Bat Predation by Spiders: Martin Nyffeler, Mirjam Kno Rnschild

Bat Predation by Spiders
Martin Nyffeler
1
*, Mirjam Kno rnschild
2
1Section of Conservation Biology (NLU), Department of Environmental Sciences, University of Basel, Basel, Switzerland, 2Institute of Experimental Ecology, University of
Ulm, Ulm, Germany
Abstract
In this paper more than 50 incidences of bats being captured by spiders are reviewed. Bat-catching spiders have been
reported from virtually every continent with the exception of Antarctica (,90% of the incidences occurring in the warmer
areas of the globe between latitude 30u N and 30u S). Most reports refer to the Neotropics (42% of observed incidences),
Asia (28.8%), and Australia-Papua New Guinea (13.5%). Bat-catching spiders belong to the mygalomorph family
Theraphosidae and the araneomorph families Nephilidae, Araneidae, and Sparassidae. In addition to this, an attack attempt
by a large araneomorph hunting spider of the family Pisauridae on an immature bat was witnessed. Eighty-eight percent of
the reported incidences of bat catches were attributable to web-building spiders and 12% to hunting spiders. Large tropical
orb-weavers of the genera Nephila and Eriophora in particular have been observed catching bats in their huge, strong orb-
webs (of up to 1.5 m diameter). The majority of identifiable captured bats were small aerial insectivorous bats, belonging to
the families Vespertilionidae (64%) and Emballonuridae (22%) and usually being among the most common bat species in
their respective geographic area. While in some instances bats entangled in spider webs may have died of exhaustion,
starvation, dehydration, and/or hyperthermia (i.e., non-predation death), there were numerous other instances where
spiders were seen actively attacking, killing, and eating the captured bats (i.e., predation). This evidence suggests that spider
predation on flying vertebrates is more widespread than previously assumed.
Citation: Nyffeler M, Kno rnschild M (2013) Bat Predation by Spiders. PLoS ONE 8(3): e58120. doi:10.1371/journal.pone.0058120
Editor: Trine Bilde, Aarhus University, Denmark
Received November 15, 2012; Accepted February 4, 2013; Published March 13, 2013
Copyright: 2013 Nyffeler, Kno rnschild. This is an open-access article distributed under the terms of the Creative Commons Attribution License, which permits
unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Funding: The authors have no support or funding to report.
Competing Interests: The authors have declared that no competing interests exist.
* E-mail: martin.nyffeler@unibas.ch
Introduction
Bats have few natural enemies [1]. The most prominent bat
enemies mentioned in the scientific literature are owls, hawks, and
snakes [13]. Predation by a few large arthropods is occasionally
documented in the literature as well [35]. In a cave in Venezuela,
giant centipedes (Scolopendridae) have been observed killing and
eating mormoopid and phyllostomid bats [5]. Whip spiders
(Phrynidae) were observed feeding on dead phyllostomid bats in
caves of the Caribbean, but it is not known whether this were cases
of predation or scavenging ([6]; B. Fenton, pers. comm.). Despite
their name, whip spiders do not belong to the order spiders
(Araneae); instead they belong to the Amblypygi which is a
separate arachnid order. Furthermore, cockroaches have been
seen feeding on bat pups which have fallen to the floor [78]. In
several technical books on chiropterology, accidental deaths of
bats in spider webs have been reported [3,912]. The observation
of bat-catching by spiders is not that peculiar if we consider the
fact that a number of larger-sized spiders are known to supplement
their arthropod diet by occasionally preying on vertebrates.
Fishing spiders (Pisauridae) have been reported capturing and
devouring fish and frogs [1314]. Some species of wolf spiders
(Lycosidae), huntsman spiders (Sparassidae), tarantulas (Thera-
phosidae) and other mygalomorph spiders were observed killing
and eating frogs and lizards [13,1519]. Predation on snakes and
mice by tarantulas and comb-footed spiders (Theridiidae) has been
mentioned in the literature [13,20]. Furthermore, there are
numerous reports of birds being killed in the large orb-webs of
araneid and nephilid spiders, whereby the birds were either eaten
by the spiders or not [2129].
Deaths of bats in spider webs have been considered to occur
very rarely. In two more recent papers, a web-building spider,
Argiope savignyi, and a theraphosid spider, Poecilotheria rufilata, were
each reported to predate on a small bat [3031]. These authors
hypothesized that bat captures and kills due to spiders might be
more frequent than previously thought. To test this hypothesis, an
extensive global literature survey on bat-catching spiders was
conducted, along with an attempt to use web-based sources as
well. The insights from this research are reviewed here.
Methods
An extensive bibliographic search was conducted in order to
find any information available on bat-catching spiders. The search
was based largely on the Thomson-Reuters data base (Web of
Science), Google Scholar, Google Books, ProQuest Dissertations &
Theses, and Flickr image-hosting website (hosting more than 6
billion images). In addition to this, an internet search for blogger
information on this topic was conducted. Bloggers who had posted
photographs and reports on bat-catching spiders on the internet
were contacted to get detailed information on their observations.
Furthermore, the staff of bat hospitals was contacted to get
information on bats rescued from spider webs. Finally, an inquiry
among fellow arachnologists and chiropterologists was carried out
to get access to unpublished reports on this topic. Many of these
experts had conducted field studies for decades, and their feedback
provided valuable information needed to assess how frequent
PLOS ONE | www.plosone.org 1 March 2013 | Volume 8 | Issue 3 | e58120
incidences of bat catches by spiders might be. Altogether, 52
reports on bat-catching spiders could be gathered (Table 1).
Twenty-three of these reports have previously been published in
scientific journals, books, and a doctoral thesis.
During these inquiries we got access to several previously
unpublished photographs of bat-catching spiders. These photo-
graphs were shown to established bat and spider taxonomists for
identification of the bats and spiders, respectively. In a few cases
photographs of habitats were sent to vegetation specialists for
proper habitat classification. Nomenclature follows [3233].
Spiders reported in this paper are divided into two major groups
based on foraging mode (sensu Gertsch [14]): Hunting spiders
(i.e., spiders that forage without the use of a catching web) and
Web-building spiders (i.e., spiders that forage using a catching
web). Data on spider weight and size as well as bat weight,
wingspan, foraging mode and echolocation call frequency were
taken from the literature when available. Report numbers used in
the results, tables and figures refer to the respective detailed report
description (see File S1).
Results
Geographic Distribution of Bat Catches by Spiders
Incidences of bats being captured by spiders have been reported
from virtually every continent with the exception of Antarctica
(Table 1). Seventy-seven percent of the incidences listed in Table 1
were witnessed in regions of tropical climate between latitude
23u N and 23u S (,90% of the cases occurred between latitude
30u N and 30u S; Fig. 1). The prevalence of such events in the
warmer areas of the globe may be explained, among others, by the
fact that the vast majority of spider species capable of catching bats
are giant theraphosids, nephilids, and araneids who have their
main area of distribution in the tropical and subtropical regions
[20,3335].
There are six reports on bats being captured by hunting spiders.
Tarantulas (Theraphosidae) of the genus Avicularia have been
observed eating small bats in tropical rainforest areas in Peru
(Fig. 2A; report # 1) and eastern Ecuador (report # 2),
respectively ([18]; Rick West, pers. website; G. Schmid, flickr
image hosting website). A large tarantula of the genus Lasiodora was
observed eating a bat on the forest floor in northeastern Brazil (R.
West, pers. comm.; report # 3). Moreover, a large Reddish
Parachute tarantula, Poecilotheria rufilata, was reported to predate on
a small bat in Kerala, India ([31]; report # 4). Furthermore, a
huntsman spider Heteropoda venatoria (Sparassidae) was observed
capturing and killing a small bat in a shed near Kolkata, India
([36]; report # 5). This spider had apparently not fed on the bat
which may be explained by the fact that the observer interfered by
capturing spider and bat and placing them into a glass jar (see File
S1). An attempt by a large fishing spider Dolomedes triton
(Pisauridae) to kill a bat pup has been witnessed below a bridge
in Indiana, USA (P. Clem & V. Brack, pers. comm.; report # 6).
However, in this latter case the predation attempt failed probably
because the spider was frightened by the presence of the
photographing observers (see File S1). All other reports refer to
web-building spiders.
There are 19 reports of Neotropical bats being captured in
orb-webs of large araneids or nephilinids. These reports refer
to incidences witnessed in Peru (report # 7), Colombia (report
# 810), Guatemala (report # 11), Belize (report # 1213),
Costa Rica (report # 1420), Panama (report # 2124), or
elsewhere in Central America (report # 25) and are depicted in
Fig. 2BE, G and H. Several of these observations were made
by researchers stationed at the Los Amigos Biological Station
in Peru, the Biological Stations La Selva and La Sirena in
Costa Rica and the Smithsonian Tropical Research Station
Barro Colorado Island in Panama, which are all located in
lowland tropical rainforest ([8,30,3738]; M. Kno rnschild,
unpubl. data; M. Eckenweber, pers. comm.; M. Mallo & C.
D ez, pers. comm.; M. Nagy, pers. comm.; H & G. Unger,
pers. comm.; M. Weber, pers. comm.). In the incidences
witnessed in Costa Rica and Panama, the spiders had
constructed their webs on the outside of or in close proximity
to buildings inhabited by bat colonies ([8]; M. Kno rnschild,
unpubl. data; C. Metcalf, pers. comm.).
A second geographic region where bat-catching by web-
building spiders has frequently been reported (13 reports; 2I and
L) is eastern and southeastern Asia including locations in China
(report # 2634), Japan (report # 35), Vietnam (report # 36),
Malaysia (report # 37), and Sri Lanka (report # 38). Here, bat-
catching by spiders have been witnessed particularly often in the
eastern coastal area of China, specifically in parks and forests of
the Greater Hong Kong area ([3940]; G. Ades, pers. comm.;
C.S.K. Liu, pers. comm.; Fig. 2L).
A third geographic region where bat-catching by web-building
spiders was repeatedly witnessed (seven reports) is the area of
Australia (report # 3944) and Papua New Guinea (report # 45).
Most Australian incidences were observed in the coastal areas of
New South Wales and Queensland (Fig. 2JK).
Three incidences of bat captures by web-building spiders have
been reported from Africa ([4142]; D. Schultz, pers. comm.;
Fig. 2F; report # 4648). Only two incidences of bat catches by
web-building spiders witnessed in North America have been
reported so far ([4344]; report # 4950), and these both refer to
warm areas in the southern USA. Incidences of bat catches by orb-
weaving spiders are unknown from the northern part of North
America (B. Fenton, pers. comm.). Likewise, incidences of this type
have not been reported in the Ukrainian and Russian scientific
literature (A.T. Bashta, pers. comm.). Only two incidences of bats
being captured in spider webs have been reported from Europe
(report # 5152). In one case, a dead bat was found entangled in
the web of an orb-weaving spider on a building site near Stuttgart,
Germany (German tabloid BILD, May 2011). Another incidence
of a bat caught in a spider web was observed on the Isle of Wight,
South East England (G. Street, pers. comm.).
Which Spider Species are Involved in Bat Catches?
Bat-catching spiders belong to the araneomorph families
Nephilidae (golden silk orb-weavers), Araneidae (orb-weaver
spiders), Sparassidae (huntsman spiders), and the mygalomorph
family Theraphosidae (tarantulas). Furthermore, an attack attempt
by an araneomorph hunting spider of the family Pisauridae
(fishing spiders) was witnessed. Seventy-three percent of the known
incidences of bat catches were attributable to orb-weaving spiders,
15% to unidentified web-building spiders, and 12% to hunting
spiders (Table 1).
The dominant group of bat-catching spiders are giant orb-
weavers of the genus Nephila (Nephilidae). These spiders are forest-
dwellers that reach a legspan of 1015 cm and a weight of ,17 g
([34,45]; Table 2). They are diurnally and nocturnally hunting
[4647]; feeding was found to be most intense in the time between
sundown and midnight [46]. Nephila spp. spin strong webs with a
diameter of up to 1.5 m at a height of 1 to 6 m above the ground
[34,4748]. On certain locations, where females aggregate, several
webs are built connected to each other, which may result in a web
area of many square meters [49]. Of the 15 valid species in the
genus Nephila (see [34]) only two species - namely Nephila clavipes
and Nephila pilipes - have been reported so far to be engaged in bat
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catching. It can be assumed, however, that other Nephila spp. catch
bats as well. Up to the present, the Neotropical Nephila clavipes has
not been proven to feed on the bats captured in its webs. This
species is, on average, significantly smaller than the Asia-Pacific
species Nephila pilipes [5052]. One might speculate that the
smaller-sized N. clavipes perhaps is less successful in subduing bats
(H. Hofer, pers. comm.). It appears that the female Nephila spiders
depicted in Fig. 2H, I, and L were each missing a leg. Leg loss in
adult female Nephila spiders is a well-known phenomenon [53]. It
cannot be ruled out that some of the leg losses occurred during
aggressive encounters between spiders and bats trying to defend
themselves after being entangled in spider webs.
A second dominant group of bat-catching spiders are large orb-
weavers of the genus Eriophora (Araneidae) that spin strong vertical
orb-webs of up to 1.5 m diameter [8,3738,5455]. These spiders,
that reach a legspan of 58 cm and a weight of .1 g at favourable
web sites (Table 2), are forest-dwellers like the Nephila spp. The
Eriophora spp. are nocturnal spiders that hide in a leaf retreat
during the daytime hours and start spinning their webs just after
sunset [8,37,54]. They stay at the hub of the web all night long, the
majority of their feeding activity taking place at the beginning of
the night [56]. The genus Eriophora is represented in the Neotropics
with four species [33]. A congener from Australia, Eriophora
transmarina, has been witnessed catching bats as well (Table 1).
Several araneids (genera Eriophora and Caerostris) spin orb-webs of
extraordinary size and tensile strength suspended upon bridge
lines of several meters, which led to the assumption that such web
gigantism might have evolved as an adaptation for capturing flying
vertebrates such as bats and birds [5759]. In the case of Caerostris
darwini, an araneid which spins giant orb-webs of up to 2 m
diameter across rivers in Madagascar suspended upon bridge lines
exceeding 20 meters in length, chiropterophagy could not be
evidenced so far [5859]. However, in this latter study the sample
size of recorded predation events was rather small [5859]. In the
habitats of Caerostris darwini, several species of small riverine bats of
the families Vespertilionidae and Emballonuridae occur who
would be available as potential prey at least for a few weeks per
year during which time their volant juveniles (weighing ,2.54 g)
are within the spiders prey size range (S. Goodman, pers. comm.;
P.A. Racey, pers. comm.). Quantitative prey analyses of Caerostris
darwini at various seasons (with a high enough sample size to not
miss rare events) are urgently needed to answer the question
whether this spider is preying exclusively on larger-sized flying
insects (as suggested by [5859]) or whether it supplements its diet
by occasional catches of flying vertebrates.
Nephilengys cruentata, another bat-catching nephilid (Table 1),
weighs 0.51.6 g (Table 2). It is a synanthrophic spider that spins
orb webs (with up to 1 m diameter) on the corners of walls and
ceilings of buildings [6062]. Nephilengys might be less efficient in
bat catching. The only reported incidence where a bat got
entangled in a Nephilengys web ended with the bats successful
escape. Nevertheless this spider species should not be ruled out as a
potential predator of small bats; it is known to catch and eat
various types of vertebrates including small birds [2526,62].
Other orb-weavers reported to have caught bats in their webs
are found in the araneid genera Argiope and Parawixia (Table 1).
These lighter spiders, in comparison to Nephila spp. and Eriophora
spp., spin webs of much smaller size and may be engaged in bat
catches less frequently, although it must be noted that Argiope spp.
have repeatedly been observed killing (and sometimes eating) small
birds in North America and elsewhere (e.g. [21,29]).
Heteropoda venatoria, a sparassid spider reported capturing a
pipistrelle in India, has an adult legspan of 712 cm and weighs
between 26.5 g [6364]. Other larger-sized cave-dwelling
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Heteropoda spp. (with a legspan of 2030 cm), likewise suspected to
be predators of bats, are expected to be even heavier though body
mass data for these latter species are missing [6566]. Based on
their estimated size, the theraphosids Avicularia spp. engaged in
bat-eating may have weighed between 1030 g (compare [67] for
body mass data of theraphosids), whereas Lasiodora parahybana,
another bat-eating theraphosid, may have weighed even more
[68]. The theraphosid Poecilotheria rufilata that was observed
preying on a bat in India may have weighted between 2885 g
[31]. These giant sparassids and theraphosids are fully equipped
with the toxins and enzymes needed to subdue and devour small
vertebrate prey [1418,20]. The occasional capture of a bat should
therefore come as no surprise, though these spiders are actually
known to feed predominantly on arthropods [14,64,69]. Another
group of hunting spiders that might be capable of attacking and
killing small bats are a few large-sized species in the superfamily
Lycosoidea (e.g., genera Hogna and Dolomedes) whose adult females
weigh between 12 g [70]. Such huge hunters might occasionally
attack neonatal bats fallen from the roost. An incident of this type,
where a large pisaurid (Dolomedes triton) was attacking an immature
bat, was witnessed in Indiana, USA (report # 6).
Which Bat Species are Captured by Spiders?
So far, microbats from five families are known to have been
captured by spiders in the field (Table 1): Vespertilionidae (plain-
faced bats), Emballonuridae (sheath-tailed bats), Rhinolophidae
(horseshoe bats), Hipposideridae (Old World leaf-nosed bats) and
Phyllostomidae (New World leaf-nosed bats). In 31% of the
reported incidences the captured bats remained unidentified. The
majority of identifiable captured bats belonged to the families
Vespertilionidae (64% of reports) or Emabllonuridae (22%),
whereas only few reports existed for Hipposideridae (8%) and
one for Rhinolophidae (3%). The capture of phyllostomid bats was
only reported once (report # 20) or maybe twice (report # 11). In
the latter, uncertain report, concerning a small brown-coloured
bat entangled in a spider web in Guatemala (Fig. 2H; report #
11), the captured bat might have been a juvenile fruit-eating
phyllostomid bat though the features needed for a positive
identification were not sufficiently recognizable in the photo (A.
Gardner, pers. comm.). The Old World flying foxes (Pteropodidae)
have never been reported to get captured or killed by spiders
except for one report from captivity. Liat [71] reported that
neonates of the Lesser Dawn Bat, Eonycteris spelaea were offered to a
theraphosid, Coremiocnemis brachyramosa (subfamily Selenocosmii-
nae), in captivity. The neonatal bats were eaten readily by this
theraphosid. It is therefore possible that small or immature flying
foxes are caught and eaten by spiders in the field as well. Due to
the fact that the incidence reported by Liat [71] was not witnessed
in the field, it was not included in Table 1.
The majority of bats captured by spiders represented species
that are among the most common bats in their respective
geographic region (e.g., Pipistrellus spp. in various parts of the
world; Myotis nigricans and Rhynchonycteris naso in the Neotropics).
Nevertheless, rare species may sometimes get caught in spider
webs as well (e.g., Centronycteris centralis; compare [72]).
The majority of bats entangled in spider webs were small species
with a wingspan of 1024 cm and an adult weight of 38 g
(Table 3). It is noteworthy that adult bat weight can vary by several
grams depending on the bats feeding status and it is plausible that
only the lightest individuals of any given bat species get entangled
in spider webs; moreover, some of the captured bats were juveniles
or subadults (reports # 7, 1516, 2023, and 50) and thus
presumably much lighter than the weights for adults presented in
Table 3. Some of the entangled bats are among the smallest bats
on earth (e.g., Tylonycteris pachypus weighs only 3 g; [73]). Likewise,
the neonatal Myotis austroriparius bats found entangled in spider
webs in a study in Florida (see [43]) weighed only 3 g [74]. Large
bats are missing in Table 3 because of these bats capability to fly
right through a web (see below). Most of the species listed in
Table 3 are known to roost in buildings, caves, and tree holes.
Eighty-five percent of the reports with identifiably bats
entangled in spider webs refer to species classified as aerial
insectivorous bats (Table 4). Such bats (e.g., Pipistrellus spp. and
Myotis nigricans) feed aerially on flying insects; non-volant prey such
as spiders is almost entirely missing in their diets [7577]. In
contrast, there are some bats that feed heavily on web-building
spiders (e.g., Phoniscus papuensis, Myotis emarginatus, Myotis nattereri,
Myotis bechsteinii, Myotis keenii, Myotis lucifugus, Myotis aurascens; see
Figure 1. Geographic distribution of bat catching spiders worldwide. The map depicts the locations were spiders were observed catching
bats (red dots). Large red dots indicate that several reports originated from the same geographic region. Numbers refer to the detailed report
description (see File S1).
doi:10.1371/journal.pone.0058120.g001
[7885]). These are all bats with an average adult weight of 510 g
and a wingspan of 2127 cm [81,86]. In particular, the Golden-
tipped Bat (Phoniscus papuensis [formerly termed Kerivoula papuensis])
and Geoffroys Myotis (Myotis emarginatus) are considered to be true
spider specialists with spiders making up.75% of their total prey
[81,85,87]. These bats, which glean spiders from webs are
characterized by high flight agility and manoeuvrability at low
speed, using high frequency echolocation calls (.100 kHz start
frequency) to detect spider prey and to avoid accidential crashes
([81;88]; G. Jones, pers. comm.). One might expect such highly
specialized foragers to be sufficiently well adapted to avoid
collisions with spider webs. However, studies in New South Wales
(Australia) and Bavaria (Germany) revealed that gleaning insec-
tivorous bats (i.e., Phoniscus papuensis and Myotis emarginatus)
captured in harp traps and mist nets, had often spider web
material attached to their body fur and wing membrane,
indicating that these bats frequently strike spider webs while
gleaning spiders suspended in webs or while accidentally
encountering webs during their flights through the cluttered forest
vegetation [87,89]. In these latter studies, the web material must
have originated from smaller-sized spider webs that were not
strong enough to withstand the bats kinetic energy without
breaking (low energy absorbing webs sensu Craig [90]).
Analyses of faecal pellets likewise confirm that these bats prey on
orb-weavers of rather small size (,210 mm in length; [80,91]),
suggesting that they select smaller-sized webs for their gleaning
attempts. Quite a number of studies on spider-eating Myotis bats
were conducted in geographic areas such as Europe and northern
parts of North America [7879,8283,85,89,9194] where huge
araneid and nephilid spiders do not occur (compare [33]),
implying that in these areas gleaning bats presumably face little
danger of being caught and killed by web-building spiders. But in
some tropical areas of Australia, where Phoniscus papuensis does
occur sympatrically with huge nephilid spiders, incidences of
Phoniscus papuensis being accidentally ensnared and killed in Nephila
webs would be imaginable, though nothing has been reported
about this so far.
Figure 2. Bats caught by spiders. A - Adult female Avicularia urticans feeding on a Greater Sac-winged Bat (Saccopteryx bilineata) on the side of a
palm tree near the Rio Yarapa, Peru (photo by Rick West, Victoria, Canada; report # 1). B - Adult Proboscis Bat (Rhynchonycteris naso) entangled in a
web of Argiope savignyi at the La Selva Biological Station, northern Costa Rica (photo by Mirjam Kno rnschild, Ulm, Germany; report # 14). C - Dead
bat (presumably Centronycteris centralis) entangled in an orb-web in Belize (photo by Carol Farneti-Foster, Belice City, Belize; report # 12). D - Dead
bat (Myotis sp.) entangled in a web of Nephila clavipes in La Sirena, Corcovado National Park, Costa Rica (photo by Harald & Gisela Unger, Ko ln,
Germany; report # 17). E - A bat caught in the web of an araneid spider (possibly Eriophora sp.) in Tortuguero National Park, Costa Rica (photo by
Cassidy Metcalf, USA; report # 18). F - Live bat trapped in web of Nephilengys cruentata in a thatch roof at Nisela Lodge, Swaziland (photo by Donald
Schultz, Hollywood, USA; report #47). G - Volant juvenile Proboscis Bat (Rhynchonycteris naso) entangled in web of Nephila clavipes photographed in
a palm swamp forest near Madre de Dios, Peru (photo by Sam Barnard, Colorado Springs, USA; report # 7). H - Dead bat entangled in web of a
female Nephila clavipes in tropical rainforest in the middle of the Rio Dulce River Canyon near Livingston, Guatemala (photo by Sam & Samantha
Bloomquist, Indianapolis, USA; report # 11). I - Dead bat (Rhinolophus cornutus orii) caught in the web of a female Nephila pilipes on Amami-Oshima
Island, Japan (photo by Yasunori Maezono, Kyoto University, Japan; report #35). J, K - A small bat (superfamily Rhinolophoidea) entangled in web of
Nephila pilipes at the top of the Cockatoo Hill near Cape Tribulation, Queensland, Australia (photo by Carmen Fabro, Cockatoo Hill, Australia; report #
39). The spider pressed its mouth against the dead, wrapped bat, indicating that it was feeding on it. A Nephila pilipes male also present in the web
(K) may have been feeding on the bat as well. L - Dead vespertilionid bat entangled in the web of a female Nephila pilipes in the Aberdeen Country
Park, Hong Kong (photo by Carol S.K. Liu from AFCD Hong Kong, China; report # 32).
doi:10.1371/journal.pone.0058120.g002
Table 2. Fresh weight and body length (cephalothorax plus abdomen) of adult spiders reported to catch bats (arranged in
alphabethical order).
Spider species Weight [g] Body length [cm] Source report #
Araneus bilineatus unknown unknown - 26
Araneus heraldicus unknown unknwon - 27
Argiope savignyi ,0.5 * 1.31.8 [129] 1415
Avicularia sp. 1030 57 [67] 12
Dolomedes triton 1 2 [130] 6
Eriophora fuliginea 1.4 3 [101] 2224
Eriophora transmarina unknown 2.5 www.findaspider.org.au/find/spiders/105.htm 42
Heteropoda venatoria 26.5 24 [63] 5
Lasiodora parahybana up to .100 910 [68] 3
Nephila clavipes 13 23.5 [4647] 79, 11, 1617,
1921
Nephila pilipes 27 45 [51,53] 2833, 3536,
3840, 45
Nephilengys cruentata 0.51.6 2.5 [60,62] 47
Parawixia dehaani unknown 2 [131] 34
Poecilotheria rufilata 2885 6.5 [31] 4
Epeira bilineata and Epeira heraldica are now termed Araneus bilineatus and Araneus heraldicus and placed in the family Araneidae under Nomina dubia [33]. * Weight
estimated using data for similar-sized adult female Argiope argentata [132].
doi:10.1371/journal.pone.0058120.t002
Size Ratio of Web-building Spiders Versus Bats
Bats trapped in spider webs are usually small-sized (38 g adult
weight; Table 3), whereas the spiders capable of overpowering bats
are giant orb-weavers (,0.57 g weight; Table 2). The largest bat
species reported to have been captured in spider webs were an
adult male Nyctophilus gouldi (a species with a known adult weight of
11 g) and a Chalinolobus gouldii of unknown age (a species with a
known adult weight of 15 g; Table 3). The tensile strength and
elasticity of silk produced by large nephilid and araneid orb-
weavers is high (e.g. [95]), enabling such high energy absorbing
webs to retain flying vertebrates whose weight exceeds by far the
spiders own weight. Large orb-weaving spiders are generally
capable of trapping and killing prey that is much larger than
themselves [96]. For instance, a Nephila sp. was reported to have
captured a 3035 g bird, a Lewins Honeyeater, Meliphaga lewinii
[24]. Furthermore it has been reported that an 18 g Grasshopper
Sparrow, Ammodramus savannarum, was trapped in a spider web in
New York [24], while a 90110 g Black-faced Cuckoo-Shrike,
Coracina novaehollandiae, was ensnared in a spider web in Australia
[24]. The exact circumstances under which these relatively heavy
birds were trapped in spider webs are not reported. Flying
vertebrates of fairly large size might be retained if such an animal
crashes into an entire aggregation of large, strong Nephila webs.
Discussion
Are the Witnessed Incidences Real Predation Attempts?
It is arguable whether all incidences reported in this paper are
real predation attempts or whether some are just deaths by web
ensnarement without the active involvement of the spider (non-
predation deaths). Begon et al. [97] define the term predation as
follows Predation, put simply, is consumption of one organism
(the prey) by another organism (the predator), in which the prey is
alive when the predator first attacks it With other words, the
definition of predation implies that a prey item must have been
killed and eaten by the predator.
With regard to bat-eating theraphosids photographed in the
Neotropics and in India, the actual killing of the bats was not
witnessed. However, it has been proven by means of observations
in captivity that large theraphosids are capable of killing bats. This
is shown in a YouTube video where a Grammostola rosea (subfamily
Theraphosinae) is killing a small bat offered to it in a cage
environment (www.youtube.com/watch?v =kmGEoaHBhew; this
report was not included in Table 1 because it occurred in
captivity). The Grammostola rosea shown in this video was identified
for us by R. West (pers. comm.). Theraphosid venom has been
proven to be active on small mammals [20]. One can therefore
assume that theraphosid predation on bats is taking place in the
field as well and that the incidences witnessed in the Neotropics
and in India may have been cases of predation and not scavenging.
With regard to the sparassid Heteropoda venatoria, reported to
have killed a small bat in India [36], the situation is somewhat
Table 3. Fresh weight and wingspan of bat species reported to be captured by spiders (arranged in alphabethical order).
Bat species Weight (g) Wingspan (cm) Source report #
Centronycteris centralis 6 ,2427* [133] 12
Chalinolobus gouldii 15 33.1 [86] 41
Cyttarops alecto 6 unknown [127] 10
Eonycteris spelaea (neonatal) 8 N/A [134] captive evidence
Glossophaga sp. 11** 25.2** [86] 20
Hipposideros ater 46 22.8 [135136] 39, 44
Hipposideros pomona 58 26.0 [137138] 31
Myotis austroriparius (neonatal) 3 N/A [74] 50
Myotis nigricans 4 21.0 [86] 2, 2224
Myotis septentrionalis (immature) 2 N/A P. Clem & V. Brack, pers.
comm.
6
Neoromicia nana 34 20.6 [86,139] 48
Nycticeinops schlieffeni 5 22.4 [86] 46
Nyctophilus gouldi 11 31.1 [86] 43
Pipistrellus abramus 58 10.0 [140] 2627, 2930
Pipistrellus ceylonicus 8 25.6 [86] 4
Pipistrellus hesperus 4 19.0 [86] 49
Pipistrellus pipistrellus 5 21.8 [86] 52
Rhinolophus cornutus 4 18.0 [141142] 35
Rhynchonycteris naso 4 23.9 [86] 7, 1416
Saccopteryx bilineata 8 27.5 [86] 1, 21
Tylonycteris pachypus 3 19.0 [73] 37
The data refer to adults if not indicated otherwise. The species name Vespertilio irretitus has changed to Pipistrellus abramus (see [32]).
*Wingspan estimation based on data from Rhynchonycteris naso (smaller than Centronycteris centralis) and Saccopteryx bilineata (larger than Centronycteris centralis)
published in [86].
**Reported values refer to data from Glossophaga soricina.
different. Here the spider did catch and kill the bat, but the
sequence of behavioral units naturally displayed during a
predation event - starting with the attack of the prey and ending
with the cessation of feeding - was disrupted as the observer
interfered by capturing spider and bat and placing them into a
glass jar. As it appears, the dead bat was subsequently not preyed
upon (non-predation death). The spider in question might have
preyed on the bat had it not been disturbed by the observer.
Heteropoda spp. often occur in high abundance in close proximity to
bat roost sites in places like caves, buildings, and trees [63,98].
Especially in caves in Southeast Asia, large-sized Heteropoda spp.
cohabit with different species of bats [63,6566,69]. Large-sized
Heteropoda spp. are powerful enough to subdue small bats,
especially if one takes into consideration that they possess
impressive chelicerae, potent venom against small vertebrates,
and the ability to move at high speed [17,19,64]. Furthermore,
these spiders show morphological characteristics suggesting that
they may be highly adapted to climb vertical surfaces and walk on
cave ceilings [69,99]. The spiders may climb cave walls and
ceilings to catch perching bats or they may search for bat pups
fallen from the roost to the floor. It is readily imaginable that the
Heteropoda spp. supplement their staple diet, made up of arthropods
(i.e., in particular cave crickets, cockroaches, arachnids, and
centipedes; e.g. [64,69]), by the occasional catch of a small bat.
One way to find out whether Heteropoda spp. feed on bats would be
to video record the behavior of these spiders at locations where
they forage in close proximity to bat colonies. In 2006 (April to
December), such a study was undertaken at several pipistrelle roost
sites in the Christmas Islands, Australia; there, five video cameras
were deployed for a total of 663 trap nights [98]. Sparassid spiders,
Heteropoda venatoria, were the most frequently recorded potential
predators on the trunks of the roost trees, with 43 individual
records. The video recordings, however, did not reveal any of
these spiders preying on bats, disturbing them or entering their
roost sites. Currently we cannot confirm whether Heteropoda spp.
are habitually engaged in bat predation or whether the incidence
witnessed by Bhattacharya [36] was just a chance occurrence.
Further studies on possible interactions between sparassid spiders
and small-sized bats are therefore of considerable interest (also see
[100]).
The Neotropical orb-weaving spider Eriophora fuliginea has been
observed to kill and eat small bats that got entangled in its webs
[8,37,101]. When a bat got caught in a web, the spider
immobilized the bat by attack-wrapping and subsequently biting
it [101]. Following this, the spiders fed on the dead bats for many
hours (D.E. Wilson, pers. comm.). The incidences of bats being
caught, killed, and eaten in webs of Eriophora spp. are without any
doubt predation events.
Likewise, the orb-weaving spider Nephila pilipes, occurring in
Asia and Australia, has many times been observed to attack, kill
and eat small bats. Contrary to Eriophora, Nephila pilipes immobilizes
its prey by always first biting and subsequently wrapping it
[38,102]. The immediate attack bite of Nephila towards bats has
been witnessed by two authors [11,103], whereas the consumption
of bats was repeatedly observed/photographed ([3940,104]; C.
Fabro, pers. comm.; C.S.K. Liu, pers. comm.). There are several
reports where Nephila pilipes was seen killing and eating small birds
as well (e.g. [28]). Thus, the behavior of Nephila pilipes towards
small bats (catching, killing, and eating them), clearly complies
with the definition of predation.
The Neotropical orb-weaving spider Nephila clavipes has been
witnessed catching bats quite frequently (9 reports), but in none of
these cases was it seen biting, wrapping or eating a bat. Likewise,
birds trapped in the webs of this spider species were apparently not
consumed [2224,2627]. Only once has a Neotropical Nephila
been observed biting a bird, but without subsequent consumption
of the prey [22]. It has been suggested that Nephila clavipes might be
unable to deal with large, aggressive prey such as bats and birds
Table 4. Foraging mode and echolocation call frequency of adult bat species reported to be captured in spider webs (arranged in
order of increasing peak frequency).
Bat species Foraging mode
Echolocation call
peak freq [kHz] Source report #
Cyttarops alecto Aerial insectivore 36 [143] 10
Centronycteris centralis Aerial insectivore 41 [143] 12
Chalinolobus gouldii Aerial insectivore 41 [144] 41
Nycticeinops schlieffeni Aerial insectivore 42 [145] 46
Pipistrellus abramus Aerial insectivore 45 [147] 2627, 2930
Saccopteryx bilineata Aerial insectivore 45 and 47 [143] 1, 21
Pipistrellus pipistrellus Aerial insectivore 46 [146] 52
Myotis nigricans Aerial insectivore 54 [148] 2, 2224
Pipistrellus hesperus Aerial insectivore 62 [149] 49
Tylonycteris pachypus Aerial insectivore 65 [73] 37
Neoromicia nana Aerial insectivore 69 [145] 48
Nyctophilus gouldi Gleaning insectivore 72 [144,150] 43
Rhynchonycteris naso Aerial insectivore 98 [143] 7, 1416
Rhinolophus cornutus Gleaning insectivore 110 [80,151] 35
Glossophaga sp. Nectarivore 117 [152] 20
Hipposideros pomona Gleaning insectivore 133 [137,153] 31
Hipposideros ater Aerial insectivore 160164 [154] 39, 44
The species name Vespertilio irretitus has changed to Pipistrellus abramus (see [32]).
[24,27]. If this latter assumption is true, then the captures of bats
in the webs of Nephila clavipes would be cases of non-predation
deaths (the bats dying of exhaustion, starvation, dehydration, and/
or hyperthermia). The two European incidences where bats were
killed in spider webs without the spiders feeding on them must be
considered to have been cases of non-predation deaths as well.
In conclusion, some of the bats entangled in spider webs are
actively killed and consumed by the spiders (i.e., predation),
whereas in other instances the entangled bats are not consumed by
the spiders (i.e., non-predation deaths). In several of the
incidences, where dead bats were found suspended in spider
webs, it could not be determined whether predation had taken
place because of the bats desiccated condition (e.g. [44]).
How Frequent are Bat Catches by Spiders?
We conducted a survey of a large number of chiropterologists
and arachnologists who had conducted extensive field investiga-
tions in the tropics/subtropics to find out how many of them have
ever witnessed a spider catching a bat. A bat trapped in a spider
web is a fairly conspicuous sight that an experienced biologist will
hardly overlook, especially since it takes a spider several hours to
handle a bat prey. Only very few bat scientists (e.g. [8,30,105])
have ever witnessed an incidence of a bat caught by a spider, while
many others who also spent decades in the field (e.g., B. Fenton,
pers. comm.) have never seen this. Likewise, very few orb-weaving
spider experts (e.g. Robinson [101]) have witnessed spiders
catching bats. The 52 incidences reported in our review, which
refers to a time period of more than 100 years (starting with
Cantors report in 1842 [106]), is very low if this figure is
compared to estimates of bat mortality attributable to the bats
chief natural enemies. For instance, Speakman [2] estimated that
,200,000 bats per year are killed as a result of predation by owls
and kestrels in Great Britain alone. The fact that bat catching by
spiders has been witnessed so infrequently suggests that this type of
bat fatality is extremely rare. This is surprising because in tropical/
subtropical areas the millions of huge webs of Nephila spp. and
Eriophora spp. stretched across the bats flight paths pose an
enormous risk to bats (especially in locations where such orb-webs
are aggregated) and, actually, one would expect large numbers of
bats being killed each night. It appears that bats are evolutionarily
well-adapted to detect and avoid spider webs [107108]. Bats are
likely capable of detecting spider webs by means of echolocation.
Though single silk strands of spiders (,0.0020.005 mm in
diameter; [109]) are probably below the detection threshold of
echolocating bats (,0.16 mm for the aerial insectivore Eptesicus
fuscus; [110]; H.-U. Schnitzler, pers. comm.), the webs as a whole,
often containing additional conspicuous, densely-woven silk
decorations or silk barriers [54,107], may present themselves as
tangible objects that presumably bounce back echolocation calls
emitted by the bats with enough intensity to be detectable by the
bats [107]. Not only the webs, but also the spiders (in particular
the huge adult females of Nephila spp. and Eriophora spp. staying at
the hub of the web all night long) presumably generate strong
echoes that the echolocating bats should be able to detect,
advertising to them the presence of obstacles that need to be
avoided. In a study in Australia, Schultz & Wainer [87] captured
more than 1200 bats (representing eleven species) and checked
them for the presence of spider web fragments around their wing
membrane and/or on the body fur. These authors found that the
vast majority of bats did not have any traces of spider web
fragments attached to their body, strongly confirming the above-
mentioned hypothesis that bats are able to largely avoid
encounters with spider webs. This is also confirmed by a study
conducted in British Columbia where no spider web material was
found attached to the body of two species of Myotis bats trapped in
mist nets [83]. However, even if some bats collide with spider
webs, a considerable proportion of these bats may be able to elude
ensnarement by the following reasons: First, only large webs from
a limited number of giant orb-weaving spider species (especially
Nephila spp. and Eriophora spp.) are strong enough to withstand the
tremendous kinetic energy of a flying bat without breaking. Such
webs that intercept heavy and fast flying prey have been termed
high energy absorbing webs (sensu Craig [90]). If bats strike a
smaller web (i.e., low energy absorbing webs, designed to
intercept light and slow flying prey), the bats fly right through it,
leaving behind a damaged or destroyed web (also see [87,89]).
Second, only small bats can be retained in orb webs. If a larger-
sized bat strikes a web of any size, the kinetic energy is too high to
be absorbed and the web will break. Thus, larger-sized bats will
usually fly right through a web, leaving behind a big hole. The
same happens when larger-sized birds fly through spider webs
[54]. Third, even if small bats get entangled in large webs, a
certain percentage of violently struggling bats is able to escape
(D.E. Wilson, pers. comm.). The same is true for small birds that
may get temporarily entangled in spider webs but are often able to
free themselves after a short time of struggling [23]. It should be
noted that various species of bats behave differently when trapped
in a web, resulting in differing chances to escape. This is known
from experience with trapping bats in mist nets, which may be
looked upon as huge artificial spider webs. For instances,
Rhynchonycteris naso does not defend itself if trapped in a net,
whereas other sympatric bats of comparable size (e.g., Rhogeessa io)
struggle violently (M. Knornschild, unpubl. data). Accordingly,
chances for a Rhynchonycteris naso to escape from a large spider web
are slim compared to other bat species. Adaptation for escape
from spider webs also exists in various taxa of flying insects (e.g.
[111112]).
Nevertheless, some bats get caught and killed in spider webs.
Such cases might be considered to be accidents. Why do such
accidents happen? There may be several possible reasons for this:
First, it is noteworthy that the majority of the identifiable bats
(65% of reports) that accidentally crashed into spider webs
echolocate at frequencies of only ,3672 kHz (Table 4). It could
be that the echolocation calls of these aerial insectivorous species
(e.g., Pipistrellus spp. and Myotis nigricans) are less well-adapted to
detecting spider webs compared to the high frequency echolocat-
ing species (G. Jones, pers. comm.). Bats such as the pipistrelles
and Myotis nigricans are relatively fast flying and may not always be
able to avoid the webs if detected only at close range (G. Jones,
pers. comm.). Thus, though we hypothesize that most bats are able
to largely avoid encounters with spider webs (see above), accidents
do happen and bats echolocating with lower frequencies might be
engaged in these accidents with a higher likelihood. Second,
Griffin [113] theorized that bats flying in familiar territory rely
heavily on spatial memory and not on echolocation. This might be
the case when bats fly in proximity to the roost or when they use
flyways from the roost to the hunting ground and vice versa [114].
In such situations of heavy reliance on spatial memory, the bats
might not notice a spider web until they have already hit it. Indeed
several incidences reported in this paper (27% of reports) occurred
while the bats were flying in proximity to their roost (i.e., buildings,
caves, and forest trees [30,44]; M. Knornschild, unpubl. data) and
thus in spots where the bats presumably did not rely heavily on
echolocation. Third, some bats found entangled in spider webs
were juveniles or inexperienced subadults (reports #7, 1516, 20
23, and 50). As unskilled flyers, young bats may be more
susceptible to accidents than adults [115117]. A special case was
the incidence observed in Florida where bat pups of the
Southeastern Brown Bat got entangled in spider webs after falling
from the roost [43]. Bat pups falling from the roost frequently
become prey of various predators [7]. Fourthly, some bats may get
captured when they crash into an aggregation of large orb-webs.
Large orb-weavers of the genus Nephila tend do aggregate under
circumstances of high prey availability [49,118]. Blackledge et al.
[119] states: While reducing the overall number of prey that
might be intercepted, these spiders gain access to larger insects that
would normally break through a web as the insects either ricochet
off or slow down as they pass through exterior webs This is
exemplified by Sue Churchills observation at Tolmer Falls,
Australia, where an adult Hipposideros ater had apparently got
partially tangled in one web and then fallen into the other webs
before being fully caught (report #44). Fifthly, bats that habitually
glean insects from spider webs while hovering in front of them
[120] may sometimes get entangled after accidentially striking the
web. This evidently happens despite the fact that these bats,
echolocating at high frequencies (.100 kHz start frequency
[81,88]), are thought to be evolutionarily adapted to avoid
collision with spider webs. The Rhinolophus cornutus orii captured
in a Nephila web in Japan (Fig. 2I) may exemplify an incidence of
this type (report # 35). Bats from this genus sometimes strike
spider webs, evidenced by the fact that 4 out of 58 specimens of
Rhinolophus megaphyllus captured in harp traps in Australia had
spider web fragments attached to their wing membrane and body
fur [87]. The spider webs in this latter study were obviously not
strong enough to withstand the bats kinetic energy without
breaking (low energy absorbing webs). Sixly, during periods of
food scarcity, weakened small bats might be unable to free
themselves if trapped in spider webs, even if the webs are not as
strong as those spun by tropical large orb-weavers. The incidences
witnessed in Germany and England, where small pipistrelle bats
were killed in webs without any involvement of the spiders, might
present examples for this type of non-predation deaths.
At the present time nothing is known about the frequency with
which theraphosids and sparassids catch bats. The cryptic life
habits of these predominantly nocturnal hunters make them
difficult animals to study. Likewise, nothing is known about the
frequency of predation on bats by pisaurids.
Are Nephilid Webs at Cave Entrances a Threat to Cave-
roosting Bats?
In the tropics, huge nephilid orb-webs (genera Nephila and
Nephilengys) sometimes block the entrances to bat caves ([61]; C.
Dietz, pers. comm.). Such cave entrance inhabiting nephilid
populations have been discovered in East and South East Asia as
well as in the Neotropics. So far it is unknown to what extend
cave-roosting bats flying back and forth between caves and
foraging areas are able to detect and avoid these webs. Since it is
hypothesized that the bats might navigate by spatial memory while
passing through cave entrances [121], it is conceivable that some
of them may crash into nephilid webs within the caves entrance
zone, given the fact that in some areas they leave caves at dusk in
gigantic swarms. Monitoring nephild webs at cave entrances by
means of video recording devices could bring an answer to this
question.
How Important is Chiropterophagy from a Point of View
of Spider Nutritional Ecology?
All five groups of bat-catching spider taxa (Nephilidae,
Araneidae, Theraphosidae, Sparassidae, and Pisauridae) are
known to be predominantly predaceous on insects
[14,53,64,69,101]. With regard to large-sized theraphosids,
sparassids, and pisaurids, their feeding behavior in the field has
not been thoroughly investigated and one cannot currently judge
whether predation on bats is of significance to them from a feeding
ecological point of view.
Our current knowledge of orb-weaver feeding ecology suggests
that these spiders depend on flying insects as main prey, whereas
bats and also birds occasionally entangled in spider webs might be
considered to be by-catch [45]. It can take a large orb-weaver
many hours to consume a bat or bird prey ([26,28,30]; D.E.
Wilson, pers. comm.) indicating that spiders might extract a
substantial amount of energy while feeding on such a large prey.
Based on field observations in Papua New Guinea, Robinson &
Robinson [53] estimated the average capture rate of Nephila pilipes
at ,0.19 g wet weight prey killed per spider per day (which
corresponds to ,0.015 g dry weight food ingested per spider per
day). According to Higgins [46], the average prey capture rate of
the significantly smaller-sized Nephila clavipes in Texas was ,1.5
2.5 times lower than the estimate by Robinson & Robinson [53].
Thus, the catch of a ,2 g bat yields a Nephila pilipes a potential
prey biomass that is about 10 times the average daily prey catch.
In recent years, the idea has been proposed that the occasional
catch of large, energetically rewarding prey may be essential in
order to fulfil the reproductive needs of large orb-weaving spiders
(rare, large prey hypothesis; see [122123]). While large orb-
weavers such as Nephila spp. capture predominantly small insects of
little energetic value, they derive the bulk of their energy from a
few rare, large prey items (see [46,49,53,122]). In this context
rare, large prey encompasses large insects (e.g., cicadas, moths,
coleopterans, orthopterans, and odonates) as well as small flying
vertebrates (bats and birds). In our opinion, the examples of bat-
eating orb-weavers reported in this paper are consistent with the
"rare, large prey" hypothesis, though one may object to this given
the rarity of such events.
Acknowledgments
Many people have contributed greatly to this review paper. First
we wish to thank all those who kindly provided us with
unpublished information and/or photographs of incidences of
bat-catching spiders. Second we wish to thank all those scientists
who identified bats or spiders based on photographs. Alfred
Gardner, Don Wilson, Richard Laval, and Adriana Bravo
identified Neotropical bats. Marc Holderied and Tigga Kingston
identified a bat from Hong Kong. A group of Australian bat
scientists - including Kyle Armstrong, Mike Craig, Chris Pavey,
Terry Reardon, Martin Rhodes, and Christopher Tidemann -
identified a bat from Queensland, Australia. Nephilid spiders
depicted in photographs were identified by Matjaz Kuntner and I-
Min Tso. We wish to acknowledge William Eberhard and G.B.
Edwards for commenting on araneid spiders depicted in photo-
graphs or shown in film clips. Samuel Zschokke identified spider
webs based on photographs and a film clip. Rick West identified a
theraphosid spider shown in a film clip. Monika Rhodes was very
helpful by posting an information request on the listserver of the
Australasian Bat Society. Appreciation is also expressed to the
following people who helped us in various ways: Brock Fenton,
Thierry Gasnier, Steve Goodman, Hubert Hofer, Gareth Jones,
Brad Law, Markus Metz, Tadashi Miyashita, Paul A. Racey,
Hans-Ulrich Schnitzler, Chung-Tong Shek, John Speakman, Erik
Thorn, John O. Whitaker Jr. and many others who cannot all be
named here. Special thanks to Derek Yalden for valuable
suggestions and comments that helped to improve the manuscript.
Supporting Information
File S1 Detailed Reports Description.
(DOCX)
Author Contributions
Conceived and designed the experiments: MN. Performed the experi-
ments: MN. Analyzed the data: MN MK. Contributed reagents/materials/
analysis tools: MN. Wrote the paper: MN MK.
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