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MAMMALIAN
SPECIES No. 260, pp. 1-4, 3 igs.
Mac ode ma gigas. By Wendy S a Hudson and Don E. Wilson
Published 16 June 1986 by The Ame ican Socie y o Mammalogis s
Mac ode ma Mille , 1906 56, 47; leng h
o an e io lobe o agus,
12, 10; leng h
o pos e io
lobe o agus, 25.5, 22.0; leng h o noselea , 16, 15; leng h o
Mac ode ma Mille , 1906:84. Type species Megade ma gigas by o ea m, 117.0, 103.5; leng h o humb, 21.0, 19.5; leng h o
o iginal designa ion. me aca pal II, 84.5, 80.5; leng h o phalanx II, 16, 17; leng h o
CONTEXT AND CONTENT. O de
Chi op e a,
Subo de me aca pal III, 69, 68; digi III, leng h o phalanx I, 47, 44; digi
Mic ochi op e a,
Family Megade ma idae.
The genus Mac ode ma III, leng h o phalanx II, 92, 80; leng h o me aca pal IV, 79, 78;
con ains
only
one Recen
species,
M. gigas, and a ossil
o m,
M. digi IV, leng h o phalanx I, 25.5, 24.5; digi IV, leng h o phalanx
god helpi Hand (1985). II, 38.5, 33.0; leng h o me aca pal V, 84.5, 82.0; digi V, leng h
o phalanx
I, 32, 33; digi
V, leng h
o phalanx
II, 28.5, 23.5;
Mac ode ma gigas (Dobson, 1880)
Ghos Ba
Megade ma gigas Dobson, 1880:461. Type locali y Moun Ma -
ga e , Wilson's
Ri e ,
cen al
Queensland, Aus alia.
CONTEXT AND CONTENT. Con ex same as o genus.
Douglas (1962) named M. g. sa u a a on he basis o specimens^ : *"
-
om Kalumbu u in no he n Wes e n Aus alia, bu Koopman <
~
..
(1984a) synonymized i wi h he nomina e
o m. _ -
DIAGNOSIS. The uppe inciso s a e absen as in all mega-
de ma ids,
bu he udimen a y
p emaxillae a e mo e highly de el-
oped in Mac ode ma han in Megade ma. M. gigas lacks P2, in
con as o species
o Megade ma
(Mille , 1906). As in o he
mem-
be s o he amily,
he mesos yles
o M1 and M2 a e displaced
lingually,
mo e
so in Mac ode ma han in La ia o Ca diode ma.
Reduc ion o loss o he
hypoconulid
o m3 cha ac e izes
he
amily,
bu
Mac ode ma has a be e
de eloped
m3 hypoconulid
han o he
gene a
in he amily
(Hand, 1985). The shield-like in e o bi al ex-
pansion
(Fig. 1) is in e media e in size be ween ha o he Asian
and A ican
megade ma ids.
The
pos o bi al
and an o bi al
p ocess-
es a e mo e
p ominen
han in Megade ma,
bu less so han in
Ca diode ma
and La ia (Mille ,
1907). The in e p e ygoid
space
ex ends o he le el o he pos e io
bo de o M2. The pos e io
lobe o he
agus
is p opo ionally
sho e han
ha
o Megade ma
and he an e io lobe is much b oade a he base, mo e con ex 96..8Z9
an e io ly,
and ob use a he
ip
(Dobson,
1880). The
second
inge
ex ends
beyond
he
middle o he i s
phalanx
o he hi d
inge ,
in con as
o he sho e
second
- inge
o Megade ma
spasma.
The ail consis s
o only wo e eb ae
hidden
be ween he wo
in egumen a y
laye s
o he la ge in e emo al
memb ane. Sho
hai s co e he ex emi y
o he ca pus,
humb,
and
p opa agium.
GENERAL
CHARACTERS.
The
Aus alian
ghos
ba ,
M.
gigas, wi h a body
mass om 130 o 170 g (Kulze
e al., 1984;
Pi lo and
Nelson, 1980; Taylo ,
1984), is one
o he la ges
o he -
mic ochi op e ans.
Head and
body
leng h
anges
om
100 o 140
mm and o ea m
leng h
om
105 o 115 mm
(Nowak
and
Pa adiso,
1983). The long
ea s
join along
he midline
o hal
he leng h
o
hei inne
ma gin
(Fig. 2). The noselea is long and has con ex
sides and i s an e io conca e disk is la ge
wi h a e ical
p ocess
abo e (T ough on, 1926). The eyes a e ela i ely la ge o mic o-
chi op e ans
and he claws a e s ong
and cu ed
(Taylo ,
1984).
The lowe
lip p ojec s
a he han he uppe
and he nos ils a e
dep essed
wi hin an a ea o naked skin. In mos
popula ions,
he
e minal
hi d o he do sal hai s is pale g ayish
b own,
and he
pale whi e o he en al
pelage
ma ches he ea s, noselea ,
and
memb anes
(T ough on,
1926). The species
also encompasses
in-
di iduals wi h da ke u and
memb anes
(Koopman,
1984a). The
wing
span
o M. gigas is app oxima ely
0.6 m (B eeden
and B ee-
den, 1967). The emale is smalle
han he male, al hough
he
measu emen s
o he ibia and i s
phalanx
o he i h
inge
a e
g ea e
in he emale
(Wai e,
1900).
Ex e nal measu emen s (in mm) o he male holo ype and o FIG. 1. Do sal, en al, and la e al iews o c anium and
a emale om he Aus alian Museum (Wai e, 1900) a e: head and la e al iew o mandible o emale Mac ode ma gigas, USNM
body leng h, 135, 110; head leng h, 48.5, 41.0 (b oken);
ea leng h, 396829. G ea es leng h o skull is 38.7 mm.
MAMMALIAN
SPECIES 260
120 140
25
25'-
0 KMS
1000 KMS
120" 140
FIG. 2. Mac ode ma gigas pho og aphed in Aus alia by
F ank G eenwell
in 1973.
ibia leng h, 44, 45; calcaneum leng h 28.5, 28.0; and leng h o
hind oo , 28.5, 23.5.
Douglas (1962) lis ed he ollowing skull measu emen s (in
mm) o a specimen om Wes e n Aus alia:
leng h, 38.5; b ead h,
24.3; in e o bi al
cons ic ion,
4; and
pala e, 13. Molna e al. (1984)
desc ibed se e al ossil specimens e e able o he Recen species,
and lis ed he ollowing measu emen s (in mm) o a Recen speci-
men om he Queensland Museum; leng h o mandibula
amus,
29.2; leng h and wid h o p4, 3.2, 1.9; leng h and wid h o ml,
3.5, 2.6; leng h and wid h o m2, 3.7, 2.6; leng h and wid h o
m3, 3.6, 2.4.
DISTRIBUTION. Mac ode ma p esen ly occu s h ough-
ou he no he n wo- hi ds
o Aus alia sou h o 28?S in Wes e n
Aus alia
and 27?S in wes e n Queensland (Fig. 3). I is ound bo h
in d y coun y and in humid ain o es s (Molna e al., 1984). A
sigh ing on M . Kenne h, Wes e n Aus alia, in 1854 is he sou h-
e nmos eco d;
howe e , mos eco ds a he sou he n limi s consis
o isola ed indi iduals a he han pe manen popula ions. Reco ds
o la ge, pe manen popula ions
a he sou he n bo de o i s ange
ex end om he Pilba a dis ic in Wes e n Aus alia o M . E na
in Queensland (Molna e al., 1984). Pleis ocene locali ies coincide
wi h he p esen dis ibu ion
only in no heas e n
Queensland.
Mac-
ode ma occu ed much a he sou h in he Pleis ocene han i
does oday (Cook, 1960). Finlayson (1961) conside ed he species
o be elic ual in cen al Aus alia and sugges ed ha i s ange is
eceding om he sou h. Baynes e al. (1976) sugges ed ha he
absence o d y, smoo h-walled
ca es in sou he n Aus alia
limi s i s
ange. The genus may occu in sui able habi a in New Guinea
(Filewood, 1983), al hough no specimens ha e been aken he e.
FOSSIL RECORD. The mos sou he ly ossil occu ence is
app oxima ely 600 km SSW o M . Kenne h, Wes e n Aus alia,
he sou he nmos eco d o li ing ghos ba s. The Wanne oo-Yan-
chep egion has yielded occasional skele al ma e ial and may ep-
esen only a ecen sou hwa d
expansion o he ange o he species
(B idge, 1975). La ge guano piles and skele al ma e ial we e ound
a he Ledge Poin -Donga a and Coo ow-Wa he oo a eas. B idge
(1975) sugges ed ha he dis ibu ion and abundance o he guano
piles and skele al ma e ial ep esen expansions and con ac ions o
he ange du ing he Holocene.
In D o e 's Ca e, Lundelius (1960) disco e ed M. gigas e-
mains a all dep hs up o 1.5 m in associa ion wi h oden and
ma supial ma e ial. Sub ossils ha e been collec ed om limes one
ca es in Wes e n Aus alia and he Flinde s Ranges o Sou h Aus-
alia (Hamil on-Smi h, 1974).
All ossil specimens ound o da e seem o di e only in mino
ways om mode n ep esen a i es o he species (Molna e al.,
FIG. 3. Dis ibu ion o Mac ode ma gigas (shaded a ea).
Open ci cles deno e ex alimi al
locali ies om which Holocene os-
sils a e known.
1984). Analyses o cu en and pas dis ibu ion
pa e ns sugges
he possibili y
o successi e wa es o expansion and con ac ion o
ange, pe haps
ela ed
o changing
clima ic
condi ions
(B idge, 1975;
Molna e al., 1984).
FORM. Al hough he p edominan pelage colo is pale g ay
o whi ish, he ela i ely long and lax hai anges in colo om
almos whi e h ough shades o g ay o almos b own (Douglas,
1962).
Al hough mos species o ba s ha e only a single pai o pec-
o al
mammae, Mac ode ma has a pai
o inguinal " alse mammae"
which se e as hold as s o he young, in addi ion o he no mally
unc ioning pec o al pai .
The skele al a angemen in he ho acic egion is somewha
unusual in megade ma ids. The an e io ly widened p es e num is
used o he i s pai o ibs, he i s ho acic e eb a, and he
se en h ce ical e eb a o o m a solid ing o bone (Mille , 1907).
The pel ic gi dle is mo e like ha o o he mic ochi op e ans,
wi h
he ischia ee pos e io ly.
The h ead-like
ibula is only hal as long
as he ibia (Koopman, 1984b).
A dis inc
sagi al c es equen ly de elops on he la ge, obus
skull o olde indi iduals. The pen agonal on al shield is la and
i s angles a e de ined dis inc ly.
The den al o mula is i 0/2, c 1/1, p 1/2, m 3/3, o al 26.
The ee h a e la ge, obus , and highly cuspida e. The me as yle o
M1 has shi ed pos e io ly, and he mesos yle, almos obli e a ed,
has shi ed lingually be ween he pa as yle and he me acone. The
an e olingual cingula cusp o C1 is g ea ly de eloped (Ande sen
and W ough on, 1907).
Volumes o 11 b ain s uc u es indica e ha M. gigas is
in e media e be ween insec i o ous mic ochi op e ans
and he ca -
ni o ous species om Sou h Ame ica (Pi lo and Nelson, 1980). M.
gigas is he leas encephalized membe o i s amily wi h an en-
cephaliza ion index o app oxima ely 142 (S ephan e al. 1981).
S ephan and Nelson (1981) desc ibed Mac ode ma as ha ing a
clea midb ain
exposu e, a la ge hemisphe ical pa o he pa a loc-
culus, and a ela i ely small lobulus pe osus. Compa ed o o he
species o ba s, highe de elopmen o he b ain was indica ed by
he pa ial co e o he midb ain, aces o a do sal sulcus, and a
well-de eloped
ce ebellum (S ephan and Nelson, 1981).
FUNCTION. Body empe a u e is main ained be ween 35
and 39?C a ambien empe a u es o 0 o 35?C. Oxygen consump-
ion inc eases abo e and below he he mal neu al zone, 30 o
35?C. As ambien empe a u es dec ease o 25?C, su icien oxygen
is ob ained h ough an inc eased dep h o b ea hing. Below 25?C,
he b ea hing a e is di ec ly p opo ional
o oxygen consump ion.
A 20?C, ele a ed oxygen equi emen s o he issues a e me by
inc eased s oke olume, he a e io enous di e ence in he blood,
and shi e ing. Wi hin he he mal neu al zone, hea a e emains
- - -0 - - -6
a_ I_ n
2
MAMMALIAN SPECIES 260
minimal and inc eases wi h oxygen consump ion. The species does
no hibe na e o unde go daily o po (Lei ne and Nelson, 1967).
When a Mac ode ma is mo ing owa ds de ec ed p ey, i
emi s echoloca ion pulses ha a e sho e in du a ion and a e age
0.8 ms (n = 11); du a ion inc eases o a mean o 1.7 ms (n = 13)
du ing landing (Kulze e al., 1984).
Mac ode ma has unusually good ision o a mic ochi op e -
an, wi h a spa ial esolu ion app oaching wo cycles/deg ee based
on e inal mo phology (Guppy and Coles, 1983). I may well use
ision ex ensi ely o loca e o disc imina e
p ey.
Mas ica ion is ho ough and passage h ough he diges i e
sys em is slow compa ed o ui -ea ing ba s (Douglas, 1967). M.
gigas also may be able o balance hei wa e budge wi hou e-
cou se o ee d inking
wa e (Kulze e al., 1984).
Douglas (1967) desc ibed a mus y odo om Mac ode ma
ha he could dis inguish
eadily om ha o o he species.
ONTOGENY
AND REPRODUCTION. Females
cong e-
ga e in ma e ni y colonies du ing he b eeding season and each
gi es bi h o a single young a he end o he b eeding season
(Tho nback
and Jenkins, 1982). The b eeding
season is la e Oc obe
o ea ly No embe in he sou h and somewha ea lie in he a
no h (Tho nback
and Jenkins, 1982). In he Pilba a
Dis ic , Doug-
las (1967) ound a emale wi h an emb yo 10 mm in c own- ump
leng h in Augus , and h ee o he s wi h emb yos o 52, 65, and 70
mm in Oc obe . In No embe , he ound neona es anging om 60
o 80 mm in leng h. In he No he n Te i o y, he ound a 45-mm
emb yo in June, and a hai less neona e o 96 mm in Sep embe .
The laccid
pec o al and inguinal
mammae o nullipa ous
Mac ode -
ma become u gid a e p egnancy. The mammae and cli o is a e
mo e easily de ec ed in pa ous han in nullipa ous
emales (Douglas,
1967).
ECOLOGY. Al hough buildings
may be used as eeding s a-
ions, Mac ode ma oos s only in ca es, ock c e ices, and mines
(Douglas, 1967). Colonies
o up o se e al hund ed
indi iduals
may
o m, bu small g oups o indi iduals a e mo e common (Taylo ,
1984).
Mac ode ma gigas eeds on mice, small ba s, small bi ds,
legless liza ds (Pygopodidae), geckos (Gekkonidae), small snakes,
and insec s. Mus musculus emains we e ound mos commonly in
disca ded ood ma e ial in ca es o he Pilba a Dis ic . Twen y
species o bi ds
also we e iden i ied in he ma e ial
(Douglas, 1967).
Wilson (1973) classi ied Mac ode ma as p ima ily a specialized
ca ni o e and as a oliage gleane seconda ily. The occasional ind-
ing o la ge quan i ies o insec chi in in he s omachs o M. gigas
sugges s ha he ba is an oppo unis ic
eede (Kulze
e al., 1984).
Ves jens and Hall (1977) epo ed e mi es in he s omach o one
indi idual.
Li e p ey is ea en mo e equen ly and no mally
is ea en
nea he poin o cap u e (Douglas, 1967).
The ghos ba has ew na u al p eda o s, Medium-sized owls
(S igidae) co'mpe e
o insec s and small mammals, bu no co e-
la ion exis s be ween p esence o hese owls and Mac ode ma. Local
mo emen s o colonies and indi iduals mos likely esul om luc-
ua ions in p ey a ailabili y
(Molna e al., 1984).
Al hough he ange seemingly has con ac ed du ing he Ho-
locene Pe iod, p obably
se e al housand Mac ode ma emain
ali e.
Some a eas o sou he n Aus alia unde wen a ansi ion om open
sa anna o dense woodland du ing he Pleis ocene and Recen .
These changes may ha e led o he con ac ion o he ange o he
species in Aus alia
(Tho nback
and Jenkins, 1982). The species is
h ea ened by andalism (McKean and P ice, 1967), mining o
limes one ca es, and qua ying
(Hamil on-Smi h,
1980). The e a e
ew na ional pa ks o ese es o p o ide e ugia (Hamil on-Smi h,
1980). M. gigas is classi ied as a ulne able species by he In e -
na ional Union o he Conse a ion o Na u e and Na u al Re-
sou ces (Tho nback
and Jenkins, 1982).
A species o ick, A gas (C.) mac ode mae, was desc ibed
in 1977 om specimens o he ghos ba (Hoogs aal
e al., 1977).
Douglas (1967) sugges ed ha Mac ode ma is conspicuously ee
o ec opa asi es in compa ison wi h o he mic ochi op e ans.
The
only epo ed endopa asi e is a ila ial
nema ode, Jose ila ia mac-
ke asae (Moo house
e al., 1979).
BEHAVIOR. Mac ode ma is inac i e du ing dayligh hou s
and lea es he oos om one o se e al hou s a e sunse . Some
indi iduals
may emain in he oos all nigh . They lea e singly, in
pai s, o in small g oups. Du ing ligh , hey hold he head high and
appea o scan he coun yside (Douglas, 1967). Hun ing beha io
began 3 o 5 h a e da k in a cap i e colony (Kulze
e al., 1984).
The ba s use a si -and-wai
hun ing s a egy. F om oos s, he ba s
look in he di ec ion o noises om p ey and di ec hei pinnae o
de ec mo emen . Hea ing is used ini ially
o de ec p ey, and pe ch
posi ion is al e ed occasionally o "lis en" o p ey om a di e en
angle. Echoloca ion emissions each a peak equency be o e he
ba lies and ho e s o e p ey. The p ecise loca ion o he p ey is
a ained h ough a pulse se ies emi ed du ing a a iable numbe o
econnoi e ing ligh s o e po en ial p ey. When loca ed, he ic im
is seized quickly om abo e, p essed o he g ound, and held by
he humb claws o he killing bi e in o he neck o head deli e ed
by he canine ee h. Small mammals a e manipula ed
by he humb
claws and ea en om head o ail (Guppy and Coles, 1983).
Cap i e M. gigas can cap u e p ey up o 80% o hei body
mass. When able o ca ch la ge p ey, Mac ode ma may no hun
daily (Kulze e al., 1984).
When ood is sca ce unde cap i e condi ions, wo o mo e
indi iduals
may igh o ob ain he same p ey i em. While igh ing,
hey emi high-pi ched squeals
(Douglas, 1967). They o en p oduce
a c icke -like chi p when hung y and he young con inuously chi p
when sepa a ed om he mo he . Usually, he ba s make no noise
a all (Douglas, 1967).
Human in ude s o a ca e colony may cause he g oup o
become ne ous and lea e inconspicuously (Douglas, 1967).
The ex en o daily and seasonal mo emen s is unknown, bu
Douglas
(1967) sugges ed ha he e we e di e ences in nigh oos -
ing beha io du ing we and d y seasons. Du ing he we season,
ba s a Kulumbu u
equen ed buildings
as eeding oos s, bu we e
no known o do so du ing he d y season.
GENETICS. No hing is known abou he gene ics o his
species.
REMARKS. The name Mac ode ma is om he G eek oo s
mak os, meaning la ge, and de ma, meaning skin. The speci ic
epi he gigas is G eek, meaning gian , and e e s o he ac ha
his is he la ges species in he amily.
LITERATURE CITED
ANDERSEN, K., AND R. C. WROUGHTON.
1907. On he ba s o
he amily Megade ma idae.
Ann. Mag. Na . His ., 19:129-
145.
BAYNES, A., D. MERRILEES,
AND
J. K. PORTER.
1976. Mammal
emains om he uppe le els o a la e Pleis ocene deposi in
De il's Lai , Wes e n Aus alia. J. Royal Sci. W. Aus alia,
58:97-126.
BREEDEN,
S., AND K. BREEDEN.
1967. Animals in eas e n Aus-
alia. Aus alasian Publ. Co., Sydney, 128 pp.
BRIDGE,
P. J. 1975. Paleo-dis ibu ion
o Mac ode ma gigas in
he sou hwes o Wes e n Aus alia. Helic i e, 13:35-36.
COOK,
D. L. 1960. Some mammal emains ound in ca es nea
Ma ga e Ri e . W. Aus alia
Na ., 7:107-108.
DOBSON,
G. E. 1880. On some new o a e species o Chi op e a
in he collec ion o he G6 ingen Museum. P oc. Zool. Soc.
London, 1880:461-462.
DOUGLAS,
A. M. 1962. Mac ode ma gigas sa u a a (Chi op e a,
Megade ma idae)
a new subspecies om he Kimbe ly
Di ision
o Wes e n Aus alia. W. Aus alia
Na ., 8:59-61.
. 1967. The na u al his o y o he ghos ba , Mac ode -
ma gigas, in Wes e n Aus alia. W. Aus alia
Na ., 10:125-
138.
FILEWOOD,
L. W. 1983. The possible occu ence in New Guinea
o he ghos ba (Mac ode ma gigas; Chi op e a, Megade -
ma idae). Aus alian
Mamm., 6:35-36.
FINLAYSON,
H. H. 1961. On cen al Aus alian mammals. Pa
IV- he dis ibu ion and s a us o cen al Aus alian
species.
Reco ds S. Aus alian
Mus., 14:141-191.
GUPPY, A., AND R. B. COLES. 1983. Feeding beha io o he
Aus alian
ghos ba , Mac ode ma gigas (Chi op e a:
Mega-
de ma idae)
in cap i i y. Aus alian
Mamm., 6:97-99.
HAMILTON-SMITH,
E. 1974. The p esen knowledge
o Aus alian
Chi op e a.
Aus alian
Mamm., 1:95-108.
1980. The s a us o Aus alian
Chi op e a. Pp. 199-
206, in P oceedings o he Fi h In e na ional Ba Resea ch
Con e ence (D. E. Wilson and A. L. Ga dne , eds.). Texas
Tech Uni . P ess, Lubbock, 434 pp.
3
MAMMALIAN SPECIES 260
HAND, S. J. 1985. New Miocene megade ma ids (Chi op e a:
Megade ma idae)
om Aus alia
wi h commen s on megade -
ma id phylogene ics. Aus alian
Mamm., 8:5-43.
HOOGSTRAAL,
H., D. E. MOORHOUSE,
G. WOLF, AND H. Y. WASSEF.
1977. Ba icks o he genus A gus (Ixodoidea: A gasidae).
A. (Ca ios) mac ode mae, new species om Queensland,
Aus-
alia. Ann. En omol. Soc. Ame ., 70:861-870.
KOOPMAN,
K. F. 1984a. Taxonomic and dis ibu ional no es on
opical Aus alian ba s. Ame . Mus. No i a es, 2778:1-48.
1984b. Ba s. Pp. 145-186, in O de s and amilies o
Recen Mammals o he wo ld
(S. Ande son and J. K. Jones,
J ., eds.). John Wiley and Sons, New Yo k, 686 pp.
KULZER, E., J. E. MCKEAN, AND F. P. MOEHRES.
1984. P ey
ca ching beha iou and echoloca ion in he Aus alian ghos
ba , Mac ode ma gigas (Chi op e a:
Megade ma idae).
Aus-
alian Mamm., 7:37-50.
LEITNER, P., AND
J. E. NELSON. 1967. Body empe a u e,
oxygen
consump ion and hea a e in he Aus alian alse ampi e
ba , Mac ode ma gigas. Comp. Biochem. Physiol., 21:65-
74.
LUNDELIUS,
E. 1960. Pos Pleis ocene aunal succession in Wes -
e n Aus alia and i s clima ic in e p e a ion. P oc. Twen y-
i s In e na l. Geol. Cong ., 4:142-153.
McKEAN, J. L., AND W. J. PRICE. 1967. No es on some Chi op-
e a om Queensland, Aus alia. Mammalia, 31:101-119.
MILLER,
G. S. 1906. Twel e new gene a o ba s. P oc. Biol. Soc.
Washing on, 19:83-85.
1907. The amilies and gene a o ba s. Bull. U.S. Na l.
Mus., 57:1-282.
MOLNAR,
R. E., L. S. HALL,
AND
J. H. MAHONEY.
1984. New
ossil locali ies o Mac ode ma Mille , 1906 (Chi op e a:
Megade ma idae)
in New Sou h Wales and i s pas and p esen
dis ibu ion in Aus alia. Aus alian
Mamm., 7:63-73.
MOORHOUSE,
D. E., O. BAIN, AND G. WOLF. 1979. Jose ila ia
macke asae gen. e sp. no . (Nema oda:Fila ioidea)
pa asi e
de la chau e-sou is Mac ode mc gigas Dobson. Ann. Pa a-
si ol. Hum. Comp., 54:645-651.
NOWAK,
R. M., AND
J. L. PARADISO. 1983. Walke 's mammals
o he Wo ld. Fou h ed. Johns Hopkins Uni . P ess, Bal i-
mo e, Ma yland, 1:1-568.
PIRLOT, P., AND
J.. NELSON. 1980. Quan i a i e
aspec s o b ain
mo phology
in Mac ode ma gigas (Megade ma idae,
Chi op-
e a). Aus alian
Mamm., 3:105-108.
STEPHAN, H., AND
J. E. NELSON. 1981. B ains o Aus alian Chi-
op e a I. Encephaliza ion
and mac omo phology.
Aus alian
J. Zool., 29:653-670.
STEPHAN, H., J. E. NELSON,
AND
H. D. FRAHM.
1981. B ain size
compa ison in Chi op e a. Z. Zool. Sys . E olu .- o sch, 19:
195-222.
TAYLOR,
J. M. 1984. The Ox o d
guide o mammals o Aus alia.
Ox o d Uni e si y P ess, New Yo k, 148 pp.
THORNBACK, J., AND
M. JENKINS. 1982. Mac ode ma gigas. Pp.
101-102, in The IUCN mammal ed da a book. Pa 1.
In e na l.
Union Conse .
Na u e, Gland, Swi ze land,
515 pp.
TROUGHTON,
E. L. G. 1926. The ba s o Aus alia and New
Guinea. Pp. 21-88, in The wild animals o Aus alasia
(A. S.
Le Soue and H. Bu ell, eds.). Geo ge C. Ha ap and Co.,
L d., Sydney, 388 pp.
VESTJENS,
W. J. M., AND L. S. HALL. 1977. S omach con en s
o o y- wo
species o ba s om he Aus alasian
egion. Aus-
alian Wildl. Res., 4:25-35.
WAITE,
E. R. 1900. Recu ence o Megade ma gigas, Dobson.
Reco ds S. Aus alian
Mus., 3:188-189.
WILSON, D. E. 1973. Ba aunas: a ophic compa ison, Sys .
Zool., 22:14-29.
Edi o s o his accoun we e B. J. VERTS
and SYDNEY ANDERSON.
Managing Edi o was TIMOTHY E. LAWLOR.
W. S. HUDSON AND D. E. WILSON,
WELLS
COLLEGE,
AURORA,
NEW
YORK
13026 AND
U. S. FISH AND WILDLIFE
SERVICE,
NATIONAL
MUSEUM OF NATURAL
HISTORY, WASHINGTON,
D.C. 20560.
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