 Panamin M s, Cali o nia, USA.
HZM 4.1 1219 Sycamo e Well, Hidalgo Coun y, New Mexico, USA. (Fig. 8d)
Pipis ellus eisen au i
BM(NH) 84.1684, BM(NH) 84.1686 Moun Came oun, Came oun. (Fig. 9g, BM(NH) 84.1684)
Pipis ellus imb ica us
BM(NH) 9.1.5.286 Bui enzo g, Ja a. (Fig. 9a)
Pipis ellus mac o is
BM(NH) 23.1.2.12 Sabang, NW Suma a. (Fig. 9b)
Pipis ellus ki chene i
BM(NH) 10.4.5.47 Boen ok, Ba i o Ri e , Kaliman an, SC Bo neo. (Fig. 8e)
Pipis ellus lophu us
BM(NH) 14.12.1.6 Maliwun, Vic o ia P o ince, Tenasse im, Bu ma (Holo ype). (Fig. 8 )
Pipis ellus s enop e us
BM(NH) 60.1537 Ins i u e o Medical Resea ch Compound, Kuala Lumpu , Malaya.
BM(NH) 65.135 Pasi Road, Kuala Lumpu , Malaya. (Fig. 7h)
Pipis ellus (Falsis ellus)
Pipis ellus a inis
BM(NH) 83.3.3.2 Wynaa d, India. (Fig. 8a)
BM(NH) 72.4224 A ga awa, Ne awa Elwa, Cen al P o ince, S i Lanka.
Pipis ellus pe e si
BM(NH) 23. 1 .2.3. Bu u I, Molucca Is (Fig. 8b)
Pipis ellus asmaniensis
HZM 1.8712 Ba ing on Tops Na ional Pa k, New Sou h Wales, Aus alia. (Fig. 8g)
Pipis ellus (Neo omicid)
Pipis ellus capensis
BM(NH) 32.9.1.249 B oken Hill, Zambia.
BM(NH) 54.859 Elizabe h ille, Zai e.
BM(NH) 61.1078 Doddiebu n Ranch, Wes Nicholson, Zimbabwe, 2300 , 2124'S, 2921'E.
BM(NH) 72.4383 E o Lake Ma gha i a, Bulcha Fo es , E hiopia, 1800 m, 06 1 1'N, 3610'E.
BM(NH) 72.4391 Didessa Ri e , Wollega P o ince, E hiopia, 1 190 m, 0902'N, 3609'E. (Fig. 12g)
BM(NH) 75.561 Mole Na ional Pa k, Ghana. (Fig. 12b)
BM(NH) 83.200 Mcheni Go ge, Chiza i a Na ional Pa k, Binga P o ince, Zimbabwe, 1740'S, 2752'E.
HZM 36.4514 40 m NW o Se owe, Bo swana.
BM(NH) 66.6057 Ambosi a, Madagasca (ma okd). (Fig. 12a)
BM(NH) 77.2.19.6 Anzahame u, Madagasca ('minu us'). (Fig. 12i)
Pipis ellus guineensis
BM(NH) 70.2224, BM(NH) 70.2228, BM(NH) 72.4373 Gambela, E hiopia, 815'N, 3435'E (BM(NH)
72.4373 a 515 m) (Fig. 12c, BM(NH) 70.2224)
BM(NH) 76.293 Shagamu, Nige ia.
BM(NH) 84.1019 Bon ioli, Bougou iba Ri e , Bu kina Faso (Uppe Vol a).
Pipis ellus melcko um
BM(NH) 83.216 Mcheni Go ge, Chiza i a Na ional Pa k, Binga P o ince, Zimbabwe, 1740'S, 2752'E.
(Fig. 12Q
VESPERTILIONINE SYSTEMATICS 275
Pipis ellus somalicus
BM(NH) 70.484 Mou h o Fincha Ri e , Blue Nile Go ge, E hiopia, 1003'N, 3720'E. (Fig. 12h)
BM(NH) 76.814 S bank o Ganale Do ia, Sidam-Bale B idge, Sidamo P o ince, E hiopia, 545'N, 3937'E.
BM(NH) 84.1016 Comoe Ri e , Bu kina Faso (Uppe Vol a), 260 m, 957'N, 438'W.
CMNH MJS 2846 Snai Suga Plan a ion, l km S, km E o Gioha , Somalia, 246'N, 453 1 'E.
Pipis ellus zuluensis
BM(NH) 83.212 Mchesu Ri e , Chiza i a Na ional Pa k, Binga P o ince, Zimbabwe, 1747'S, 2739'E.
BM(NH) 83.215 Singama, Sibuwa, Binga P o ince, Zimbabwe, 17 C 36'S, 2751'E. (Fig. 12d)
Pipis ellus endalli
BM(NH)89.12.12.1 Ba hu s , Gambia.
BM(NH) 7.12.17.1-2 Gondoko o, Whi e Nile, Sudan.
BM(NH) 23.4.12.1-2 Bugala, Sesse Is, Vic o ia Nyanza, Uganda. (Fig. 12e, BM(NH) 23.4.12.2)
BM(NH) 48.702 N'ko, Obub a Di ision, S Nige ia (Ib unneus). (Fig. 14b)
Pipis ellus enuipinnis
BM(NH) 47.350 Umuahia, E Nige ia.
BM(NH) 54.917 Bon he, Sie a Leone.
BM(NH) 67.1734 Bo a, Vic o ia, Came oun, 400'N, 905'E. (Fig. 12j)
Pipis ellus (A ielulus)
Pipis ellus ci cumda us
BM(NH) 73.618 Telecommunica ions Towe , F ase 's Hill, Pahang, Malaya. (Fig. 2e)
Pipis ellus cup osus
BM(NH) 83.351 Sepilok, Sabah, Bo neo, 552'N, 1 1756'E (Holo ype). (Fig. 9h)
Pipis ellus socie a is
BM(NH) 67.1605 Base Camp, Gunong Benom, Pahang, Malaya, 800 (Holo ype). (Fig. 9c)
Nyc alus noc ula
BM(NH) - - Locali y unknown.
HZM 10.613 Bo isham, Camb idgeshi e, England.
HZM 33.8888 Winchelsea Beach, Sussex, England. (Fig. 100
Laepho is bo swanae
BM(NH) - - Zomba, Malawi (o iginal No. 2269; damaged).
Laepho is win oni
HZM 1.3020 Nye i, Moun Kenya, Kenya. (Fig. 160
Glisch opus ylopus
BM(NH) 10.4.5.136 Uppe Ba i o Ri e , Kaliman an, SC Bo neo. (Fig. 18a)
Sco ozous do me i
BM(NH) 12.3.8.30 Fu dapu , Ajan a, Khandesh, India.
BM(NH) - - Ka hiawa , India (O iginal No. BNHS 2007). (Fig. 1 6d)
Sco eanax ueppellii
BM(NH) 80.3.25.1 Richmond Ri e , New Sou h Wales, Aus alia. (Fig. 16i)
Sco o epens bals oni
BM(NH) 10.6.21.9 He mannsbu g, No he n Te i o y, Aus alia. (Fig. 16g)
Sco o epens g eyii
BM(NH) 75.2261 Pine C eek, 20 m ESE o Candy's Hill, No he n Te i o y, Aus alia, 1349'S, 13149'E.
(Fig. 16h)
Nyc iceinops schlie enii
BM(NH) 14.7.3 1.14 Wei Wei Ri e , Kenya.
BM(NH) 15.3.6.66 Kamisu, Dinda Ri e , Sudan.
BM(NH) 71.675 Awash, Filhoa, E hiopia, 0900'N, 3858'E.
HZM 5.2120 Ikau, Rukwa, Tanzania. (Fig. 16e)
Sco oecus albigula
BM(NH) 63.1042 Calundo, Lunda, Angola. (Fig. 20a)
276 J. E. HILL & D. L. HARRISON
Sco oecus albo uscus
BM(NH) 96. 1 2.3 1 . 1 Sie a Leone. (Fig. 20e)
Sco oecus hindei
BM(NH) 14.7.31.13 30m NW o Ba ingo, Kenya. (Fig. 20d)
BM(NH) 66.1466 Jos, Nige ia ( alabae). (Fig. 20b)
Sco oecus hi undo
BM(NH) 76.771 Mole Na ional Pa k, Ghana. (Fig. 20c)
Sco oecus pallidus
BM(NH) 86.531 A ghanis an (damaged).
Phile o b achyp e us
BM(NH) - - New Guinea. (Fig. 16b)
Hespe op enus (Mili h onyc e is)
Hespe op enus blan o di
BM(NH) 83.853 Sepilok, Sabah, Bo neo, 552'N, 1 1756'E. (Fig. 21g)
Hespe op enus ickelli
BM(NH) 71.12.26.1 S i Lanka. (Fig. 21b)
Hespe op enus omesi
BM(NH) 7.1.1.428 Malacca, Malaya (Holo ype). (Fig. 2 la)
Chalinolobus gouldi
BM(NH) 71.1504 Wes wood, nea Rockhamp on, Queensland, Aus alia. (Fig. 17b)
Chalinolobus mo io
BM(NH) 6.8.1.60 (King Ri e , Wes e n Aus alia. (Fig. 17a)
Chalinolobus nig og iseus
BM(NH) 44.6.13.2 Po Essing on, No he n Te i o y, Aus alia ( oge si).
BM(NH) 75.2260 Pine C eek, 20 m ESE o Gandy's Hill, No he n Te i o y, Aus alia, 1349'S, 13149'E.
( oge si). (Fig. 17c)
Chalinolobus pica us
BM(NH) 9.3.7.2 Gunnamulla, Queensland, Aus alia. (Fig. 17d)
Chalinolobus ube cula us
BM(NH) 89.10.27.1 Ou lying islands nea S ewa I, New Zealand. (Fig. 17e)
Nyc ophilinae
Nyc ophilus bi ax
BM(NH) 67.5.6.5 Cape Yo k, Queensland, Aus alia.
BM(NH) 77.3.28.1 Islands o To es S ai s, Aus alia.
BM(NH) 86.1 1.8.12 Some se , Cape Yo k, Queensland, Aus alia.
BM(NH) 15.3.13.1 Cloncu y, Queensland, Aus alia.
BM(NH) 15.3.13.3 He be on Dis ic , Queensland, Aus alia (Holo ype). (Fig. 22a)
Nyc ophilus daedalus
BM(NH) 47.7.2 1 . 1 6, BM(NH) - - Po Essing on, No he n Te i o y, Aus alia. (Fig. 22g, BM(NH)
47.7.21.16)
BM(NH) 97.4.12.5 Daly Ri e , No he n Te i o y, Aus alia.
Nyc ophilus gouldi
BM(NH) 15.3.13.7 Ash I, Hun e Ri e , New Sou h Wales, Aus alia (damaged, pa los ).
BM(NH) 15.3.13.8 Sydney, New Sou h Wales, Aus alia.
BM(NH) - - Bo any, Sydney, New Sou h Wales, Aus alia (O iginal No. 164) (Fig. 22d)
HZM 1.12085 We ikimbe, Has ingsshi e, New Sou h Wales, Aus alia. (Fig. 16c)
BM(NH) 52. 1 . 1 5.30 Tasmania (she ini). (Fig. 220
Nyc ophilus geo oyi
BM(NH) 15.3.13.1 1 Kosciusko, New Sou h Wales, Aus alia (paci icus).
BM(NH) - - Tasmania (paci icus) (O iginal No. M.I 735).
VESPERTILIONINE SYSTEMATICS 277
BM(NH) - - Launces on, Tasmania (paci icus) (O iginal No. M. 168) (Fig. 22e)
BM(NH) 7.1.4.3 Alexand ia, No he n Te i o y, Aus alia (pallescens). (Fig. 22b)
Nyc ophilus mic o is
BM(NH) 88.4.18.1 Soge i, Papua New Guinea (Holo ype). (Fig. 22c)
Pha o is imogene
BM(NH) 97.8.7.21 Kamali, Papua New Guinea. (Fig. 22h)
278 J. E. HILL & D. L. HARRISON
VESPERTILIONINE SYSTEMATICS 279
280 J. E. HILL & D. L. HARRISON
Table 2 Usual incisi e and p emola den al o mulae in he Vespe ilioninae and Nyc ophilinae. To al
numbe o ee h (including ou canines and wel e mola s) in pa en heses. Den al no a ion o Mille ( 1 907).
(38) Myo is, Pizonyx
(36) Lasionyc e is, Pleco us, Idionyc e is, Eudiscopus
(34) Eude ma, Ba bas ella, la, Pipis ellus, Glisch opus, Sco ozous, Nyc alus,
Chalinolobus
(32) Ep esicus, Vespe ilio, His io us, Tylonyc e is , Mime illus, Glauconyc e is,
Pipis ellus, Laepho is, Phile o , Hespe op enus
(32) Lasiu us
(30) Rhogeessa, Baeodon, Nyc iceius, O onyc e is, Dasyp e us, Sco omanes,
Sco ophilus, Sco eanax, Sco o epens, Nyc iceinops, Sco oecus,
Nyc ophilus, Pha o is
(28) An ozous, Baue us
Table 3 Classi ica ions o he Vespe ilioninae and Nyc ophilinae. Tha o Ta e ( 942a) is conce ned p ima ily wi h
O ien al and Aus alasian axa, hose o Koopman wi h Aus alasian (1973) and p edominan ly A ican (1975) o ms.
Ta e(1942a) Koopman (1973, 1975) Hill & Ha ison
Pipis ellus
ab amus g oup
ab amus
akokomuli
bancanus
camo ae
i e i us
pa e culus
pumiloides
pipis ellus g oup
pipis ellus (Including
bac ianus)
na husii
co omand a g oup
aladdin
angula us
collinus
co omand a
imb ica us
meyeni
mic opus
mu ayi
poncele i
po ensis
egulus
s u deei
subulidens
ama us
enuis g oup
mimus (Including
glaucillus)
ni idus
papuanus (Including
o ien alis)
p incipulus
enuis
ceylonicus g oup
ceylonicus (Including
ch yso h ix, indicus,
subcanus)
Pipis ellus
Amalgama es pipis ellus, ab amus
( =ja anicus), co omand a and
enuis g oups o Ta e (1942a)
pipis ellus g oup
imb ica us
ja anicus (Including
ab amus)
meyeni
nanus (Including
(?) Helios)
pe mix us
enuis (Including angula us,
collinus, ni idus, papuanus,
poncele i, mu ayi, sewelanus,
subulidens, wes alis [Koopman,
1984c])
ceylonicus g oup
ceylonicus
Pipis ellus
Pipis ellus (Pipis ellus)
pipis ellus g oup
pipis ellus subg oup
pipis ellus (Including aladdin,
bac ianus, lac eus,
medi e aneus)
na husii
pe mix us
ja anicus subg oup
ab amus (Including
akokomuli, i e i us,
pumiloides)
babu
endoi
ja anicus (Including bancanus,
camo ae, meyeni,
' ala i ius")
pa e culus
peguensis
co omand a subg oup
adamsi
angula us (Including poncele i)
collinus
co omand a (Including a ghanus,
po ensis, ama us)
mimus (Including
glaucillus, p incipulus)
mu ayi
papuanus
s u deei
enuis (Including ni idus,
sewelanus, subulidens)
wa si
wes alis
ceylonicus subg oup
ceylonicus (Including bo neanus,
ch yso h ix, indicus, ap o ,
shano um, subcanus)
(?) minahassae
VESPERTILIONINE SYSTEMATICS 281
Table 3-con .
Ta e(1942a) Koopman (1973, 1975) Hill & Ha ison
minahassae g oup
minahassae
ueppellii g oup
coxi
kuhlii g oup
babu
canus
kuhlii (Including
ikhwanius, lepidus)
leuco is
loba us
Ep esicus
pumilus g oup
pumilus (Including
cau inus, da ling oni,
ul u nus)
pygmaeus
Pipis ellus
sa ii g oup
aus enianus
cado nae
cu a us
mac o is
sa ii
o de manni
minahassae g oup
minahassae
ueppellii g oup
ueppellii (Including (?)
uscipes; pulche )
kuhlii g oup
ae o
anchie ae
dese i
inexspec a us
kuhlii (Including (?)
aegyp ius; usca us)
us icus (Including ma ensis)
sa ii g oup
a iel
mac o is
made ensis
Hespe us g oup
hespe us
musciculus
jo ei g oup
an honyi
jo ei
s enop e us
jo ei g oup
s enop e us
ueppellii g oup
c assulus
nanulus
ueppellii (Including coxi,
uscipes, leucomelas, pulche ,
senegalensis , e nayi)
kuhlii g oup
ae o
dese i
inexspec a us
kuhlii (Including (?)
aegyp ius; usca us,
ikhwanius)
made ensis
us icus (Including ma ensis)
Pipis ellus ( Vespadelus)
douglaso um
pumilus (Including
da ling oni)
egulus
sagi ula
ul u nus
Pipis ellus (Pe imyo is)
sub la us
Pipis ellus (Hypsugo)
sa ii g oup
sa ii subg oup
anchie ae ( = 'bicolo '7)
a iel
aus enianus
bodenheime i
sa ii (Including caucasicus,
da wini, mau us)
nanus subg oup
a abicus
helios
musciculus
nanus (Including culex,
s amp lii)
pul e a us subg oup
pul e a us
hespe us subg oup
hespe us
eisen au i subg oup
eisen au i
imb ica us subg oup
cu a us
imb ica us
mac o is
o de manni
lophu us subg oup
cado nae
ki chene i
lophu us
s enop e us g oup
an honyi
jo ei
s enop e us
^.......:n.<^ ,,.:... ; :... 1 ......,.,. " w " '^ >
Fig. 5 Baculum (D, RL) o a, Pipis ellus kuhlii; b, P. made ensis; c, P. dese i; d, P. us icus.
Scale = 0-5 mm.
VESPERTILIONINE SYSTEMATICS 289
Fig. 6 Baculum o a, Pipis ellus sa ii (D, RL); b, P. HO/IMS (D, RL); c, P. us icus (D, RL); d, P. helios
(D, RL); e, P. anchie ae (D, LVL, e e sed). Scale = 0-5 mm.
S?> ^KW^^ .,,,...-,.,.:;:.,. , : -..,.. wmc. .l.UW
Fig. 7 Baculum o a, Pipis ellus a abicus (D, RL, RVL); b, P. co omand a ( ama us) (D, RL); c, P.
co omand a (D, RL); d, P. ceylonicus (D, RL); e, P. c assulus (D); , P. nanulus (D, RL); g, P. mimus
(D, RL); h, P. s enop e us (D, RL, RVL). Scale = 1 mm.
Fig. 8 Baculum o a, Pipis ellus a inis (D, RL); b, P. pe e si (D, RL); c, P. pul e a us (D, RL, RVL); d,
P. Hespe us (D, LL, e e sed, LVL); e, P. ki chene i (D, RL, RVL); , P. lophu us (D, RL); g, P.
asmaniensis (D, RL, V). Scale = 1 mm.
Fig. 9 Baculum (D, RL excep whe e s a ed) o a, Pipis ellus imb ica us; b, P. mac o is; c, P. socie a is,
d, P. ennis (ni idus) (D, RL, RVL); e, P. anchie ae ('Vespe us' bicolo ); , P. bodenheime i; g, P.
eisen au i; h, P. cup osus. Scales a-g= 1 mm; h = 0-5 mm.
_l
Fig. 10 Baculum (D, RL) o a, Pipis ellus ueppellii (pulche ); b, P. ueppellii; c, P. adamsi; d, P.
wes alis; e, P. ja anicus; , Nyc alus noc ula; g, P. wa si; h, P. mackenziei (c, d, g, h om Ki chene
e al, 1986). Scales = a, b, e, =2mm;c, d, g, h= 1 mm.
294 J. E. HILL & D. L. HARRISON
Fig. 1 1 Baculum (V, RL) o a, Pipis ellus pumilus pumilus; b, P. pumilus (cau inus); c, P. ul u nus; d, P.
douglaso um; e, P. egulus; , P. sagi ula (a-c, e, om McKean e al., 1970; d om Ki chene , 1976).
Scale = 2 mm.
Fig. 12 Baculum (D, RL) o a, Pipis ellus capensis (ma okd); b, P. capensis; c, P. guineensis; d, P.
zuluensis; e, P. endalli (wi h an e io iew); , P. melcko um; g, P. capensis; h, P. somalicus', i, P.
capensis ('minu us'); j, P. enuipinnis; k, P.pumilus. Scale = 1 mm.
Fig. 13 Baculum (D, RL) o a, Ep esicus uscus; b, E. ho en o us (megalu us); c, E. u inalis; d, E.
b asiliensis (andinus); e, E. bob inskoi; , E. lowe i; g, E. se o inus; h, E. se o inus (isabellinus); i, E.
uscus (hispaniolae); j, . bo ae (innesi); k, . b asiliensis: 1, E. lowe i(lowei). Scale = 1 mm.
VESPERTILIONINE SYSTEMATICS 297
Fig. 14 Baculum (D, RL excep whe e s a ed) o a, Ep esicus bo ae (omanensis); b, Pipis ellus endalli
(? b unneus); c, Ep esicus nasu us; d, Pleco us ene i ae (D) ( om Ibanez & Fe nandez, 1986).
Scales = 1 mm.
d %**^
I I
Fig. 21 Baculum (D, RL excep whe e s a ed) o a, Hespe op enus omesi, b, H. ickelli, c, H. do iae (a-c
om Hill, 1 976); d, la io (D) ( om Topal, 1 970); e, Sco o epens o ion, , S. sanbo ni (e, om Ki chene
& Capu i, 1985); g, Hespe op enus blan o di ( om Hill & F ancis, 1984); h, Sco oecus pallidus ( om
Ag awal & Sinha, 1973); i, Vespe ilio mu inus (V, RL) ( om Topal, 1958); j, V. o ien alis ( om
Wallin, 1969). Scales a-c = 2 mm; d-h, j= 1 mm; i = 0-5 mm.
_- ' -
Fig. 22 Baculum (D, RL) o a, Nyc ophilus bi ax; b, AT. geo oyi (pallescens); c, M mic o is; d, A . gouldi;
e, A 1 , geo oyi (paci icus); , A 7 ^. gouldi (she ini); g, A 7 , daedalus; h, Pha o is imogene. Scale = 2 mm.
Hill, J. E. and Ha ison, Da id L. 1987. "The baculum in he Vespe ilioninae
(Chi op e a: Vespe ilionidae) wi h a sys ema ic e iew, a synopsis o
Pipis ellus and Ep esicus, and he desc ip ions o a new genus and
subgenus." Bulle in o he B i ish Museum (Na u al His o y) Zoology 52, 225–305.
h ps://doi.o g/10.5962/p.18307.
View This I em Online: h ps://www.biodi e si ylib a y.o g/i em/19520
DOI: h ps://doi.o g/10.5962/p.18307
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