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2021. Journal of Arachnology 49:407–409
Arachnids that feed on vertebrate carrion: necrophagy by the whip spider Paraphrynus raptator
(Amblypygi: Phrynidae) and its relation to the feeding behavior of the woolly false vampire bat
Chrotopterus auritus (Chiroptera: Phyllostomidae).
Luis A. Trujillo
,Rony E. Trujillo
and Rodrigo A. Medell´
Laboratorio de Ecolog´
on de Vertebrados Terrestres, Instituto de Ecolog´
ıa, Universidad Nacional Aut ´
onoma de M´
Exterior s/n, Ciudad Universitaria, C.P. 04510, Coyoaca
´n, Ciudad de M´
exico; E-mail: trujillososaluis@gmail.
Posgrado en Ciencias Biol´
ogicas, Universidad Nacional Aut ´
onoma de M´
exico, Av. Ciudad Universitaria 3000,
Ciudad Universitaria, C.P. 04510, Coyoaca
´n, Ciudad de M´
Sistema de Colecciones Biol ´
ıa, Universidad de San Carlos de Guatemala, Ediﬁcio T-10, Ciudad Universitaria Zona 12, Ciudad de
Abstract. Necrophagy is a feeding strategy in which animals feed on carrion; most scavengers are facultative and can also be
predators or consumers. For amblypygids, necrophagy is a poorly documented phenomenon and there are literature records of
individuals of three different species feeding on dead bats inside caves. In the present note, we document for the ﬁrst time a
necrophagic behavior in the whip spider Paraphrynus raptator (Pocock, 1902) which was observed feeding on Otonyctomys
hatti Anthony, 1932 (Rodentia: Cricetidae) and a yucatan poorwill, Nyctiphrynus yucatanicus Hartert, 1892 (Caprimulgi-
formes: Caprimulgidae) carrion. We made the observations inside a small chamber in an ancient Mayan temple inhabited by a
group of woolly false vampire bats (Chrotopterus auritus Peters, 1856) in southeastern Mexico. Carrion consumption in P.
raptator is directly related with the carnivorous feeding behavior of the C. auritus group with which they coexist.
Keywords: Amblypygid, carrion, facultative, scavenger.
Necrophagy is a feeding strategy that allows access to high-quality
food resources and it is deﬁned as the consumption of carrion by
animals. Only a few species feed exclusively on carrion; most of them
are facultative and can also be predators or consumers (Wilson &
Wolkovich 2011). Amblypygids are mainly predators of other
arthropods, although there are some records of them preying on
vertebrates (e.g., Owen & Cokendolpher 2006). In general, the diet
and feeding behavior of amblypygids are poorly documented in
nature (Weygoldt 2000; Chapin & Hebets 2016). There are only four
published records of necrophagy by three different species of
amblypygids (Table 1), all belonging to the family Phrynidae, feeding
on dead bats (Peck 1974; Armas & Abreu-Collado 1999; Garc´
Rivera et al. 2009; Prous et al. 2017).
In this note, we document the ﬁrst observation of necrophagy by a
fourth species of whip spider, and the ﬁrst time this behavior is
observed north of the Isthmus of Panama, in southeastern Mexico.
We made our observations at the ‘‘El Hormiguero’’ archaeological
site, in Ejido Eugenio Echeverr´
ıa Castellot II, south of Xpujil,
Campeche, Mexico (18.4085778N, 89.4902508W), speciﬁcally, inside a
small chamber in an ancient Mayan temple inhabited by a group of
six woolly false vampire bats Chrotopterus auritus (Peters, 1856).
Chrotopterus auritus is a large carnivorous bat that feeds on a great
variety of small vertebrates, such as doves, tanagers, warblers,
gekkonid lizards, rodents, shrews, mouse opossums, frogs and small
ın 1988, 1989; Bonato et al. 2004; Witt & Fabia
Frequently, individuals carry prey to their roost to consume it later or
share it with other group members. The prey are eaten from the head
down and during the process some parts are discarded (Medell´
1988, 1989). The discarded parts, along with feces, accumulate on the
roost ﬂoor, creating a suitable microhabitat for arthropods and
microorganisms that decompose the carrion and other organic
On April 26, 2019, we observed a whip spider feeding on a rodent
skin with muscle and fat attached. After being photographed, the
individual ran away, carrying the remains with it and hiding inside a
wall-cavity. Later, on July 11, 2019, we observed some bird remains
jammed inside the same wall-cavity. We collected both bird and
rodent remains that were being consumed by amblypygids in order to
identify the species. Based on fur characteristics and size and feather
patterns, Fernando Gual-Sua
´rez identiﬁed the rodent remains as
belonging to Otonyctomys hatti Anthony, 1932 (Cricetidae) (Pardi ˜
et al., 2017) and the bird remains as belonging to Nyctiphrynus
yucatanicus (Hartert, 1892) (Caprimulgidae) (Cleere, 1999). We have
found these and around 60 other species of vertebrates as part of the
diet of C. auritus, as well as some large invertebrates; however, we
have not found evidence of C. auritus feeding on P. raptator.
This is the ﬁrst time that bird and rodent necrophagy has been
documented for wild amblypygids. Before these records, dead bats
inside caves had been the only group of vertebrates documented
unequivocally in the necrophagic diet of whip spiders (Peck 1974;
Armas & Abreu-Collado 1999; Garc´
ıa-Rivera et al. 2009; Prous et al.
2017). However, there are other groups of vertebrates such as
amphibians and reptiles documented in the diet of amblypygids, but
in most cases, it is not known if these records correspond to predation
events or necrophagy (Chapin & Hebets 2016; Torres et al. 2019).
This issue is discussed by Owen and Cokendolpher (2006) in their
report of an amblypygid feeding on a hummingbird in which they
argue about the fact that they are not able to determine if the
amblypygid had actually hunted the hummingbird.
On July 13, 2019, one of the amblypygids inhabiting the chamber
was collected near the aforementioned wall-cavity. Usually, two or
three individuals can be seen on the walls upon entering the chamber.
We transported the individual to the Institute of Ecology, UNAM;
Rony E. Trujillo identiﬁed the specimen as an adult female of
Paraphrynus raptator (Pocock, 1902). Paraphrynus raptator is a
moderately large species (20–30 mm total length) distributed in the
U.S. (Florida), Mexico, Guatemala (Pet´
en), and Belize. This species is
a troglophile, is synanthropic and is adapted to a wide variety of
habitats, mainly in tropical humid to very humid forests, between 100
and 1500 m.a.s.l. (Armas 2006; Armas et al. 2018).
Necrophagy is a poorly documented phenomenon in amblypygids
and appears to be an opportunistic behavior: under suitable
conditions, individuals of some species can become facultative
scavengers. Carrion is a high-quality, donor-controlled resource
which depends on factors such as prey availability, hunting events
and subsequent prey consumption behavior (Wilson & Wolkovich
2011). In this particular case, carrion availability for individuals of P.
raptator depends completely on the predatory activity and prey
consumption behavior of the C. auritus group with which they
coexist. We hypothesize that P. raptator maintains a commensal
relationship with C. auritus, in which the latter provides a high-
quality resource on a regular basis. It is not known how frequently
these individuals of P. raptator feed on carrion, but based on the
knowledge that C. auritus takes food to their roost throughout the
ın 1988), we suggest that carrion represents an important
supplementary food resource for these amblypygids.
We would like to thank the people of ejido Eugenio Echeverr´
Castellot II and ejido 20 de Noviembre for their warm hospitality
during ﬁeldwork. We would also like to thank Abiael Illescas, Sabina
erez, David Herna
´ndez, Alejandra Gallegos and
Javier Torres for their help and effort during ﬁeldwork. Many thanks
to National Geographic Society for partially funding this study.
Finally, we would like to thank two anonymous reviewers and the
Table 1.—Records of necrophagy by amblypygids around the world.
Species Number Location Habitat Behavior Food Source
[identified by Peck as
2 Puerto Rico
Cave Feeding on dead
Unidentified bats Peck 1974
Cave Feeding on hanging
Unidentified bat Armas &
3 Cuba Cave Lifting carcasses Phyllonycteris poeyi
et al. 2009
1 Brazil (Amazon
Cave Lifting carcass Unidentified bat Prous et al. 2017
1 Mexico (Yucatan
Lifting and hiding
- Mexico (Yucatan
Lifting and hiding
Figure 1.—Paraphrynus raptator feeding on Otonyctomys hatti remains. Photograph: Fernando Gual-Sua
´rez. Low resolution (72 dpi) due to
the opportunistic nature of the observation; shot with iPhone XR 4mm, 1/48s, f/1.8, ISO-25, handheld, auto-ﬂash.
408 JOURNAL OF ARACHNOLOGY
editors Deborah Smith and Rick Vetter for their valuable comments
to improve this manuscript.
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Manuscript received 11 September 2020, revised 28 October 2020.
TRUJILLO ET AL.—NECROPHAGY BY THE WHIP SPIDER PARAPHRYNUS RAPTATOR 409