Molinari & Gutiérrez
28
ANARTIA
Publicación del Museo de Biología de la Universidad del Zulia
ISSN 1315-642X (impresa) / ISSN 2665-0347 (digital)
https://doi.org/10.5281/zenodo.16501801 / Anartia, 40 (junio 2025): 28-38
Mortality in three species of bats of the genus Pteronotus
Gray, 1838 (Mammalia, Chiroptera, Mormoopidae) due to
overpopulation, and harassment by blaberid cockroaches,
in a Venezuelan cave
Mortalidad en tres especies de murciélagos del género Pteronotus Gray, 1838
(Mammalia, Chiroptera, Mormoopidae) debida a sobrepoblación, y hostigamiento
por cucarachas blabéridas, en una cueva venezolana
Jesús Molinari1 & Eliécer E. Gutiérrez2
1Departamento de Biología, Facultad de Ciencias, Universidad de Los Andes, Mérida 5101, Venezuela. orcid.org/0000-0002-9393-5483
2Programa de Pos-Graduação em Biodiversidade Animal, Centro de Ciências Naturais e Exatas, Universidade Federal de Santa Maria,
Rio Grande do Sul, Brazil. orcid.org/0000-0001-6790-8185
Correspondence: jmvault@gmail.com
(Received: 06-01-2025 / Accepted: 17-03-2025 / On line: 31-07-2025)
ABSTRACT
Nearly half of the world’s bat species roosts in caves. Despite the availability of caves being thought to limit their abun-
dance, roost overpopulation has not been listed as a cause of mass mortality in cavernicolous bats, suggesting that predation
and other factors will normally prevent them from surpassing the carrying capacity of caves. We report mortality likely due
to roost overpopulation in three species of insectivorous bats inhabiting a small cave located in a non-karstic outcrop in
the Venezuelan Llanos. Outside the cave, abundant bat remains were found washed away by rainwater. At the entrance of
the cave, bats were roosting exposed to sunlight (a highly unusual behavior for the species involved), which thus risked de-
hydration, and detection by diurnal predators. Inside the cave, the ceiling, walls, and part of the oor were densely packed
with bats. On the oor, bats were resting in quadrupedal position and interacted agonistically with large guanophilic
cockroaches. Most of the bats had skin lesions apparently caused by cockroach bites. We attribute these ndings to roost
overpopulation, and suggest that, owing to the orography, lithology and high primary production in the region, caves used
as roosts by insectivorous bats are in much shorter supply than insect prey.
Keywords: Blaberidae, Blattodea, Dictyoptera, guano invertebrates, interspecic aggression, predation, Pteronotus davyi,
Pteronotus gymnonotus, Pteronotus personatus, Venezuelan Llanos.
RESUMEN
Casi la mitad de las especies de murciélagos del mundo se refugia en cuevas. Pese a que se cree que la disponibilidad de
cuevas limita su abundancia, la sobrepoblación de refugios no ha sido listada como una causa de mortalidad en masa de
murciélagos cavernícolas, lo cual sugiere que la depredación y otros factores normalmente les impiden superar la capacidad
de carga de las cuevas. Reportamos mortalidad probablemente debida a sobrepoblación del refugio en tres especies de mur-
ciélagos insectívoros que habitan una pequeña cueva ubicada en un aoramiento no kárstico en los Llanos venezolanos.
Fuera de la cueva se encontraron abundantes restos de murciélagos arrastrados por el agua de lluvia. En la entrada de la
cueva, los murciélagos dormían expuestos a la luz solar (un comportamiento altamente inusual para la especie involucrada),
corriendo el riesgo de deshidratarse y ser detectados por depredadores diurnos. Dentro de la cueva, el techo, las paredes y
Bat mortality in a Venezuelan cave
29
ANARTIA
Publicación del Museo de Biología de la Universidad del Zulia
ISSN 1315-642X (impresa) / ISSN 2665-0347 (digital)
https://doi.org/10.5281/zenodo.16501801 / Anartia, 40 (junio 2025): 28-38
parte del suelo estaban densamente cubiertos por murciélagos. En el suelo, los murciélagos estaban descansando en posición
cuadrúpeda e interactuaban de forma agonística con grandes cucarachas guanólas. La mayoría de los murciélagos tenían
lesiones en la piel aparentemente causadas por mordeduras de cucarachas. Atribuimos estos hallazgos a la sobrepoblación
de refugios y sugerimos que, debido a la orografía, la litología y la elevada producción primaria en la región, las cuevas utili-
zadas como refugio por murciélagos insectívoros son escasas relativamente a la abundancia de insectos presa.
Palabras clave: agresión interespecíca, Blaberidae, Blattodea, depredación, Dictyoptera, invertebrados del guano, Llanos
venezolanos, Pteronotus davyi, Pteronotus gymnonotus, Pteronotus personatus.
INTRODUCTION
With 1,475 currently recognized species, bats (Chi-
roptera) are the second most diverse order of living mam-
mals (ASM 2024). Almost all bat species are exclusively
nocturnal (Speakman 1995), and about 48.5% of them
are dependent on caves as diurnal retreats (Tanalgo et al.
2022). Cavernicolous bats form the largest aggregations
of vertebrates, ranging from tens of thousands to millions
of individuals (Kunz 1982, Russell & McCracken 2006,
Furey & Racey 2016). Some bat species are found only in
few caves (e.g., the Paraguana Common Mustached Bat,
Pteronotus paraguanensis Linares & Ojasti, 1974 in Para-
guana Peninsula, Venezuela; Gutiérrez & Molinari 2008),
or even in just one (e.g., the Cuban Greater Funnel-eared
Bat, Natalus primus Anthony, 1919, in Guanahacabibes
Peninsula, Cuba; Tejedor 2011).
While noting that bats are the only major vertebrate
group that continuously and extensively uses caves as
shelter, Jepsen (1970) suggested that ancestral bats took
advantage of their capacity for echolocation to become
cavernicolous, which in turn allowed them to evade avian
predators, and to conserve energy and body water when
not foraging. Humphrey (1975) proposed roosts, espe-
cially caves, to be more important than the growing sea-
son and insect diversity as determinants of the distribu-
tion and abundance of temperate bats. Caves and articial
cave-like structures such as tunnels and abandoned build-
ings are deemed fundamental for the conservation of bats
worldwide (Arita 1993, Furey & Racey 2016, Medellín
et al. 2017, Tanalgo et al. 2022, Guixé et al. 2024, Meier-
hofer et al. 2024), and the construction of articial caves
has been proposed to protect some bat species (Lavoie &
Northup 2009, Gulickx et al. 2011, Slider & Kurta 2011,
Molinari et al. 2012, Albayrak 2013, Gibbons 2013, Mer-
ing & Chambers 2014).
By bringing huge amounts of nutrients into caves in the
form of guano and their own carcasses, bats have created
the conditions for the evolution and survival of unique
and highly diverse communities of cavernicolous organ-
isms, ranging from bacteria and yeasts to arthropods and
vertebrates (Ferreira & Martins 1999, Gnaspini & Trajano
2000, Fenolio et al. 2006, Biswas & Shrotriya 2010, Cunha
et al. 2020, Pimentel et al. 2022, Lundberg & McFarlane
2024). Cavernicolous bats provide major ecosystem ser-
vices outside caves, including pollination, seed dispersal,
and control of insect populations (Jones et al. 2009, Kunz
et al. 2011, Ghanem & Voigt 2012, Medellín et al. 2017,
Ramírez-Fráncel et al. 2021).
Bats congregating in caves are aected by higher levels
of parasitism and are exposed to a greater risk of epidem-
ics than solitary and foliage-roosting bats (Patterson et al.
2007, O’Shea et al. 2016, Hoyt et al. 2021). Within and
around caves, bats serve as prey for cockroaches (Rice
1957, Wilson 1971, Braack 1989, Bell et al. 2007, López-
Wilchis et al. 2023), mealworms (Hermanson & Wilkins
1986), spiders (Nyeler & Knörnschild 2013, Dias et al.
2015, Leivers et al. 2021), and centipedes (Molinari et al.
2005); and for predatory vertebrates, including sh and
amphibians (Yager & Williams 1988, Mikula 2015), liz-
ards and snakes (Herreid 1962, Hammer & Arlettaz 1998,
Esbérard & Vrcibradic 2007, Barti et. al. 2019, Clarkson
& Massyn 2020, Tanalgo et al. 2020), raptors and other
birds (Twente 1954, Harden 1972, Looney 1972, Bar-
clay et al. 1982, Fenton et al. 1994, Lenoble et al. 2014,
Spitzenberger et al. 2014, Mikula et al. 2016, Sieradzki &
Mikkola 2020), and mammals (Urbanczyk 1981, Kokure-
wicz 2004, Rodríguez-Durán et al. 2010, McAlpine et al.
2011, Mas et al. 2015, Haarsma & Kaal 2016, Cichocki
et al. 2021). e latter include both specialized and op-
portunistically carnivorous bat species (Fischer et al. 1997,
Bordignon 2005, Oprea et al. 2006, Rodríguez-Durán &
Rosa 2020). Remarkably, there is a record of diurnal birds
entering caves to catch and eat hibernating bats (Estók et
al. 2010).
In caves, cockroaches are known to eat bat pups that
have fallen on guano before the latter can climb to a safe
place (Rice 1957, Wilson 1971, Bordon 1995, Bell et al.
2007). e bat genus Pteronotus Gray, 1838 is divided into
three subgenera: Pteronotus; Chilonycteris Gray, 1839;
and Phyllodia Gray, 1843 (Smith 1972). e subgenus
Pteronotus includes three species of naked-backed bats,
Molinari & Gutiérrez
30
whose wing membranes are attached to the midline of the
dorsum and fully cover their backs, which thus appear to
be hairless though they are not. ere is a single report of
possibly predatory behavior of cockroaches on adult bats
(López-Wilchis et al. 2023). It involved a single individu-
al of each species, namely the cockroach, Nyctantonina az-
teca (Saussure & Zehnter, 1893) (Dictyoptera, Blattodea,
Nyctiboridae), and the omas’s Naked-backed Bat, Pter-
onotus fulvus (omas, 1892) (Mammalia, Chiroptera,
Mormoopidae) in a Mexican cave. A large N. azteca was
found lodged in the interstice between the lower surface
of the le-wing membrane and the back of the P. fulvus.
Because both organisms had mutually caused injuries that
were deemed not to be recent, the authors interpreted this
interaction as a case of accidental ectoparasitism.
Venezuela is the country in which Pteronotus is most di-
verse. Seven species of this bat genus were included in the
latest national list (Boher-Bentti et al. 2023): 1) the Davy’s
Naked-backed Bat, Pteronotus (Pteronotus) davyi Gray,
1838; 2) the Big Naked-backed Bat, P. (Pt.) gymnonotus
(Wagner, 1843); 3) the Wagner’s Lesser Mustached Bat,
P. (Chilonycteris) personatus (Wagner, 1843); 4) the Al-
len’s Common Mustached Bat, P. (Phyllodia) fuscus (J. A.
Allen, 1911); 5) the Wagner’s Common Mustached Bat,
P. (Ph.) rubiginosus (Wagner, 1843); 6) P. (Ph.) paragua-
nensis; and 7) the Amazonian Common Mustached Bat, P.
(Ph.) alitonus Pavan, Bobrowiec & Percequillo, 2018.
Species of the genus Pteronotus are dependent on caves
and articial cave-like structures as diurnal retreats (e.g.,
Silva-Taboada 1979, Molinari et al. 2012, Soto-Centeno
et al. 2015). Members of the subgenus Phyllodia show a
tight dependence on forest (Gutiérrez & Molinari 2008,
Oliveira et al. 2015, Martino et al. 2019), thus are largely
absent from the savanna corridor (where caves are also
scarce) of the Venezuelan Llanos region (Gutiérrez & Mo-
linari 2008), which cover approximately 240,000 km2 and
are bounded to the west by the Andes, to the north by the
Venezuelan Coast Range, and to the south by the Orinoco
River (Huber et al. 2006).
Despite the availability of suitable caves being thought
to limit their abundance (Humphrey 1975, Rodríguez-
Durán 2009, Furey & Racey 2016, Vargas-Mena et al.
2020), roost overpopulation has not being listed as a
cause of mass mortality in cavernicolous bats (O’Shea et
al. 2016). is suggests other causes of mortality, the most
important of which likely is predation by natural enemies
(Mikula et al. 2024), to normally prevent the size of the
colonies of cavernicolous bats from surpassing the carrying
capacity of caves.
We report the rst known case of multiple mortality
in adult bats due to the carrying capacity of a cave being
surpassed, and to harassment by a dense population of
cave-dwelling cockroaches. e observations involve three
species of Pteronotus and were recorded in the Venezuelan
Llanos.
STUDY AREA AND METHODS
e observations reported here were carried out in a
small cave (Fig. 1) locally known as Cueva del Viejo [=
Old Man’s Cave], or Cueva del Ermitaño [= Hermit’s
Cave], which despite these names is not suitable for hu-
man habitation owing to its harsh interior environment.
is cave is located at 8.98° N and 68.27° W, 2.7 km ENE
El Baúl, Estado Cojedes, Venezuela, in El Baúl Massif, a
complex of scattered hills composed of igneous, metamor-
phic and metasedimentary rocks of late Cambrian to early
Permian age that emerges from beneath the mainly ua-
ternary alluvial deposits of the Venezuelan Llanos (Bucher
1952, Huber et al. 2006, Viscarret et al. 2012). It is at an
elevation of 235 m, 500 m to the east of the road (Carret-
era Regional El Baúl-Las Galeras) and at lowlands with
elevations of 60–70 m. e climate of the region is char-
acterized by a single rainy season (April-May to October)
alternating with a dry season (November to April-May)
(Huber et al. 2006).
At dawn on 19 January 2001, we ascended from the
road to the cave following a rocky path. We stayed at the
cave for 6 hours, capturing bats with a hand net and per-
forming observations. e cave consists of a single cham-
ber approximately 12 m long, 5 m wide, and 3 m high. e
entrance (Fig. 1, top le) is its highest point, and the oppo-
site end is its lowest point. us, the posterior of the cave
functions as a cul-de-sac in which the layer of bat guano
becomes progressively thicker, wetter, and more decom-
posed while going deeper. At the bottom of the cave, the
concentration of ammonia made the air unbreathable for
humans; thus, to observe the interaction between bats and
cockroaches, we visited this portion of the cave multiple
times while holding our breath. We had planned a subse-
quent visit with better equipment, but this was not pos-
sible, thus we decided to prepare this report.
RESULTS
As we ascended to the cave, at about 50 m from its
entrance, we began to nd on the ground disarticulated
bones and whole skeletons belonging to the same bat spe-
cies found in the cave, which were Pteronotus davyi, P.
gymnonotus, and P. personatus. Of the three, the last species
was the least abundant. No other bat species seemed to be
present in the cave. e number of skeletons increased as
Bat mortality in a Venezuelan cave
31
we were getting closer to the cave. We assumed that these
remains were washed away by rainwater owing along and
around the path to the cave.
Upon reaching the cave, we discovered the likely source
of the skeletons: numerous bats roosting at the entrance
of the cave despite being exposed to sunlight (Fig. 1, top
right). is was entirely unexpected, because species of
Pteronotus, as other bats with small eyes, are normally
found only in the dark part of caves. Bats roosting at the
entrance of caves can be exposed to diurnal predators and,
if not adapted to roost outside caves, may lose body water.
Bats dying inside the cave were unlikely to be the source of
the skeletons because any water entering the cave would
drain inwards together with the guano, since as mentioned
above the cave decreases in height from entrance to bot-
tom.
Figure 1. Top le) Entrance of the cave photographed from inside the cave. Note that the interior of the cave is lower than its entrance.
e human silhouette is that of EEG. Top right) Bats roosting exposed to sunlight at the entrance of the cave, photographed from
outside the cave. Bottom) Bats on the oor of the cave, which had started to escape when approached to obtain the photograph.
Molinari & Gutiérrez
32
e ceiling and walls of the cave were densely packed
with roosting bats. e guano on the oor of the cave
and the low areas of the cave walls were covered by dense
groups of cockroaches of an unidentied species (Dicty-
optera, Blattodea, Blaberidae; Fig. 1, bottom), most of
which were too large to be potential prey for these bats.
Another unexpected nding was an aggregation of bats
resting in a quadrupedal position on the oor, deep in the
cave (Fig. 1, bottom). ey were tightly packed, and all had
their heads pointing in the direction of the entrance of the
cave, which is not evident in Fig. 1 because they started to
y whenever we approached them at a short distance (~1
m) to obtain photographs. We were able to observe the
behavior of these bats by staying 2–3 m away. e aggrega-
tion covered a semicircular area of 5–6 m2. Its curved edge
was composed of bats with their hindquarters in contact
with the posterior walls of the cave. Its straight edge was
composed of bats arranged to form a sharp line limiting
with cockroach-covered guano. e bats, especially those
in the front edge, were alert (heads continually moving,
mouths open) and agonistic (pouncing and retreating) to-
wards the cockroaches. Aer a short while, the bats in the
front edge would take ight and land on the backs of bats
in the rear of the aggregation, thus were replaced by the
bats that previously were just behind them, which in turn
repeated this behavior. us, the aggregation functioned
as a continuous track in which the treads were the bats.
As the bats took ight, it was evident that they had cock-
roaches below them.
We hand-netted ~120 bats within the cave. To comple-
ment material from this location already in museums, we
selected 33 specimens as vouchers and released the rest.
Most of the bats (>70%) that we handled had erosive
wounds (1–5 mm wide) on their wing and tail membranes
that may have been caused by cockroach gnawing.
DISCUSSION
Numerous instances of three species of Pteronotus
inhabiting the same diurnal roost have been reported
throughout the Neotropics: 1) e Macleay’s Mustached
Bat, P. (Ch.) macleayi (Gray, 1839), the Sooty Mustached
Bat, P. (Ch.) quadridens (Gundlach, 1840), and the Par-
nell’s Common Mustached Bat, P. (Ph.) parnellii (Gray,
1843) in 15 Cuban and two Jamaican caves (Silva-Taboa-
da 1979, Genoways et al. 2005, Tejedor et al. 2005); 2) P.
(Pt.) fulvus, the Dobson’s Lesser Mustached Bat, P. (Ch.)
psilotis, and the Mexican Common Mustached Bat, P.
(Ph.) mexicanus in ve caves and one abandoned mine in
Mexico (Bateman & Vaughan 1974, Torres-Flores et al.
2012, Ayala-Téllez et al. 2018); 3) P. gymnonotus, P. per-
sonatus, and the Mesoamerican Common Mustached Bat,
P. (Ph.) mesoamericanus in a Costa Rican cave (Deleva &
Chaverri 2018); and 4) P. gymnonotus, P. personatus, and
P. rubiginosus in two Brazilian caves (Zortéa et al. 2015,
Barros & Bernard 2023). In Venezuela, the only reports of
roost co-occurrence between species of Pteronotus involve
P. davyi and P. paraguanensis (the latter misidentied as ‘P.
personatus’: Bonaccorso et al. 1992, de La Torre & Medel-
lín 2010) in the three main caves of Paraguaná Peninsula
(Molinari et al. 2012). e Cueva del Viejo (Fig. 1, top
le) is smaller than all those caves and mines, and diers
from many of them in not being a hot cave (Silva-Taboada
1979, Rodríguez-Durán 2009). e non-karstic lithol-
ogy of El Baúl Massif (Viscarret et al. 2012) may limit the
number, size and suitability of caves available to bats of the
genus Pteronotus.
Savannas represent ~17% of the area and contribute
~30% of the primary production of the Earth’s terrestrial
vegetation (Grace et al. 2006). As in other biomes (Borer
et al. 2012, Cusens et al. 2012), in grasslands primary pro-
duction determines the abundance of phytophagous in-
sects (Prather & Kaspari 2019, Welti et al. 2020, Delabye
et al. 2022). ese arthropods are the dominant herbivores
in terms of biomass in savannas worldwide (Andersen &
Lonsdale 1990, Lewinsohn & Price 1996). Consequently,
in savanna regions like those of the Venezuelan Llanos, ow-
ing to the combined eect of orography (in our case also
the non-karstic lithology) and primary production, caves
used as roosts by insectivorous bats can be in much shorter
supply (relatively to the needs of the bats) than insect prey.
We believe that this is the situation that applies to the spe-
cies of Pteronotus at our study site, where three clear signs
of overpopulation were evident: rst, abundant remains
of these species were found outside the cave; second, bats
had to roost at the entrance of the cave, thus were exposed
to death by dehydration and diurnal predators; and third,
bats also had to roost on the oor of the cave, thus were
suering harassment by guanophilic cockroaches result-
ing in metabolic energy loss, and skin lesions apparently
caused by the bites of these insects.
Bats of the genus Pteronotus have a highly manoeuvrable
ight and forage in background-cluttered space: those of
the subgenera Pteronotus and Chilonycteris hunt arthropod
prey in proximity to but not within vegetation, whereas
those of the subgenus Phyllodia do so within dense vegeta-
tion (Jennings et al. 2004, Mancina et al. 2012, Oliveira et
al. 2015, Martino et al. 2019, personal observation). For
this reason, species of the rst two subgenera, such as P.
davyi, P. gymnonotus, and P. personatus, have the potential
to act as insect pest controllers at a low height above agri-
cultural land, even gleaning ightless pests directly from
Bat mortality in a Venezuelan cave
33
the vegetative organs of crop plants. Because in the Ven-
ezuelan Llanos the scarcity of caves likely limits their pop-
ulations, it should be feasible to construct articial caves
to increase the local densities of these bats, thus boosting
their eect as natural pest controllers. Such caves should
be positioned near farms and be designed to contain hot
chambers, thus making them ideal for mormoopids and
unsuitable for vampire bats, and to facilitate the harvest of
guano to be used as a fertilizer (Molinari et al. 2012).
Cockroaches are related to praying mantises, with
which they form the superorder Dictyoptera (Evangelista
et al. 2019, Ma et al. 2023). In the Mesozoic, the superor-
der included several lineages of cockroach-like predatory
species, possessing mantis-like forelegs (Dittmann 2015,
Vrsanský & Bechly 2015, Liang et al. 2018). e mandibu-
lar apparatus of cockroaches allows them to chew all sorts
of materials (Weihmann et al. 2015). Accordingly, extant
cockroach taxa are scavengers or detritivores, but there are
reports of them predating on other insects, and they can be
cannibalistic (Roth & Willis 1960, Bordon 1995, Persad
& Hoy 2004, Bell et al. 2007, Pfannenstiel et al. 2008).
Cockroaches are known to feed on corpses, and to bite
sleeping persons, carving signicant skin lesions (Roth &
Willis 1957, Denic et al. 1997, Uieda & Haddad 2014,
Viero et al. 2019). A dead mouse placed into a dense cock-
roach culture was skeletonized overnight, suggesting that
cockroaches might be used to clean osteological material
(Bell et al. 2007).
Cockroaches can be superabundant in caves, where
they feed on guano, but also on dead and live invertebrates
and vertebrates, including bats (Rice 1957, Roth & Wil-
lis 1960, Wilson 1971, Schal et al. 1984, Braack 1989,
Bordon 1995, Bell et al. 2007, López-Wilchis et al. 2023).
Cockroaches cause mortality in the cave swilets of tropi-
cal Asia by eating their nests, which are largely composed
of the birds’ salivary proteins (Cruz et al. 2008, Manchi
& Sankaran 2009). In the caves of Paraguaná Peninsula,
we have witnessed the American cockroach, Periplaneta
americana (Linnaeus, 1758), quickly taking away (or
tearing them apart when two or more cockroaches were
competing for the victim) wounded conspecics. Bordon
(1995) provided an impressive account of the behavior of
cockroaches feeding on dead bats in a Venezuelan cave. An
abridged translation follows: ‘Near the entrance some bats
were collected (Artibeus and Phyllostomus), which were le
on the ground in individual plastic bags to later obtain ec-
toparasites. en we entered the area of complete darkness.
Aer returning to the entrance, we noticed that the bags
with the bats had disappeared. A little further away we saw
something like large self-propelled balls that moved irregu-
larly om one place to another. ey were hundreds of giant
wingless cockroaches (probably Megaloblatta) which, aer
having broken the plastic bags, ercely and antically fought
over what was le of the bats; there were manifestations of
cannibalism’. erefore, under propitious circumstances,
such as those that we witnessed (Fig. 1), cave-dwelling
cockroaches can be expected to bite adult bats.
Similarly to López-Wilchis et al. (2023), we have dif-
culties to characterize the interaction between bats and
cockroaches at our study site. Because according to their
observations the cockroach had been living in symbiosis
with the bat for some time, they concluded that it was
an unusual case of accidental parasitism rather than pre-
dation. Based on our observations, we conclude that the
cave was overcrowded with bats, which thus were forced
to come into close contact with the cockroaches on the
guano. e agonistic behavior of the bats towards the
cockroaches suggests that their skin lesions were caused
by these insects. Given that there was no symbiosis be-
tween both kinds of organisms, if the assumption is made
that the cockroaches were biting the bats to opportunisti-
cally feed on them, their behavior could be classied as
accidentally predatory. Although cockroaches cannot kill
active adult bats, if present in huge numbers as observed
in this cave, they could harass them, thus contributing to
bat mortality.
ACKNOWLEDGMENTS
anks are due Dominic A. Evangelista, of the Depart-
ment of Entomology of the University of Illinois-Urbana,
for conrming (based on Fig. 1) the familial identity of
the cockroaches; to Antonio J. González-Fernández, of
the Universidad Nacional Experimental de los Llanos Oc-
cidentales “Ezequiel Zamora”, for his hospitality at the
Hato Mataclara, Carretera Regional El Baúl-Las Galeras,
Cojedes, Venezuela; and to Ángel L. Viloria, of the Insti-
tuto Venezolano de Investigaciones Cientícas (IVIC), for
bringing to our attention a rare but important piece of in-
formation (Bordon 1995).
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