MAMMALIAN SPECIES
Chae ephon pumilus.
No. 574, pp.
1-6,
3 igs.
By Syl ie Boucha d
Published 1
June
1998
by he Ame ican Socie y o Mammalogis s
Chae ephon
Dobson,
1874
Chae ephon Dobson, 1874:144. Type species Molossus (Nyc ino-
mus) joho ensis Dobson.
Nyc inomus Dobson, 1878:420.
Chae ephon Ande sen, 1907:35.
CONTEXT
AND CONTENT. O de Chi op e a, Subo de
Mic ochi op e a, Family Molossidae. The genus Chae ephon con-
ains 13 species: C. aloysiisabaudiae, C. anso gei, C. bemmelini,
C. bi i a a, C. chapini, C. gallaghe i, C. jobensis, C. joho ensis,
C. majo , C. nige iae, C. plica a, C. pumilus, and C. ussa a
(F eeman, 1981; Koopman, 1993, 1994). The ollowing key o spe-
cies is de i ed om cha ac e s p o ided in Co be and Hill (1992),
Ha ison (1975), Hayman and Hill (1971), Hill (1974):
1 Ea s conjoined 2
Ea s sepa a ed 6
2Ea s conjoined by iangula lappe o skin 3
Ea s conjoined by uns uc u ed band o skin 5
3 La ge pai ed nasal in la ions; leng h o o ea m
37-39
mm
____________________________________________________________________________
C. gallaghe i
No nasal in la ions; leng h o o ea m
42-50
mm 4
4 Colo uni o mly b own wi h whi e belly s ipe; lack o well
de ined in e au al pocke ; leng h o o ea m
42-44
mm
____________________________________________________________________________________
C.
majo
Colo blackish b own do sally and no whi e s ipe en al-
ly; well de ined in e au al pocke ; leng h o o ea m
46-
50 mm C. joho ensis
5 Colo uni o mly b own; leng h o o ea m
48-54
mm
_________________________________________________________________________________
C. jobensis
Colo da k b own do sally and ligh e en ally; leng h o
o ea m
40-50
mm C. plica a
6 Size la ge , leng h o o ea m
42-52
mm 7
Size smalle , leng h o o ea m 34--42 mm 12
7Pai ed caudal glands opening as sli s on ei he side o ail
oo below; leng h o o ea m
46-47
mm C. bemmelini
Wi hou such glands; leng h o o ea m
42-52
mm 8
8Whi e spo s, s ipes, o la e al
bands
9
Wi hou whi e ma ks 10
9 Colo umbe b own wi h a iable pa e n o whi e spo s o
sho s ipes a anged la e ally on c own and some imes
also on shoulde s and lanks; leng h o o ea m
46-51
mm C. bi i a a
Conspicuous whi e la e al band below, along lanks a
junc-
ion wi h memb ane, con as ing wi h e y da k b own
colo abo e and below; leng h o o ea m
44-50
mm
________________________________________________________________________________
C. nige iae
10 Colo umbe b own, op o head, neck, and h oa da ke ,
almos black; leng h o o ea m
46-48
mm C. anso gei
Colo uni o mly usse b own; leng h o o ea m
42-52
mm
_______________________________________________________________________________________________
11
11 Leng h o o ea m
42-46
mm C. ussa a
Leng h o o ea m
50-52
mm C. aloysiisabaudiae
12 Leng h o o ea m
34-39
mm; males wi h long
(13-14
mm)
bicolo ed pos au al c es (da k basally, pale apically);
ligh wings; colo below ligh wi h whi ish band down o
he mid-belly C. chapini
Leng h o o ea m
37-42
mm; males wi h sho (:510 mm)
unicolo ed pos au al c es ; wings da k o ligh ; colo be-
low a iable, wi h o wi hou whi e on he median a ea
_________________________________________________________________________________
C.
pumilus
Chae ephon
pumilus
(C e zschma ,
1826)
Li le F ee- ailed Ba
Dysopes
pumilus
C e zschma , 1826:69. Type locali y "Massawa,"
E i ea.
Dysopes limba us Pe e s, 1852:56. Type locali y "Mozambique Is-
land" (15°S, 40
042'E),
Mozambique.
Nyc inomus leucogas e G andidie , 1869:337. Type locali y "Ma-
hab," nea Menabe, eas o Mo onda a, Madagasca .
Nyc inomus
gambianus
De Win on, 1901:39. Type locali y Gam-
bia.
Nyc inomus pusillus Mille , 1902:245. Type locali y, Aldab a Is-
land.
Nyc inomus
hindei
Thomas, 1904:210. Type locali y "Fo Hall,"
Kenya.
Chae ephon webs e i Dollman, 1908:546. Type locali y "Yola,"
no he n Nige ia.
Chae ephon
pumilus
nai ashae Hollis e , 1916:4. Type locali y
"Nai asha S a ion," Kenya.
Chae ephon
a e
Allen e al., 1917:456. Type locali y "Malela,"
nea Boma, Bas Zai e, Zai e.
Chae ephon (Lophomops) c is a us Allen e al., 1917:463. Type
locali y "Boma," Bas Zai e, Zai e.
Chae ophon (sic.)
pumilus
elphicki Robe s, 1926:245. Type lo-
cali y "Malelane Es a e," Ba be on dis ic , sou heas e n
T ans aal, Sou h A ica.
Chae ophon (sic.) (Lophomops) nig i Ha , 1928:53. Type locali y
"Bou em," Timbue u dis ic , Mali.
Chae ephon (Lophomops)
langi
Robe s, 1932: 17. Type locali y
"Tso so oga Pan," no he n Bo swana.
Tada ida (Chae ephon)
aini
Hayman, 1951:82. Type locali y
"Wago o es ," Hau Zai e, Zai e.
CONTEXT
AND CONTENT. Con ex no ed in gene ic
summa y abo e. No subspecies a e a ibu ed o C.
pumilus
due o
i s high dich oma ism (Ansell, 1986; Hayman and Hill, 1971). Pe-
e son e al. (1995) di ides C.
pumilus
in o ou species: Tada ida
leucogas e , T. limba a, T. nai ashae and T. pumila.
DIAGNOSIS. Chae ephon
pumilus
can be dis inguished
om o he membe s o he genus by lack o lobe p ojec ing be ween
he inne bases o ea s, small size, leng h o o ea m
37-42
mm,
and male's unicolo pos au al c es , which measu es :510 mm (Fig.
1; Hayman and Hill, 1971). The ollowing skull cha ac e is ics also
dis inguish his species: educed an e io pala al ema gina ion
which does no ex end behind
uppe
inciso s and is sepa a ed om
incisi e o amina by a bony ba ; hi d commissu es o he las
uppe
mola s well de eloped and almos as long as he second commis-
su e; basisphenoid pi s shallow and sepa a ed a mos by a low
idge; o ehead dis inc ly ele a ed; leng h o maxilla y oo h ow
<6.3
mm; in e den al pala e
30%
sho e in wid h han in leng h
(Fig. 2; Mees e e al., 1986).
FIG. 1. Male Chae ephon
pumilus
om Sou h A ica.
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2MAMMALIAN SPECIES 574
40
....
30
-
20-
20
10
0
10
20
30
(--J'
, , ,
~~'"
')'
<>=e'
-30
~
-20
20
10
oI
10
I
20
,
30
I
40
I
50
FIG. 2. Do sal, en al, and la e al iews o c anium and la -
e al iew o mandible o Chae ephon
pumilus
om Bo swana
(20
000'S,
23°25'E;
male, Royal On a io Museum #62 639). G ea -
es leng h o c anium is 17.6 mm.
GENERAL CHARACTERS. Chae ephon
pumilus
is one o
he smalles species wi hin he genus Chae ephon, wi h a o al
leng h om 54 o 102 mm (Kingdon, 1974; Smi he s, 1983). The
do sal u is sho blackish-b own and he en al u is ligh e .
Whi e o pale hai s a e p esen on he en al su ace whe e wings
join lanks, and da k-winged and whi e-winged o ms ha e
been
eco ded (Happold, 1987; Hayman and Hill, 1971; Kingdon, 1974;
Smi he s, 1983). A inge o pale b is les is p esen on he ou e -
mos oes (Kingdon, 1974). The ea s a e ounded and la ge o he
size o he head. The an i agus is la ge and conceals he iny agus
which is asymme ically bilobed a he ip (Smi he s, 1983). Males
ha e a o ehead u ha makes
hei
silhoue e ecognizable in
ligh (Kock, 1969).
Ranges o c anial measu emen s (in mm) in Uppe Vol a a e
as ollows: o al leng h,
15.2-16.2;
zygoma ic b ead h,
9.1-9.8;
mas oid b ead h,
9.0-9.2;
leng h o mandibula oo h ow,
5.3-5.8;
M3-M3,
6.1-7.2;
leng h o maxilla y oo h ow,
9.9-10.5
(Poche,
1975). Means and anges o ex e nal measu emen (in mm) o 10
males and 20 emales, espec i ely, om Zimbabwe a e as ollows:
FIG. 3. Dis ibu ion o Chae ephon
pumilus
in A ica and
he
nea
eas .
o al leng h, 93
(90-98),
95 (90-100); leng h o ail, 36
(35-39),
35 (31-39); leng h o o ea m, 38
(37-39),
38 (37-39). A e age
and anges o mass (in g) o he same specimens a e 11.1
(10.1-
12.7), 11.3 (10.1-14.8) and o g a id emales, 12.5
(11.2-15.1,
n
=
19-5mi he s,
1983). In Yemen, leng h o
ea
is 12 mm, agus
2.5 mm, and he wingspan anges om
250
mm o
260
mm (n
~
3-AI-Sa adi,
1991).
DISTRIBUTION.
Chae ephon
pumilus
occu s om Senegal
o Yemen, sou h o Sou h A ica, Bioko,
Pemba
and Zanziba , Com-
o o Isls, Aldab a and Ami an e IsIs (Seychelles), and Madagasca
(Fig. 3; Koopman, 1993). No ossils o his species a e known.
FORM
AND FUNCTION. The den al o mula o he li le
ee- ailed ba is i 1/2, c 1/1, p 2/2, m 3/3, o al 30 (Pe e s, 1852).
Ju eniles ha e ou milk ee h. The e eb al column has 7 C, 13
T, 6 L, 5 S, and 10 Ca, o al 41. The keel o he s e num is weak
and a icula es wi h six pai s o ibs. The second inge has one
phalange, he hi d inge has h ee, and he ou h and i h also
ha e h ee phalanges, bu he las digi is sho and ben (Pe e s,
1852).
The an e io po ion o he ongue has la , scale-like papillae,
whe eas he pos e io po ion has la ge , mo e widely spaced, and
wa -shaped papillae. Two sali a y glands, ca. 7 mm in diame e ,
lie on
each
side o he neck. The la ynx is ca. 5 mm long. The le
lung has a single lobe and he igh lung has wo small lobes. The
hea is ca. 10 mm long and 5 mm wide. The esophagus opens
nea
he diaph agm and in o a la ge saclike s omach which con in-
ues in o an alimen a y canal which is ca. 130 mm wide. The li e
has a weak inden a ion in he middle and a
deep
inden a ion on
he igh side whe e i s g ea es mass lies. The gall
bladde
is
loca ed in he
deep
inden a ion o he li e . The spleen is ca. 15
mm long and i s g ea es wid h is ca. 4 mm. Kidneys a e
bean-
shaped
and measu e ca. 6 mm long and 4 mm wide. The igh
kidney lies much
deepe
han
he le one. C.
pumilus
has a hin-
walled u ina y
bladde
which in males is enci cled om i s neck
a ound bo h sides by he p os a e glands (Pe e s, 1852).
The penis o he li le ee- ailed ba is so and does no ha e
abaculum. Tes es a e long (ca. 4 mm), a e lodged beside he oo
o he penis, and p o ude in o he
skin
(Pe e s, 1852). Combined
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MAMMALIAN SPECIES
574
weigh o es es in adul males does no show seasonal a ia ion in
Uganda, bu emains cons an h oughou he yea a ca. 0.1 g (Mu-
e e, 1973). Adul males possess ma u e epididymidal spe ma ozoa
all yea , bu hey do no unde go con inuous spe ma ogenesis (Ma -
shall and Co be , 1959).
Females ha e a bico nua e u e us. No sign o a ophica ion is
p esen
in he le ho n o nonp egnan indi iduals, and he size
and
appea ance
o bo h ho ns a e simila ; howe e , implan a ion
occu s almos unila e ally in he igh ho n (Mu e e, 1973). His o-
logical examina ion o a p egnan and lac a ing emale showed
mamma y glands ha we e g ea ly hype ophied, measu ing 14
mm by 11 mm in diame e and ex ending o wi hin 5 mm o he
spine
pos e io ly. Many glandula acini we e ully
dis ended
wi h
milk and had a lining epi helium ha appea ed la .
O he
acini
we e emp y and possessed sec e o y columna epi helium whe e he
dis al
pa
o he epi helial cells con ained
ei he
globules o se-
c e ion o we e b oken o . The igh ho n o he u e us was swollen
due o an ea ly p egnancy. A a c own- ump leng h o 3.3 mm, limb
buds
o he emb yo we e
jus
de eloped and sagi al sec ions e-
ealed ha di e en ia ion o o eb ain, ongue, hea , and li e we e
comple e (Mu e e, 1973).
Specimens o
ei he
sex o he li le ee- ailed ba collec ed
in Kenya p io o he d y season ha e la ge quan i ies o subcu a-
neous a . Females ha e hea y a deposi s du ing bo h ges a ion
and lac a ion (O'Shea and Vaughan, 1980). In cap i i y li le ee-
ailed ba s became o pid du ing he day a
21-24°C
(Happold and
Happold, 1988).
Chae ephon
pumilus
is
adap ed
o as ligh in open a eas.
I uses high in ensi y, shallow equency-modula ed calls o low
equency and long du a ion. These calls, al hough suscep ible o
in e e ence om clu e , a e ideally sui ed o long- ange de ec ion
o p ey in open a eas. C.
pumilus
has na ow, long- ip wings o
i s size, a high aspec a io (8.90), a high wing loading (12.6 N
m"),
and a low wing ip
shape
index (1.2) which indica es a low
le el o maneu e abili y (Ald idge and Rau enbach, 1987; Findley
e al., 1972; O'Shea and Vaughan, 1980; Vaughan, 1966). The min-
imum dis ance be ween s ings (moun ed in he labo a o y) ha C.
pumilus
can nego ia e is 44 em, whe eas o he A ican ba s
can
ly
be ween s ings 11 e n apa (Ald idge and Rau enbach, 1987). The
ligh memb ane is s ong, lea he y, and elas ic. Ea s a e b oad
ela i e o hei leng h and lie agains he
head
when in ligh o
educe
d ag. Ai oils o high
cambe
a e o med by connec i e is-
sue ha b aces an e io and en al bo de s o he pinnae. Weigh
o he
head
is compensa ed by ca ying he occipi al po ion agains
he in e scapula dep ession, placing he
head
close o he body's
cen e
o g a i y. The w inkled lips
sp ead
ou wa d when in ligh
and inc ease he a ea a ailable o ca ching insec s. This may com-
pensa e
o lack o maneu e abili y du ing as ligh (Vaughan,
1966). Roos ing ba s sho en
hei
wings by olding hem. Join s o
he hi d inge lex upon one ano he and he i s join o a es
conside ably on he me aca pal, in
o de
ha he old may be
ucked sideways
benea h
he wing (Rose ea , 1965).
ONTOGENY AND REPRODUCTION. Chae ephonpum-
ilus is polyes ous wi h emales expe iencing pos pa um es us
(Happold and Happold, 1989; McWilliam, 1987; an
de
Me we e
al., 1986, 1987). Females become sexually ma u e in he b eeding
season ollowing hei bi h, bu he age a which his occu s a ies
wi h loca ion. In he T ans aal, sexual ma u i y is a ained a
5-12
mon hs ( an
de
Me we e al., 1986). McWilliam (1987) epo ed
a3-mon h-old sexually ma u e emale in Ghana ha ga e bi h a
5mon hs. In Malawi, sexual ma u i y in emales is a ained a
6-
10 mon hs (Happold and Happold, 1989). Sexual ma u i y in ou
banded
males occu ed ca. 5 mon hs in Ghana (McWilliam, 1987),
whe eas
da a
a e inconclusi e in Malawi (Happold and Happold,
1989).
P egnancies ha e
been
epo ed a a ious imes h oughou
he yea , iming ha may be a unc ion o ain all pa e ns (Happold
and Happold, 1989). S udies in Bu kina Faso, Nige , Nige ia, and
Sudan
(all no h o
lOON
la i ude) epo collec ion o p egnan e-
males o emales simul aneously p egnan and lac a ing a di e en
imes o he yea : in Nige (Poche, 1975) and Nige ia (Ha ison,
1958), one p egnan and one p egnan and lac a ing emale, e-
spec i ely, in May; in Bu kina Faso (Koopman e al., 1978), one
p egan emale a he end o Augus ; in Sudan (Kock, 1969), 23
p egnan emales ou o 32 emales wi h imma u es in Oc obe .
Naked young we e ound in
Sudan
in June. Rainy season in hese
3
coun ies begins in May and
ends
in Sep embe , excep o
Sudan
whe e i s a s and
ends
one mon h la e .
Be ween looN and 10
0S,
eco ds o p egnancies and pa u i-
ion a e as ollows: in May, one lac a ing
and
p egnan emale in
Somalia (Ha ison, 1958); in No embe and Ma ch-Ap il, pa u i-
ion in
Kenya- imes
which co espond o he pe iods o
peak
ain-
all (O'Shea and Vaughan, 1980); in No embe , Janua y, and
June,
>80%
o emales p egnan in Uganda (Mu e e, 1973). Females
om Uganda a e polyes us and in ep oduc i e synch ony (Ma -
shall and Co be , 1959). In Uganda ain occu s h oughou he yea
wi h a small
peak
in Oc obe and No embe and a la ge
peak
in
Ap il and May (Happold and Happold, 1989). In Ghana mos e-
males gi e bi h a leas h ee imes a yea , mos ly du ing he we
season, which las s om Ap il o Oc obe ; synch onized peaks oc-
cu
in June, Augus , and ea ly No embe . The cap u e o a ju enile
and a emale ha was simul aneously lac a ing and p egnan in
Janua y du ing he d y season sugges s ha pa ous emales could
po en ially gi e bi h o i e young
pe
yea , al hough he ex en o
b eeding occu ing ou side he we season is s ill unknown
(McWilliam, 1987).
Sou h o 10
0S
la i ude, eco ds o ep oduc i e indi iduals in-
clude he ollowing: in Mozambique (Law ence and Lo e idge,
1953), h ee emales wi h la ge emb yos in Janua y; in Zimbabwe
(Smi he s and Wilson, 1979), p egnan emales in Ap il; and in
Bo swana (Smi he s, 1971), ap egnan emale in Augus , 47 p eg-
nan emales om Oc obe o No embe , and hai less young in
No embe and Feb ua y. The we season in hese h ee coun ies
las s om No embe o Ma ch. In Zambia p egnan emales we e
caugh in Sep embe , No embe , and Decembe ; bo h p egnan and
lac a ing o only lac a ing emales in Decembe , Feb ua y, and
Ma ch; and sexually inac i e emales in Ap il. F om hese da a,
Ansell (1986) concluded ha h ee bi hs
pe
emale ake place
du ing he ainy season, which las s om No embe o Ap il. P eg-
nancies
occu
om Augus o Ap il in K uge Na ional Pa k, Sou h
A ica. Rain he e s a s in Oc obe and
ends
in Ma ch; howe e ,
minimum empe a u e may play a mo e impo an ole in de ining
he b eeding season han ain all,
because
i di ec ly a ec s insec
abundance
a he highe al i udes ound in he pa k ( an
de
Me we
e al., 1986, 1987).
The li le ee- ailed ba is a mono ocous species, wi h emb y-
os almos always occu ing in he igh
hom
o he u e us (Happold
and Happold, 1989; McWilliam, 1987; Mu e e, 1973; an
de
Me -
we e al., 1986, 1987), al hough winning has
been
epo ed on one
occasion (McWilliam, 1987). Ges a ion las s o ca. 60 days in
Sou h A ica ( an
de
Me we e al., 1986) and Malawi (Happold
and Happold, 1989) and
67-72
days in Ghana (McWilliam, 1987).
Fe al weigh s ange om 0.1 g o 3.5 g, he
la e
being hose close
o he ime o pa u i ion (Mu e e, 1973). Young a e bo n naked
(Kock, 1969). Lac a ion las s
2-3
weeks (Happold and Happold,
1989; Mu e e, 1973; an
de
Me we e al., 1986, 1987). Al hough
he possibili y o i e bi hs/yea has been sugges ed, emales gen-
e ally gi e bi h h ee imes, wi h synch oniza ion dec easing
each
ime (McWilliam, 1987; Mu e e, 1973; an
de
Me we e al., 1986).
ECOLOGY. The li le ee- ailed ba is ound in a ied hab-
i a om semia id egions in he no h o clea ed ain o es egions
in he sou h (Happold, 1987). These ba s
occu
widely in sa annas
o Guinea and Sudan and in o es s o he Congo basin in Zai e
(Rose ea , 1965). They a e ound in sa anna woodlands in Zambia,
and in he Cape P o ince hey
occu
in moun ainous a eas in he
Cape Macchia Zone, as well as in mo e a id open coun y u he
no h. They show a
clea
p e e ence in Zimbabwe o low eld
condi ions and a e ne e
aken
on he pla eau o e he
1,000
m
le el (Smi he s, 1983).
Na u al oos s o C.
pumilus
a e hollows and c e ices in ees,
c owns o a ious species o palm (Kingdon, 1974), and ocky en-
i onmen s wi h an
abundance
o c e ices in which colonies can
ake e uge in la ge
numbe
(Pienaa e al., 1987). In addi ion, C.
pumilus
is o en ound in human dwellings, mainly
unde
ha ched
o co uga ed i on oo s (Happold, 1987; Mu e e, 1973;
Pienaa
e
al., 1987; an
de
Me we, 1986, 1987). Condi ions o a oos
unde
a oo in Uganda a ied o e he cou se o 1 yea om 18°C o
50°C and om
35%
o
90%
ela i e humidi y (Mu e e, 1973). In
Sudan li le ee- ailed ba s p e e he no h side o a building when
i is shady and p o ec ed om he wind (Kock, 1969).
Size o he oos ing colony a ies om a ew indi iduals
(5-
20) o
hund eds
(Happold and Happold, 1989; Kingdon, 1974;
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4
Kock, 1969; Mu e e, 1973; Smi he s, 1971; Smi he s and Wilson,
1979; Ve schu en, 1957). La ge colonies oos ing in buildings a e
o en conside ed anuisance because o he smell o hei d oppings
(Ma shall and Co be , 1959). Seg ega ion acco ding o age o sex
does no occu , as adul s o bo h sexes and ju eniles a e ound
oge he (Happold and Happold, 1989; Ma shall and Co be , 1959;
Mu e e, 1973).
Chae ephon
pumilus
o en oos s wi h a la ge molossid, Mops
condylu us (Happold, 1987; Happold e al., 1987; Mu e e, 1973;
O'Shea and Vaughan, 1980).
When
oos ing in ees, C.
pumilus
can be ound wi h he palm swi , Cypsiu us pa us, which eeds
on he same insec s and like C.
pumilus
alls p ey o he ba -hawk
(Kingdon, 1974).
The li le ee- ailed ba is insec i o ous, ea ing a wide a ie y
o small, so -bodied insec s (Kingdon, 1974) ha a e consumed in
ligh (Pienaa e al., 1987). Lygaeidae (Hemip e a) cons i u es
45%
o he die o ba s cap u ed in Sep embe -Oc obe in Kenya, while
he emainde o he die is dis ibu ed among he ollowing: Co-
leop e a, 18%; Lepidop e a, 13%; O hop e a, 8%; Dip e a, 4%;
o he Hemip e a (Pen a omidae), 3%; Homop e a, 1%; Hymenop-
e a, 0.5%; Neu op e a, 0.5%; uniden i ied, 5% (Whi ake and
Mum o d, 1978). Al hough insec a ailabili y is simila du ing d y
and we seasons in Sou h A ica, mean p ey size o Chae ephon
pumilus
inc eases om
6.8
mm du ing he d y season o 8.5 mm
in he ainy season (Ellis, 1995). One C.
pumilus
oos ing in a
building had appa en ly ed upon an s, as 21 heads we e ound
a ached o a ious pa s o he ba 's body (Whi ake and Mum o d,
1978). Al hough C.
pumilus
and Mops condylu us a e sympa ic
and o en oos oge he , compe i ion o esou ces is limi ed be-
cause he la e consumes la ge p ey i ems wi h ha d in egumen s
(F eeman, 1981; Whi ake and Mum o d, 1978).
S omachs o ju eniles con ain milk and emains o la ge cock-
oaches (Leucophaea), which ha e no
been
epo ed in adul s. The
cock oaches can a ain leng hs o 60 mm and a e common in ceil-
ings in which he young hang (Ma shall and Co be , 1959).
A equi emen o d inking wa e in his species is hough o
be minimal as h ee indi iduals ha we e kep in cap i i y o
10-
60 days we e ne e obse ed o d ink. One emale unde wen p eg-
nancy and lac a ion wi hou d inking (Happold e al., 1987).
The ba -hawk (Machei amphus alcinus), hobby alcon (Falco
subbu eo), Wahlbe g's eagle (Aqui a wahlbe gi),and he A ican
goshawk (Accipi e achi o) egula ly p ey upon C.
pumilus
(Fen on
e al., 1994; Kingdon, 1974;
Pienaa
e al., 1987; Rau enbach e
al., 1990). Li le ee- ailed ba s also occasionally all p ey o he
black ki e, Mil us mig ans (McWilliam, 1989). One A ican gos-
hawk pai ed upon a la ge colony o C.
pumilus
oos ing
unde
a
b idge in summe in Sou h A ica. Cap u e a e was
1-5
ba s
pe
bi d
pe
e ening. No p eda ion on he same colony was obse ed
du ing win e when eme gence had d opped o
<5%
o ha o
summe (Rau enbach e al., 1990). Ec opa asi es o C.
pumilus
include he mi e Chelanyssus ae hiopicus (Mac onyssidae; Whi-
ake and Mum o d, 1978) and he lea A aeopysylla
aini
(Is-
chnopsyllidae; Beauco nu, 1981).
BEHAVIOR. Chae ephon
pumilus
is a g ega ious species.
Colonies become e y noisy and es less, squeaking and jockeying
o posi ion, 1 hou be o e indi iduals lea e he oos (Smi he s,
1983). Li le ee- ailed ba s lea e he oos in a line in small
g oups a e sunse (Kingdon, 1974). In Uganda, ime o eme gence
is no ela ed o seasonal a ia ion in ligh in ensi y bu a he o
sunse , he ba s lea ing
31-32
min a e sunse h oughou he yea
(Ma shall and Co be , 1959). In con as , eme gence beha io in
Ghana is in luenced by en i onmen al, social, and ep oduc i e ac-
o s (McWilliam, 1989). The ba s s a o aging ea lie on o e cas
e enings han on
clea
o moonli nigh s. Ha em males usually
come ou i s , which could e lec he isk o a acks associa ed
wi h i s eme gence. Females eme ge ea lie du ing he i s 3
weeks o lac a ion han du ing hei p egnancies, p obably in e-
sponse o highe ene gy demands. The p esence o a ap o modi ies
eme gence beha io as ba s inc ease synch onici y and depa in
all di ec ions (McWilliam, 1989).
Chae ephon
pumilus
o ages singly and ligh is as and e -
a ic. I hun s high abo e g ound (2:70 m) in open sky, bu also
close o he g ound, making
sudden
swoops o wi hin 3 m o i
(Kingdon, 1974; Smi he s, 1971, 1983). This species is ac i e
h oughou he nigh wi h g ea es ligh ac i i y du ing he hou
ollowing sunse , a e which ac i i y d ops o a low a ound mid-
MAMMALIAN SPECIES
574
nigh ollowed by an e en lowe le el be o e sunse (Mu e e, 1969).
Females a e p edominan ly caugh du ing he i s hou a e sunse
(Happold e al., 1987; Mu e e, 1969).
Recap u e a es indica e ha li le ee- ailed ba s a e capable
o lea ning. Ra es o cap u e dec ease signi ican ly om he i s o
second nigh i ne ing si es a e no a ied, sugges ing ha eleased
indi iduals emembe he placemen o a ap. Li le ee- ailed ba s
also show homing abili ies, as specimens eleased 70 km away om
poin o cap u e we e ecap u ed a he o iginal si e 1 h la e (Mu-
e e, 1969).
In Ghana sex a ios in se en ha ems o C.
pumilus
ound
unde
oo s a ied be ween 1:3 (male: pa ous emale) and 1:21;
young we e also p esen (McWilliam, 1988). The hea ies males
a e associa ed wi h he la ges ha ems. Roos ideli y is obse ed,
as nine ou o 13 emales om one ha em we e ound oge he o
>1 yea . Few mo emen s a e made be ween adjacen oos s. Dis-
u bed colonies show g oup cohesion; se en o eigh indi iduals
om he same oos we e ecap u ed oge he a e
hei
p e ious
oos was in aded by an s.
The
same long e m oos ing associa ion
occu s in Sou h A ica; colonies make no seasonal o local mig a-
ions (Pienaa e al., 1987). In Ghana some emales a e ec ui ed
in o hei na al g oup, al hough mos young lea e wi hin hei i s
yea ollowing sexual ma u i y. The ma ing sys em o C.
pumilus
is
based on emale-de ense polygyny, al hough some esou ce de ense
migh be in ol ed as he numbe o oos s a ailable becomes lim-
i ing (McWilliam, 1988).
Chae ephon
pumilus
is a ac ed o wa e holes, especially
du ing he d y season. Indi iduals ha all in o he wa e do no
d own bu swim o sa e y (Ve schu en, 1976).
GENETICS. The ka yo ype o C.
pumilus
exhibi s adiploid
numbe o 48 and a undamen al numbe o 58. The ka yo ype
shows one la ge and wo medium me acen ic ch omosomes, h ee
medium sub elocen ic ch omosomes, and 17 medium-small ac o-
cen ic ch omosomes. The Xch omosome is subme acen ic, whe e-
as he Y is ac ocen ic (Smi h e a1. 1986). C.
pumilus
is no known
o hyb idize wi h o he species.
REMARKS. Chae ephon was o me ly included in Tada ida
as a subgenus, as skull cha ac e s used o dis inguish i om Ta-
da ida we e no cons an (Simpson, 1945; Ta e, 1941). Chae ephon
has been e-ele a ed o he le el o genus by F eeman (1981:60,
133, 150), bu o he au ho s do no ag ee (Co be and Hill, 1986;
Legend e, 1984; Mees e e al., 1986). Mos ecen ly, Pe e son e
a1. (1995) do no e ain Chae ephon as he genus because some
species sha e cha ac e s om bo h he Chae ephon and Mops
g oups (J. L. Ege , pe s. comm.). Ne e heless, Chae ephon has
been he e e ained (Koopman, 1993). The speci ic name
pumila
was changed o
pumilus
o ma ch he masculine gene ic name,
Chae ephon (K. F. Koopman, pe s. comm.). The gene ic name
Chae ephon e e s o he iend o Soc a es who is depic ed as
"Chae ephon he
ba "
in A is ophanes' comedy, A es (Dobson,
1874). The species name
pumilus
comes om he La in, meaning
"a dwa ."
LITERATURE CITED
ALDRIDGE,
H. D. J. N.,
AND
I. L.
RAUTENBACH.
1987. Mo phology,
echoloca ion and esou ce pa i ioning in insec i o ous ba s.
Jou nal o Animal Ecology,
56:763-778.
ALLEN,
J. A., H.
LANG,
AND
J. P.
CHAPIN.
1917. The Ame ican
Museum Congo expedi ion collec ion o ba s. Bulle in o he
Ame ican Museum o Na u al His o y,
37:405-478.
AL-SAF
ADI,
M. M. 1991. Chi op e an auna o Yemen A ab Re-
public. Mammalia,
55:269-274.
ANDERSEN,
K. 1907. Chi op e an no es. Annali del Museo ci ico
di s o ia na u ale Giacomo Do ia,
3:5-45.
ANSELL,
W. F. H. 1986. Some Chi op e a om sou h-cen al A -
ica. Mammalia,
50:507-519.
BEAUCORNU,
J. C. 1981. A new lea om Rwanda. Re ue Zoo-
logique A icaine, 95:
195-199.
CORBET,
G. B.,
AND
J. E.
HILL.
1986. A wo ld lis o mammalian
species. Second ed. Fac s on File Publica ions, New Yo k, 254
pp.
.1992. The mammals o he Indomalayan egion: a sys-
ema ic e iew. Ox o d Uni e si y P ess,
Ox o d,488
pp.
CRETZSCHMAR,
P. J. 1826. SaUge hie e, in A las zu
de
Reise im
no dlichen A ika (Edua d Huppell, ed.). Senckenbe gische
na u o schende Gesellscha , F ank u am Main, 78 pp.
Downloaded om h ps://academic.oup.com/mspecies/a icle/doi/10.2307/3504344/2600749 by gues on 30 Ap il 2024
MAMMALIAN SPECIES 574
DE
WINTON,
W. E. 1901. No es on ba s o he genus Nyc inomus.
Annals and Magazine o Na u al His o y, se e 7, 7:36-42.
DOBSON,
G. E. 1874. Desc ip ion o a new species o Chi op e a
om India and Yunan. Jou nal o he Asia ic Socie y o Ben-
gal, 43:237-238.
· 1878. Ca alogue o he Chi op e a in he collec ion o
he B i ish Museum. Second ed. B i ish Museum (Na u al His-
o y), London, 567 pp.
DOLLMAN,
G. 1908. On a collec ion o ba s om Yola, No he n
Nige ia collec ed by M . G. W. Webs e . Annals and Magazine
o Na u al His o y, se ies 8, 2:545-547.
ELLIS,
S. E. 1995. Seasonal esponses o Sou h A ican ba s o
insec densi ies and ligh s. M. S. hesis, Yo k Uni e si y, No h
Yo k, Canada, 47 pp.
FENTON,
M. B., I. L.
RAUTENBACH,
S. E.
SMITH,
C. M.
SWANEPOEL,
J.
GROSELL,
AND
J.
VAN
J
AARSVELD.
1994. Rap o s and ba s:
h ea s and oppo uni ies. Animal Beha io ,
48:9-18.
FINDLEY,
J. S., E. H.
STUDIER,
AND
D. E.
WILSON.
1972. Mo -
phologic p ope ies o ba wings. Jou nal o Mammalogy, 53:
429-444.
FREEMAN,
P. W1981. A mul i a ia e s udy o he amily Molos-
sidae (Mammalia, Chi op e a): mo phology, ecology, e olu ion.
Fieldiana Zoology, 7:
1-173.
GRANDIDIER,
A. 1869. 1. T a aux inedi s, Desc ip ion de quel-
ques animaux nou eaux decou e e, pendan I'annee 1869, su
la co e oues de Madagasca . Re ue e Magasin de Zoologie
Pu e e Appliquee,
21:337-341.
HAPPOLD,
D. C. D. 1987. The mammals o Nige ia. Ox o d Uni-
e si y P ess, Ox o d, 402 pp.
HAPPOLD,
D. C. D.,
AND
M.
HAPPOLD.
1988. Renal o m and
unc ion in ela ion o he ecology o ba s (Chi op e a) o Ma-
lawi. Jou nal o Zoology (London), 215:629-655.
· 1989. Rep oduc ion o Angola ee- ailed ba s (Tada ida
condylu a) and li le ee- ailed ba s (Tada ida pumila) in
Malawi (Cen al A ica) and elsewhe e in A ica. Jou nal o
Rep oduc ion and Fe ili y,
85:133-149.
HAPPOLD,
D. C. D., M.
HAPPOLD,
AND
1. E.
HILL.
1987. The ba s
o Malawi. Mammalia,
51:337-414.
HARRISON,
D. L. 1958. A no e on successi e p egnancies in an
A ican ba (Tada ida
pumila
webs e i Dollman). Mammalia,
22:592-595.
· 1975. A new species o A ican ee- ailed ba (Chi op-
e a: Molossidae) ob ained om he Zai e i e expedi ion.
Mammalia,
39:313-318.
HATT,
R. T. 1928. No e su un chei op e e nou eau du Soudan
F ancais, Bulle in de la Socie e Zoologique de F ance, 53:
374-376.
HAYMAN,
R. W. 1951. A new A ican molossid ba . Re ue de
Zoologie e de Bo anique A icaines, 45:82-83.
HAYMAN,
R. W.,
AND
J. E.
HILL.
1971. Pa 2: O de Chi op e a.
Pp.
61-64,
in The mammals o A ica: an iden i ica ion man-
ual (J. Mees e and H. W. Se ze , eds.). Smi hsonian Ins i u ion
P ess, Washing on, Dis ic o Columbia, 73 pp.
HILL,
J. E. 1974. New eco ds o ba s om sou h-eas e n Asia,
wi h axonomic no es. Bulle in o he B i ish Museum o Na -
u al His o y (Zoology),
27:127-138.
HOLLISTER,
N. 1916. Desc ip ion o a new genus and eigh new
species and subspecies o A ican mammals. Smi hsonian Mis-
cellaneous Collec ions, 66:
1-8.
KINGDON,
J. 1974. Eas A ican mammals, an a las o e olu ion
in A ica. Volume IIA. Academic P ess, London, 341 pp.
KOCK,
D. 1969. Die Flede maus-Fauna des Sudan. Abhandlun-
gen de Senckenbe gischen Na u o schenden Gesellscha ,
521:1-238.
KOOPMAN,
K. F. 1993. O de Chi op e a. Pp.
137-241,
in Mam-
mal species o he wo ld: a axonomic and geog aphic e e -
ence. Second ed. (D. E. Wilson and D. M. Reede , eds.).
Smi hsonian Ins i u ion P ess, Washing on, Dis ic o Colum-
bia, 1206 pp.
· 1994. Chi op e a: sys ema ics. Wal e de G uy e , Be lin,
217 pp.
KOOPMAN,
K. F., R. E.
MUMFORD,
AND
J. F.
HEISTERBERG.
1978.
Ba eco ds om Uppe Vol a, Wes A ica. Ame ican Museum
No i a es,
2463:1-6.
LAWRENCE,
B.,
AND
A.
LOVERIDGE.
1953. Zoological esul s o a
i h expedi ion o Eas A ica: 1. Mammals om Nyasaland
5
and Te e. Bulle in o he Museum o Compa a i e Zoology,
Ha a d, 110:1-80.
LEGENDRE,
S. 1984. E ude odon ologique des ep esen an s ac-
uels du g oupe Tada ida, (Chi op e a: Molossidae). Impli-
ca ions phylogene iques, sys ema iques e zoogeog aphiques.
Re ue Suisse de Zoologie,
91:3~9-442.
MARSHALL,
A. J.,
AND
P. S.
CORBET.
1959. The b eeding biology
o equa o ial e eb a es. Rep oduc ion o he ba Chae ephon
hindei a la i udes 0°26' N. P oceedings o he Zoological So-
cie y o London, 132:607-616.
MCWILLIAM,
A. N. 1987. Polyoes y and pos pa um oes us in
Tada ida (Chae ephon)
pumila
(Chi op e a: Molossidae) in
no he n Ghana, Wes A ica. Jou nal o Zoology (London),
213:735-768.
· 1988. Social o ganiza ion o he ba Tada ida (Chae e-
phon)
pumila
(Chi op e a: Molossidae) in Ghana, Wes A ica.
E hology, 77:115-124.
· 1989. Eme gence beha iou o he ba Tada ida (Chae-
ephon)
pumila
(Chi op e a: Molossidae) in Ghana, Wes A -
ica. Jou nal o Zoology (London),
219:698-701.
MEESTER,
J. A. J., I. L.
RAUTENBACH,
N. J.
DIPPENAAR,
AND
C. M.
BAKER.
1986. Classi ica ion o sou he n A ican mammals.
T ans aal Museum, Monog aph
No.5,
P e o ia,
359
pp.
MILLER,
G. S., JR. 1902. Two new opical Old Wo ld ba s. P o-
ceedings o he Biological Socie y o Washing on, 15:245-246.
MUTERE,
F. A. 1969. Fligh ac i i y o opical Mic ochi op e a,
Tada ida (Chae ephon)
pumila
C e zschma and Tada ida
(Mops) condylu a A. Smi h. Lynx,
10:53-59.
· 1973. Rep oduc ion in wo species o equa o ial ee-
ailed ba s (Molossidae). Eas A ican Wildli e Jou nal, 11:
271-280.
O'SHEA,
T. J.,
AND
T. A.
VAUGHAN.
1980. Ecological obse a ions
on an Eas A ican ba communi y. Mammalia, 44:485-496.
PETERS,
W. C. H. 1852. Na u wissenscha liche Reise nach Mos-
sambique, Zoologie: 1. Sauge hie e, Be lin, 202 pp.
PETERSON,
R. L., J. L.
EGER,
AND
L.
MITCHELL.
1995. Chi op-
e es. Faune de Madagasca , 84. Museum na ional d'His oi e
na u elle, Pa is, 204 pp.
PIENAAR,
U.
DE
V., S. C. J.
JOUBERT,
A.
HALL-MARTIN,
G. DE
GRAAFF,
AND
I. L.
RAUTENBACH.
1987. Field guide o he
mammals o he K uge Na ional Pa k. C. S uik Publishe s
and he Na ional Pa ks Boa d o T us ees, P e o ia, 176 pp.
POCHE,
R. M. 1975. The ba s o Na ional Pa k W, Nige , A ica.
Mammalia,
39:39-50.
RAUTENBACH,
I. L., M. B.
FENTON,
A. C.
KEMP,
AND
S. J.
VAN
JAARSVELD.
1990. Home ange and ac i i y o A ican gos-
hawks Accipi e achi o in ela ion o hei p eda ion on ba s.
Koedoe,
33:17-21.
ROBERTS,
A. 1926. Some new S. A ican mammals and some
changes in nomencla u e. Annals o he T ans aal Museum,
11:245.
· 1932. P elimina y desc ip ion o i y-se en new o ms
o Sou h A ican mammals. Annals o he T ans aal Museum,
15:1-19.
ROSEVEAR,
D. R. 1965. The ba s o Wes A ica. B i ish Museum
(Na u al His o y), London, 418 pp.
SIMPSON,
G. G. 1945. The p inciples o classi ica ion and a clas-
si ica ion o mammals. Bulle in o he Ame ican Museum o
Na u al His o y, 85:60.
SMITH,
S. A., J. W.
BICKHAM,
AND
D. A.
SCHLITTER.
1986. Ka -
yo ypes o ele en species o molossid ba s om A ica (Mam-
malia: Chi op e a). Annals o he Ca negie Museum,
55:125-
136.
SMITHERS,
R. H. N. 1971. The mammals o Bo swana. Memoi es
o he Na ional Museum o Rhodesia,
4:1-339.
· 1983. The mammals o he sou he n A ican sub egion.
Uni e si y o P e o ia P ess, P e o ia, 736 pp.
SMITHERS,
R. H. N.,
AND
V. J.
WILSON.
1979. Check lis and a las
o he mammals o Zimbabwe Rhodesia. Memoi es o he Na-
ional Museum o Zimbabwe-Rhodesia,
9:1-193.
TATE,
G. H. H. 1941. Molossid ba s o he A chbold collec ions.
Ame ican Museum No i a es, 1142:1.
THOMAS,
O. 1904. New ba s om B i ish Eas A ica A ica col-
lec ed by M s. Hinde and om he Came oons by M . G. L.
Ba es. Annals and Magazine o Na u al His o y, se e 7, 13:
206-210.
VAN
DER
MERWE,
M., S. R.
GIDDINGS,
AND
I. L.
RAUTENBACH.
Downloaded om h ps://academic.oup.com/mspecies/a icle/doi/10.2307/3504344/2600749 by gues on 30 Ap il 2024
6
1987.
Pos -pa um
oes us
in
he
li le
ee- ailed
ba ,
Ta-
da ida (Chae ephon)
pumila
(Mic ochi op e a: Molossidae) a
24°S.
Jou nal
o Zoology (London),
213:317-326.
VAN
DER
MERWE,
M., I. L.
RAUTENBACH,
AND
W. V
AN
DER COLF.
1986.
Rep oduc ion
in
emales
o
he
li le
ee- ailed
ba ,
Tada ida (Chae ephon) pumila, in
he
eas e n
T ans aal,
Sou h
A ica.
Jou nal
o
Rep oduc ion
and
Fe ili y,
77:355-
364.
VAUGHAN,
T. A.
1966.
Mo phology
and
ligh
cha ac e is ics
o
molossid
ba s.
Jou nal
o Mammalogy,
47:249-260.
VERSCHUREN,
J.
1957.
Ecologie, biologie e
sys ema ique
des
Chi op e es. Explo a ion du
Pa e
Na ional de
Ga amba.
Ins i-
u
des
Pa es
Na ionaux du Congo Belge,
7:1-473.
MAMMALIAN
SPECIES
574
.
1976.
Les chei op e es du Mon
Nimba
(Libe ia). Mam-
malia,
40:615-632.
WHITAKER,
J.
0.,
JR.,
AND
R. E.
MUMFORD.
1978.
Foods
and
ec-
opa asi es
o
ba s
om Kenya,
Eas
A ica.
Jou nal
o Mam-
malogy, 59:632--634.
Edi o s
o
his
accoun
we e
CYNTHIA
E.
REBAR,
KARLF.
KOOPMAN,
ELAINE
ANDERSON,
VIRGINIA
HAYSSEN,
and
ALICIA
V.
LINZEY.
Managing
edi o
was
BARBARA
H.
BLAKE.
S.
BOUCHARD,
DEPARTMENT
OF
BIOLOGY,
YORK
UNIVERSITY,
NORTH
YORK,
ONTARIO,
CANADA
M3J
IP3.
Downloaded om h ps://academic.oup.com/mspecies/a icle/doi/10.2307/3504344/2600749 by gues on 30 Ap il 2024
