Predation by giant centipedes, Scolopendra gigantea, on three species of bats in a Venezuelan cave

Abstract

We report the first known cases of predation by centipedes, Scolopendra gigantea (Chi-lopoda, Scolopendromorpha, Scolopendridae), on three species of bats (Mammalia, Chiroptera), Mor-moops megalophylla and Pteronotus davyi (Mor-moopidae), and Leptonycteris curasoae (Phyllo-stomidae). Our observations were made in Cueva del Guano, a limestone cave in Paraguaná Peninsula, Venezuela, that harbors important colonies of five bat species. These observations show that, noctur-nally and diurnally, centipedes can perform two ac-tions that most other bat predators cannot. First, they climb cave ceilings to catch and eat flying or perch-ing bats. Second, they subdue bats substantially heavier than themselves. Such capabilities may al-low large centipedes to prey on bats in what other-wise would be safe roosts.

Full text

Caribbean Journal of Science, Vol. 41, No. 2, 340-346
Copyright 2005 College of Arts and Sciences
University of Puerto Rico, Mayagu¨ez
Predation by Giant Centipedes,
Scolopendra gigantea, on Three
Species of Bats in a
Venezuelan Cave
JESÚSMOLINARI,
1
ELIÉCER E. GUTIÉRREZ,
1,2
ANTONIO A. DEASCENÇÃO,
1
JAFET M. NAS-
SAR,
2
ALEXIS ARENDS,
3
ROBERT J. MÁR-
QUEZ,
11
Departamento de Biología, Facultad de
Ciencias, Universidad de Los Andes, Mérida
5101, Venezuela,
2
Centro de Ecología, Instituto
Venezolano de Investigaciones Científicas,
Caracas 1020, Venezuela,
3
Centro de Investiga-
ciones en Ecología y Zonas Aridas, Universidad
Nacional Experimental Francisco de Miranda,
Coro 4101, Venezuela. Corresponding author:
molinari@ula.ve
ABSTRACT.—We report the first known cases of
predation by centipedes, Scolopendra gigantea (Chi-
lopoda, Scolopendromorpha, Scolopendridae), on
three species of bats (Mammalia, Chiroptera), Mor-
moops megalophylla and Pteronotus davyi (Mor-
moopidae), and Leptonycteris curasoae (Phyllo-
stomidae). Our observations were made in Cueva del
Guano, a limestone cave in ParaguanáPeninsula,
Venezuela, that harbors important colonies of five
bat species. These observations show that, noctur-
nally and diurnally, centipedes can perform two ac-
tions that most other bat predators cannot. First, they
climb cave ceilings to catch and eat flying or perch-
ing bats. Second, they subdue bats substantially
heavier than themselves. Such capabilities may al-
low large centipedes to prey on bats in what other-
wise would be safe roosts.
KEYWORDS.—Leptonycteris,Mormoops, Paraguaná
Peninsula, predation, Pteronotus,Scolopendra.
Compared to other mammals, bats suffer
low predation rates. However, because bats
are long-lived and reproduce slowly, the
impact of predation on their populations is
probably greater than assumed (Tuttle and
Stevenson 1982). A large variety of verte-
brates prey on bats (Hardy 1957; Barr and
Norton 1965; Gillette and Kimbrough 1970;
Hopkins and Hopkins 1982; Rodríguez and
Reagan 1984; Rodríguez-Durán and Lewis
1985; Rodríguez-Durán 1996; Hutterer and
Ray 1997; Souza et al. 1997; Sparks et al.
2000) while the few reports of invertebrate
predation involve cockroaches, ants, larval
beetles, and araneomorph spiders (Rice
1957; Gillette and Kimbrough 1970; Wilson
1971; McCormick and Polis 1982; Herman-
son and Wilkins 1986).
Because of their carnivorous tendency,
terrestrial habitat, and either gregarious
foraging or large body size, a diversity of
arthropods, including praying mantises,
wasps, mygalomorph spiders, scorpions,
solpugids, decapod crustaceans, and scolo-
pendrid centipedes, are potential bat
predators (Hutton 1843; Millot 1943; Grant
1959; Cloudsley-Thompson 1968; Banta
and Marer 1972; Gertsch 1979; McCormick
and Polis 1982, 1990). Scolopendrid centi-
pedes prey on frogs and toads up to 95 mm
long, small lizards, snakes up to 247 mm
long, birds up to the size of a sparrow, and
both field and house mice (Wells-Cole 1898;
Okeden 1903; Shugg 1961; Cloudsley-
Thompson 1968; Easterla 1975; Clark 1979;
Lewis 1981; McCormick and Polis 1982;
Carpenter and Gillingham 1984).
We report predation under natural con-
ditions by the large scolopendrid centi-
pede, Scolopendra gigantea (Chilopoda, Sco-
lopendromorpha, Scolopendridae), on
three bat species (Mammalia, Chiroptera).
Each of these predation cases resulted from
an independent fortuitous observation. The
study site, Cueva del Guano, is a limestone
cave located in a nearly flat thorn-scrub re-
gion, in ParaguanáPeninsula, Venezuela,
at 69°56’44’’W, 11°53’50’’N, and elevation
120 m. Before entering the cave, one must
descend 10 m through a 15 ×6 m sink that
leads to the southern edge of a 24 ×20 m
antechamber. The cave’s entrance is ap-
proximately 1 m wide and high, and opens
on the western edge of the antechamber.
The cave, which consists of a 30 m main
gallery followed by two 20 m secondary
galleries, harbors 45,000-50,000 bats (SVE
1972; Matson 1974). At daytime, five bat
species roost in this refuge: Mormoops mega-
ms. received August 11, 2004; accepted February 18,
2005
NOTES340

lophylla,Pteronotus parnellii and P. davyi
(Mormoopidae); Leptonycteris curasoae
(Phyllostomidae); and Natalus tumidirostris
(Natalidae) (Linares and Ojasti 1974; Mat-
son 1974; Genoud et al. 1990; Bonaccorso et
al. 1992; Arends et al. 1995). Scolopendra gi-
gantea, the world’s largest centipede (maxi-
mum length >300 mm; Shelley and Kiser
2000), is common in Paraguanáand easily
found in the antechamber.
Nocturnal predation on Mormoops megalo-
phylla.—On 21 December 2000, at 21:45 h,
we found a S. gigantea perching from the
ceiling of the northeastern antechamber’s
edge, 5 m north and 22 m east of the cave’s
entrance, while feeding on a dead M. mega-
lophylla (Fig. 1). The centipede was 0.3 m
from the nearest vertical wall and 2 m from
the ground. Since the centipede and bat
were easily visible, we are confident that
we did not overlook them during a careful
examination of this area concluded at 19:30
h. Therefore, the centipede had been ma-
nipulating the bat where we found it (no
guano, sand, or clay on the pelage) for not
longer than 2 h and 15 min. Before collect-
ing both specimens, we observed them us-
ing flashlights for 30 min from less than 1 m
away. During this period, the centipede re-
mained attached to the ceiling with only
the last five pairs of legs, held the bat using
its first eight pairs of legs (excluding the
forcipules, or poison claws), and fed while
it moved its head from side to side. The bat,
a non-reproducing adult female, had blood
in the wounds, lax wings and legs, patagia
fully retained their natural elasticity, and
fecal pellets protruded from the anus and
adhered to the uropatagium. Its forearm
was 55.9 mm long, which equals the mean
for adult females of M. megalophylla from
Cueva del Guano. Mean body mass for
these females is 16.5 g. An ectoparasitic bat
fly, Nycterophilia mormoopsis (Diptera, Stre-
blidae) was attached to the bat’s abdominal
hair. The heel, toes, and claws of the feet
had tangled agglomerations of hair from
the upper abdomen and lower chest of the
bat (Fig. 1). Embedded in these agglomera-
tions, was another N. mormoopsis (the ali-
mentary tract of the centipede also con-
tained flies of this species; Table 1). Fecal
pellets found inside the bat’s intestine con-
tained the remains of a noctuid moth (Lepi-
doptera).
The centipede was a 15.2 g (probably 9.0-
9.5 g before eating) and 145 mm female that
had mature reproductive organs. We di-
vided its 6.3 g (including bat remains) and
124 mm alimentary tract into nine sections,
which we dissected separately (Table 1). To
determine the bat’s body region from
which hair in the alimentary tract came
from, we compared it with hair still on the
bat and with that of complete specimens of
M. megalophylla from the same locality and
date.
The centipede started eating around the
bat’s neck, continued into the chest, and
then into the abdominal region (Table 1).
When we collected the two specimens, the
centipede had devoured about 35% of the
bat’s body mass. Necropsy of the bat
showed that it was in the following condi-
tions: left temporal region of cranium with-
FIG. 1. A centipede, Scolopendra gigantea, holding
and eating a freshly-killed bat, Mormoops megalophylla,
while hanging from the ceiling in the antechamber of
Cueva del Guano, ParaguanáPeninsula, Venezuela.
NOTES 341

out muscle; left mandible disarticulated,
without muscle covering the masseteric
fossa and the angular and coronoid pro-
cesses; left ectotympanic bone displaced
from its natural position; cervical vertebrae
ventrally and sinistrally without muscle
and vascular tissues, no vestiges of trachea;
left half of neck devoid of a circular patch (6
mm diameter), carved from inside, of dor-
sal skin; left propatagium partially missing
along its external edge; left half of thorax
almost completely devoid of skin; right half
of thorax devoid of skin near sternum; left
upper abdomen without skin; left pectoral
muscles almost completely missing; right
pectoral muscles retaining about one-
fourth of their mass; left ribs exposed, de-
tached from sternum, and displaced to
form a large opening to the middle of the
pectoral cavity; right ribs exposed, in their
natural position; pectoral cavity empty; left
scapula ventrally without muscle; hypo-
dermis of skin between scapulae exposed,
skin intact; most of digestive tube missing;
liver and left kidney in their natural posi-
tion, retaining most of their mass, right kid-
ney intact; no bones missing.
Diurnal predation on Leptonycteris
curasoae.—On 7 December 2001, at 12:00 h,
we found a ∼210 mm S. gigantea feeding on
a dead L. curasoae on the floor of the ante-
chamber’s tunnel connecting to the cave,
nearly directly below the cave’s entrance.
Mean forearm length for adult L. curasoae
from Cueva del Guano is 54.3 mm, and
mean body mass is 26.5 g. The centipede
used its first eight pairs of legs (excluding
forcipules) to hold the bat. We touched the
centipede several times with a stick, at-
tempting to scare it away and release the
bat, but contrary to what we expected, the
centipede responded by partly coiling
around the bat and holding it more tena-
ciously. During 30 min of observations,
the centipede fed without interruption,
carving a wound in the bat’s lower abdo-
men. The bat also had a second and smaller
wound on the upper back, which prob-
ably represented the initial bite of the cen-
tipede. The wings were lax, the exposed
TABLE 1. Sequence of consumption of body parts of a preyed bat individual, Mormoops megalophylla,as
reflected by the contents of nine dissected sections, ordered from anus to mouth, of the alimentary tract of a
centipede predator, Scolopendra gigantea.
Part of alimentary tract
Anatomic region Hindgut Mid-gut Esophagus
Dissected section 9
th
8
th
7
th
6
th
5
th
4
th
3
rd
2
nd
1
st
Length of section (mm) 130 130 136 136 136 136 136 150 150
Contents of alimentary tract
Hair tufts without skin ●●
Hair tufts joined by skin ●●䡩 䡩䡩 䡩 䡩
Hair (neck) + +
Hair (chest) +++++
2
Hair (lower chest) + + + +
Hair (upper abdomen) + + + +
Hair (abdomen) ++ +
Ectoparasitic bat flies
1
䡩
Propatagium 䡩
Muscle ●●● ●● 䡩 䡩
Heart 䡩
Lung ●●
Gut ●● ●
Liver 䡩
Kidney 䡩
●= Predominant tissue fragments; 䡩= minor tissue fragments, or other lesser components; + = present.
1
Two flies (Nycterophilia mormoopsis) adhered to hair tufts;
2
chest hair prevails over hair from other body
regions of the bat.
NOTES342

flesh was bloodstained, and the pelage was
clean.
Diurnal predation on Pteronotus davyi.—
On 13 March 2003, at about 11:30 h, we
found a ∼160 mm S. gigantea feeding on a
dead P. davyi while perching from the ceil-
ing of the antechamber’s tunnel connecting
to the cave, 1 m away from the cave’s en-
trance, 0.2 m from the nearest vertical wall,
and 1.5 m above the cave’s floor. Mean
forearm length for adult P. davyi from
Cueva del Guano is 47.7 mm, and mean
body mass is 9.7 g. The centipede handled
the bat in a posture similar to that shown in
Fig. 1. However, the centipede remained at-
tached to the ceiling with the last eight
pairs of legs, and held the bat using its first
seven pairs of legs (excluding forcipules).
During 30 min of observations, the centi-
pede fed on the bat’s chest and abdomen.
The bat also had a large wound on the back
resulting from previous eating by the cen-
tipede. As in the other two cases, the dead
bat was lax, bleeding, and had no dirt on
pelage.
Discussion.—We are convinced that the
centipedes killed the three bats in situ
shortly before we found them. Absence of
rigor mortis, presence of fresh blood in
wounds, and cleanliness of pelage indicate
that the bats died shortly before and were
not transported from other places. Since ec-
toparasitic flies stay on a dead bat only
while it remains warm, the N. mormoopsis
on the M. megalophylla and in the alimen-
tary tract of the associated centipede show
that the bat died just before being rapidly
eaten. Fecal pellets found on this bat sug-
gest that it had foraged and fully digested
prey the same night. Further evidence of
predation on this bat is the tangled pectoral
hair found in its toes (Fig. 1), which indi-
cates an energetic attempt to dislodge the
centipede. Carrion-eating insects in Cueva
del Guano are superabundant and quickly
feed on fallen bats, thus their absence on
the L. curasoae indicates a recent death and
fall to the floor. Resistance of the centipede
to release this bat suggests that it confused
human interference with struggling by
prey. This behavior would not be expected
if the centipede had not killed the bat.
The centipedes most likely seized the
bats in the ceiling of the antechamber of
Cueva del Guano by one of three methods.
Firstly, the centipedes may have crawled
along the ceiling, with their venters facing
upwards, actively searching for perching
bats. However, scolopendrids do not usu-
ally pounce upon prey from a distance
(Manton 1965). Secondly, the centipedes
may have waited statically on the ceiling
for a bat to perch within striking distance
(we have seen centipedes “resting”for
hours on the walls and ceiling of the ante-
chamber). This could be the case because
scolopendrids learn to climb high sub-
strates where airborne insects alight, and
even hang from such substrates using their
rear legs (Remmington 1950). Thirdly, the
centipeds may have employed a strategy
used by snakes to seize bats in caves and
trees (e.g., Hardy 1957; Barr and Norton
1965; Hopkins and Hopkins 1982; Ro-
dríguez and Reagan, 1984), i.e., hang from
the ceiling and wait for a bat to fly close
enough to catch it in mid-air (as suggested
by the posture in which we observed the
centipede eating the M. megalophylla and
the P. davyi, Fig. 1). In spite of the vigor
expected from a flying bat, this could also
be the case because scolopendrids (1) can
use multiple pairs of legs to get firmly an-
chored to substrates, (2) can quickly immo-
bilize prey with their legs and with their
venom, (3) seize flying prey in mid-air by
raising the anterior part of the body, (4)
hold struggling prey tenaciously (Cloud-
sley-Thompson 1968; Lewis 1981), and (5)
the tunnel in which L. curasoae and the P.
davyi were being eaten is a high traffic fly-
way in which bats seldom perch. Which-
ever was the case, our observations on pre-
dation on M. megalophylla and P. davyi
show that, unlike most predators, large sco-
lopendrids can crawl, with their ventral re-
gion facing upwards, along cave ceilings in
search of flying or perching bats in what
otherwise would be safe roosts. In this con-
text, the L. curasoe eaten on the ground was
likely a result of the centipede’s fall while
struggling with the bat.
Venomous arthropods capture larger
prey than non-venomous ones (McCormick
and Polis 1982). Therefore, the more ven-
omous the arthropod the larger the prey it
NOTES 343

should be able to catch. Two of our cases
(M. megalophylla,L. curasoe), in which prey
may have nearly doubled the body masses
of their victimizers, show the extraordinary
predatory capabilities of an arthropod with
a potent venom. These cases are more re-
markable because of the difficulties in-
volved in capturing energetic and volant
prey while hanging from cave ceilings.
Centipedes are known to use up to eight
pairs of their anterior legs to manipulate
prey, with the number of pairs involved
varying proportionally with prey size
(Lewis 1961, 1981; Elzinga 1994). Therefore,
in both cases, use of the first eight pairs of
legs to hold the bats suggests that such
prey may have been around the maximum
permissible size for the centipedes.
Scolopendrids often inflict an initial
puncture with their forcipules on the neck
of vertebrate prey (Shugg 1961; Cloudsley-
Thompson 1968). This should quickly im-
mobilize the prey because the neurotoxic
venom (Bücherl 1946; Stankiewicz et al.
1999) is injected near the brain. The con-
sumption of the M. megalophylla began be-
hind the bat’s head. This may also have
happened to the L. curasoae and the P. davyi,
as suggested by the wounds on their backs.
Therefore, injection of venom near the
brain may have occurred and facilitated
subduing the three bats. Although the tox-
icity of centipede venom to bats is un-
known, intravenous injection of the venom
gland extract of Scolopendra viridicornis kills
mice in 20-32 sec, and direct puncture of the
forcipules of this centipede in the tail of
mice is always lethal, causing death in a
mean time of 3 min (Bücherl 1946).
Even though arthropods comprise the
bulk of the diet of all centipedes (Lewis
1981; Cloudsley-Thompson 1968), large
scolopendrids do not miss the opportunity
to feed on vertebrates (Wells-Cole 1898;
Okeden 1903; Shugg 1961; Easterla 1975;
Clark 1979; Lewis 1981; McCormick and
Polis 1982; Carpenter and Gillingham
1984). Mice were used as the main food for
captive S. gigantaea, which “devoured them
with alacrity”(Cloudsley-Thompson 1968).
A 120 mm centipede consumed a large
chunk of flesh from a human corpse (Ha-
rada et al. 1999). Our observations took
place in an environment providing extraor-
dinary abundance of arthropod prey, espe-
cially large (45 mm) cockroaches, Blaberus
discoidalis (Blattaria, Blaberidae) which we
have seen caught and eaten by S. gigantea.
This suggests that vertebrates may be par-
ticularly rewarding prey for centipedes be-
cause of their nutritional composition, or
because the quantity of nutrients per indi-
vidual prey exceeds that provided by in-
vertebrates.
Scolopendrids belong to a major and an-
cient arthropod lineage, are morphologi-
cally disctinctive, some are among the larg-
est terrestrial arthropods, are common in
the tropics, and are of medical and phar-
macological interest (Manton 1965; Cloud-
sley-Thompson 1968; Lewis 1981; McCor-
mick and Polis 1982; Knysak et al. 1998;
Stankiewicz et al. 1999). Despite their im-
portance, little information exists on their
behavior under natural conditions. On the
other hand, most observations of predation
on bats involve easily observed raptorial
birds around North American caves (Gil-
lette and Kimbrough 1970; Sparks et al.
2000). There is little information on other
predators, particularly in the tropics where
bats and their enemies are most diverse.
Research is needed both to learn more
about the foraging ecology of scolopen-
drids, especially of large ones, and to evalu-
ate the impact of diverse predators on
tropical bat populations.
Acknowledgments.—The International
Foundation for Science (Stockholm), and
the Consejo de Desarrollo Científico Hu-
manístico y Tecnológico (Universidad de
Los Andes) provided financial support to J.
Molinari. G. de Couto, M. Figuera, M.
Ghodke, A. Moreno, J. Lupi, and E. Santana
sent us literature. J. Aranguren assisted A.
Arends during fieldwork. The reviewers,
especially R. Shelley, provided valuable
comments.
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