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Herpetological Review 48(3), 2017
AMBYSTOMA CALIFORNIENSE (California Tiger Salamander).
PREDATION. Although little appears to be reported on predation
of Ambystoma californiense, the larvae are known to succumb to
predatory attacks from Rana draytonii (California Red-legged
Frog; Baldwin and Stanford 1987. Herpetol. Rev. 18:33), Sterna
forsteri (Forster’s Tern), and Recuvirostra americana (American
Avocet; Allaback et al. 2005. Herpetol. Rev. 36:50). During sur-
veys for special-status species at stock ponds in the eastern San
Francisco Bay region (Contra Costa County, California, USA), we
encountered two native predators that are frequently sympatric
with A. californiense.
While conducting nighttime surveys on 90 ponds in the up-
per Kellogg Creek watershed we detected movement on the wet
edge of a pond (37.80140°N, 121.7163°W, WGS 84; elev. 210 m).
Upon close inspection we found a late-stage A. californiense lar-
va (approximately 10 cm total length) emerging from the aquatic
habitat. The larva had fully-formed gills and was also the color
and form of a life stage that suggested it should remain in the
pond for several more weeks (Alvarez and Foster 2016. Herp Na-
tion Magazine 19:28–30). When we attempted to collect the spec-
imen by hand for examination, we found an adult Actinemys pal-
lida (Western Pond Turtle) securely attached to the larvas caudal
fin (distal portion of the tail) in a firm bite. We grasped the turtle
and lifted it along with the larval A. californiense, and within one
or two seconds the turtle released the salamander. After a brief
inspection, the turtle was released. The A. californiense larva had
sustained a significant laceration to the distal portion of the tail,
deep enough to penetrate the skin and into the musculature. As
a result, a portion of the tail was partially disconnected, with a
gap of approximately 1 cm wide on the dorsal ridge of the tail.
Despite this substantial injury, the animal moved as would be
expected for a non-injured salamander, walking and dragging
its tail. The wound, despite its severity, did not appear to be life-
threatening and the larva was released back into the edge of the
pond where within two or three seconds it moved into deeper
water and out of sight.
At another pond (37.79000°N, 121.71880°W, WGS 84; elev.
319 m), we encountered a Thamnophis atratus (Aquatic Garter-
snake) moving through the vegetation along the shoreline, pre-
sumably preying upon the larvae of extremely numerous Pseud-
acris sierra (Sierra Treefrog) and Anaxyrus boreas ( Western Toad).
Our attention was drawn to commotion in the adjacent upland,
where a second adult Aquatic Gartersnake (approximately 60
cm) was partially tucked into a hoof print from grazing cattle. On
closer inspection, the snake appeared to be bracing itself as it at-
tempted to subdue and swallow a late-stage A. californiense larva
(approximately 13 cm total length). The prey item was grasped
by the head and periodically shook its tail from side to side, pre-
sumably attempting to escape. We observed the interaction for
several minutes before the A. californiense larva was swallowed
beyond its mid-section. At that point we presumed that the larva
would be fully consumed, and resumed surveying the remainder
of the pond. Preston and Johnson (2012. J. Herpetol. 46:221–227)
investigated the feeding preferences of T. atratus at this site and
others, and found that this species primarily feeds upon P. sierra
and A. boreas. Among 60 snakes and 258 food items reported by
Preston and Johnson (op. cit.), A. californiense was unrecorded in
the diet of T. atratus.
Our observations indicate that A. pallida and T. atratus will
attempt to prey upon larval A. californiense in aquatic breeding
habitat. In addition, the predatory incident with A. pallida oc-
curred at night (approximately 2100 h); this may have increased
the turtle’s potentially successful predation attempt over a day-
time event, when salamander larvae can more rapidly perceive
and evade predators.
JEFF A. ALVAREZ, The Wildlife Project, P.O. Box 188888, Sacramento,
California 95818, USA (e-mail: je; MARY A.
SHEA, P.O. Box 1632, Brentwood, California 94513, USA.
AMBYSTOMA CALIFORNIENSE (California Tiger Salaman-
der). TERRESTRIAL ECOLOGY. Although recent research has
begun to determine the extensive amount of terrestrial habitat
that Ambystoma californiense requires relative to breeding loca-
tions, little information is available regarding upland site fidel-
ity following breeding migrations (Trenham et al. 2005. Ecol.
Appl. 15:1158–1168; Searcy et al. 2011 in Alexander and Schlis-
ing [eds.], Research and Recovery in Vernal Pool Landscape, pp.
73–78. Chico State University, California; Orloff 2011. Herpetol.
Conserv. Biol. 6:266–276). During the winter of 2008–2009, while
conducting a mark-recapture study, we observed aspects of up-
land habitat use by A. californiense.
On 2 November 2008, we captured an adult A. californiense in
a pitfall trap 580 m from the only breeding location on the Buena
Vista Unit of the Ellicott Slough National Wildlife Refuge in Santa
Cruz County, California, USA (36.93530°N, 121.82640°W, WGS84,
elev. 132 m). The individual was captured within a narrow, 4 m-
wide strip of nonnative grass and landscaped plants between an
isolated single-family residence and a small, 0.21-ha drift fence
enclosure (Fig. 1). The male A. californiense was reproductive and
detected early in the season outside the enclosure during only
the second significant rainfall event of the year (> 2 mm). These
data, combined with the constrained capture location, indicated
that it over-summered within a few meters of the original cap-
ture site. The individual was released 37 m away on the opposite
side of the enclosure at an oak woodland/nonnative grassland
ecotone in the presumed direction of travel towards the breeding
pond. Much of the remainder of the property, including the land
between the release location and breeding pond, was considered
atypical for the species, as it consisted of dense coastal scrub,
closed-canopy coast live-oak woodland, closed-cone coniferous
forest, maritime chaparral, and non-native Acacia forest with
limited grassland or other open habitat.
Herpetological Review 48(3), 2017
On 6 February 2009, we captured the same A. californiense,
identified by a photograph of the dorsal spot pattern and a single
toe-clip, emigrating from the breeding pond where we were con-
ducting a mark-recapture population study of both A. californi-
ense and A. macrodactylum croceum (36.93050°N, 121.82650°W,
WGS84; elev. 89 m; Fig. 1). On 23 February 2009, the individual
was captured again in a pitfall trap at the upland enclosure with-
in 5 m of its initial release location.
Reproductive A. californiense will emerge from over-sum-
mering retreats early in the rainy season and may spend several
weeks or months to reach breeding sites. Ninety-six days elapsed
between the time this individual was trapped at the upland
site in November and when it was captured emerging from the
breeding pond. It subsequently exhibited upland site fidelity by
returning to its original upland location 17 d later. The round-
trip breeding migration for this individual required 113 d.
Upland habitats used by A. californiense are typically open,
including annual grassland and oak savannah, although chapar-
ral is used in Monterey County, California (Wang et al. 2009. Mol.
Ecol. 18:1356–1374). Our data show that A. californiense are able
to migrate at least 550 m through dense coastal scrub and closed-
cone coniferous forest during its breeding migration. Conserva-
tion planners must ensure that no barriers to terrestrial move-
ments are introduced that might inhibit breeding migrations or
post-metamorphic dispersal to specific upland areas where A.
californiense spends the majority of its life cycle. This observa-
tion supports the contention that focused studies are required
to estimate density in occupied uplands and identify movement
corridors (Searcy et al. 2008. Conserv. Biol. 22:997–1005), espe-
cially where A. californiense persists within a mosaic of vegetation
communities or in habitat patches fragmented by human uses.
This work was conducted under the authority of a United
States Fish and Wildlife Service Endangered Species Recovery
Permit, as well as California Department of Fish and Wildlife sci-
entific collecting permits and associated Memorandums of Un-
derstanding. We are grateful to Ellicott Slough National Wildlife
Refuge Manager Diane Kodama for reviewing this manuscript,
and Rachel Tertes, Meg Marriott, and Patrick Kearns of the Unit-
ed States Fish and Wildlife Service for acquiring funding and as-
sistance with drift fence installation.
M. LAABS, Biosearch Associates, P.O. Box 1220, Santa Cruz, California
95061, USA; CHAD STEINER, 221 Baldwin Street, Santa Cruz, California
95060, USA.
dile Newt). MORPHOLOGY. Spiny Crocodile newts of the genus
Echinotriton are very secretive and their ecology and natural his-
tory are poorly known (Hernandez 2016. Crocodile Newts: The
Primitive Salamandridae from Asia: the Genera Echinotriton and
Tylototriton. Edition Chimaira, Frankfurt, Germany, 415 pp.). The
recent description of a new species, Echinotriton maxiquadra-
tus, likewise encompasses a remarkably restricted geographic
range in the mountains of South-eastern China. It is possible
that the populations of these species are declining due to habitat
disturbance (Hou et al. 2013. Zootaxa 3895: 89–102). However,
despite the increasing attention on conservation of E. maxiqua-
dratus, nothing is known about its larval stage and morphology.
Such information is crucial to protect breeding habitats for this
endangered species.
On 11, 12, and 13 July 2015, seven larvae were discovered
in three small ponds of a subtropical perhumid evergreen
Fig. 1. Initial and recapture locations of adult male Ambystoma
californiense relative to the breeding pond, Buena Vista Unit, Ellicott
National Wildlife Refuge, Santa Cruz County, California, USA.
Fig. 1. Echinotriton maxiquadratus larvae and their habitat in south-
western China. Upper left: old larva of 43 mm; Upper right: hatching
larva of 16 mm; Lower: Small pond covered with dense vegetation
were larvae were found.
Herpetological Review 48(3), 2017
broad-leaved forest of a mountain range in Southwestern China
between Zhejiang, Fujian, Jiangxi and Guangdong provinces at
1240 and 1320 m elev. Coordinates are not provided for conser-
vation purposes, following the suggestions of Hou et al. (2013, op.
cit.). The air temperature was 19.6°C at night and 26.6°C during
the daytime, that of the water 21.2°C (with larvae), and humidity
79%. The average depth of water was 6–10 cm. pH recorded at the
spawning site was between 7.3–7.6. All of these ponds were veg-
etated by Sphagnum moss and grasses, and populated by seven
larvae of 1.6 to 4.8 cm in total length, which implied they were ca.
4–6 weeks old when assuming a growth rate similar to E. chin-
haiensis (Hernandez 2016, op. cit.). The hatching larvae measure
16 mm in total length, are yellowish to orange in color and turn
pale grey with darker black coloration on the dorsum with age.
Gills are present and show a reddish coloration. Larvae also have
small white spots disposed irregularly on the dorsal part and gills.
Brown to dark spots are also present on the tail and disappear
with age (Fig. 1). The larvae were assigned to E. maxiquadratus
based on genetic testing (Hernandez et al., in press). Moreover,
I also found two adults by night depositing eggs near the ponds
at the studied sites. These observations suggest that larval mor-
phology of E. maxiquadratus is quite similar to its congeners but
differs by the bright yellowish coloration of hatching larvae and
by having more small irregular white spots on the dorsum.
AXEL HERNANDEZ, Department of Environmental Sciences, Faculty
of Sciences and Technics, University Pasquale Paoli of Corsica, France; e-
EURYCEA AQUATICA (Brown-backed Salamander). NESTING.
On 8 March 2016, we flipped a large rock to reveal two separate
Eurycea aquatica nests numbering 67 and 80 eggs in a spring-fed
stream in Knox County, Tennessee, USA (35.93830°N, 83.84720°W;
WGS 84). Under a second smaller rock that was directly adjacent
and under the first, we found a third nest numbering 89 eggs. We
determined the sex of attending adults by the presence of mental
glands in males. With these three nests, we found a total of one
adult female and one adult male E. aquatica. Nearby, we flipped
another large rock and the several smaller rocks underneath it.
In total, these rocks revealed six nests and five adult female E.
aquatica, as well as two dead and decomposing gravid females.
In all cases, determining exactly which adults were with which
nests was impossible due to the flow of the stream and disrup-
tion caused by lifting the rocks. We photographed and counted
eggs from two of these nests, and they numbered 154 and 80, re-
Proposed communal nesting has been previously reported
for other populations of the Eurycea bislineata species complex,
including some in New York (Bishop 1941. New York State Mus.
Bull. 324:1–365), Virginia (Wood 1953. Chicago Acad. Sci., Nat.
Hist. Misc. 122:1–7), Ohio (Baumann and Huels 1982. J. Herpe-
tol. 16:818–823), Illinois (Jakubanis et al. 2008. Northeast. Nat.
15:131–140), Ontario (LeGros 2011. Can. Field-Nat. 125:363–365),
and Connecticut (Ferguson et al. 2014. JNAH 2014:87–92), al-
though some of these reports fail to distinguish between nests
laid under the same object but separated by a small distance and
those truly intermixed.
The taxonomy of the E. bislineata species complex is con-
tentious, and our taxonomic assignment of this population to E.
aquatica reflects the best application of the available taxonomy
to evolutionary relationships suggested by genomic data (Pier-
son et al., unpubl. data). Graham et al. (2010. IRCF Reptiles &
Amphibians 17:168–172) report on the nesting behavior of E.
aquatica, demonstrating both male and female attendance of
clutches, with a mean clutch size of 65.93 (range 31–138). They
speculated that their largest clutches might represent evidence
of communal nesting, but they did not provide definitive proof.
We likewise cannot be certain as to whether any of our individual
nests contain eggs from different females, but the large size of
one nest suggests the possibility. Regardless, the close proximity
and dense aggregation of nesting females we observed is remark-
able even for members of the E. bislineata species complex, and
we do not know of any previous records of deceased females in
nesting aggregations.
SON, Department of Ecology and Evolutionary Biology, University of Ten-
nessee, Knoxville, Tennessee 37996, USA (e-mail:
EURYCEA AQUATICA (Brown-backed Salamander) and EURY-
CEA CIRRIGERA (Southern Two-lined Salamander). HATCH-
ING. On 13 March 2016, we lifted a small rock in a spring-fed
stream to reveal an adult female Eurycea aquatica attending
a clutch of eggs in Knox County, Tennessee, USA (35.95239°N,
83.88860°W; WGS 84). We momentarily positioned the rock at a
90° angle, with the nest suspended vertically in shallow, slowly
flowing water. Upon first examination, the nest consisted of ap-
proximately 40 eggs, at least half of which were occupied by vi-
able larvae. Within one minute, we noticed several of the larvae
within the eggs squirming vigorously and with increasing inten-
sity. In the next 5 min., we watched as all but one larva pushed
their way out of the eggs and swam into the water. Upon hatch-
ing, these larvae moved immediately to the gravel at the stream
bottom and attempted to find refuge in interstitial space. On 29
March 2016, we uncovered a female Eurycea cirrigera guarding a
clutch of eggs under a rock in a second-order stream in Haber-
sham County, Georgia, USA (34.50396°N, 83.48103°W; WGS 84).
This nest consisted of at least 44 eggs, and we did not notice any
eggs that had already hatched. We temporarily flipped the rock
upside down and rested the rock out of the water, and several of
the larvae began to hatch. We placed the rock in shallow, slowly
flowing water at a 90° angle. Within 5 min., approximately two-
thirds of the larvae had hatched and buried themselves in the
silt below the rock. Warkentin (2011. Integr. Comp. Biol. 51:111–
127) reviewed plasticity of hatching in amphibians in response
to environmental stimuli, including disturbance by potential
predators. However, relatively few observations of the hatching
of aquatic plethodontid salamander nests have been recorded
(Petranka 1998. Salamanders of the United States and Canada.
Smithsonian Institution Press, Washington D.C. 587 pp.), and ex-
pedited hatching in response to a potential predator is not yet
documented. We acknowledge that we may have happened to
uncover these nests in the process of hatching, but the speed with
which the larvae hatched seems instead to suggest a response to
the disturbance similar to that exhibited by other amphibians.
The taxonomy of the Eurycea bislineata species complex is under
investigation, and our identifications reflect the best application
of the available taxonomy to evolutionary relationships suggest-
ed by genetic and genomic data (Timpe et al. 2009. Mol. Phylo-
genet. Evol. 52:368–376; Pierson et al., unpubl. data).
TODD W. PIERSON, Department of Ecology and Evolutionary Biology,
University of Tennessee, Knoxville, Tennessee 37996, USA (e-mail: tpierso1@; NATALIA J. BAYONA-VÁSQUEZ, Laboratorio de Genética
de Organismos Acuáticos, Instituto de Ciencias del Mar y Limnología,
Universidad Nacional Autónoma de México, México D.F. 04510, México
Herpetological Review 48(3), 2017
DICAMPTODON TENEBROSUS (Coastal Giant Salamander).
FEEDING BEHAVIOR AND DIET. Dicamptodon tenebrosus is
an opportunistic predator that eats a wide variety of prey (Nuss-
baum et al. 1983. Amphibians and Reptiles of the Pacific North-
west. University Press of Idaho, Moscow. 332 pp.; Petranka 1998.
Salamanders of the United States and Canada. Smithsonian In-
stitution Press, Washington, DC. 587 pp.). However, terrestrial
earthworms (Class Oligochaeta: Order Megadrilacea) are rarely
reported as part of the diet of wild D. tenebrosus. Here, we relate
observations of D. tenebrosus feeding on two species of terres-
trial earthworms (Lumbricidae). These observations were made
along small forest streams in northwestern Oregon, USA, which
are described in detail elsewhere (Rombough 2017. Herpetol.
Rev. 48:154–155). Several species of earthworms, including Lum-
bricus terrestris and Octolasion cyaneum, are locally abundant in
this area.
Between 1840 and 2120 h on 30 October 2014, we observed
four terrestrial (metamorphosed) D. tenebrosus (SVL = ca. 80 to
> 120 mm) feeding along the edge of small streams in the afore-
mentioned area. These Dicamptodon were between 2 and 8 m
from the stream when first observed. All were walking slowly over
the surface; periodically, they were seen lowering their heads
into the moss and wet leaves that covered the substrate. At ca.
2100 h, one of these Dicamptodon (SVL = ca. 120 mm) caught
a L. terrestris that was crawling over the wet leaves. The worm
thrashed and attempted to crawl away with its anterior end, but
the Dicamptodon swallowed it using quick jerks of its head. The
entire process took <5 min.
On 07 February 2015, between 1930 and 2200 h, we observed a
larval (SVL = ca. 80 mm SVL) and two gilled adult (neotenic; SVL =
ca. 120 and 150 mm) D. tenebrosus feeding on L. terrestris in a small
stream. All Dicamptodon were in the shallow margins (depth < 20
cm) of slow pools below debris (leaf and stick) jams, less than 30
cm from the stream bank. Several additional Dicamptodon lar-
vae and gilled adults (SVL = ca. 50 to > 100 mm) were seen along
the shallow margins of similar pools; L. terrestris experimentally
dropped into these pools were immediately caught and eaten.
At 2115 h, we found a larval D. tenebrosus (SVL = ca. 80 mm)
on the gravelly bank of the stream, approximately 15 cm from
the water. As we watched, it moved forward and caught an Oc-
tolasion cyaneum (Fig. 1). The Dicamptodon immediately swal-
lowed the anterior end of the worm. However, the worm was
too large to eat quickly, and a battle ensued, in which the worm
repeatedly dug into the gravel with its posterior end, dragging
the Dicamptodon’s head down into the gravel, and partially
pulling itself out of the Dicamptodon’s mouth. Periodically, the
Dicamptodon jerked its head back, pulling the worm back out
of the gravel, and re-swallowing the escaped portion. This pro-
cess continued for ca. 15 minutes, at which point we retreated
so as not to disturb the salamander further. When we checked
on the salamander approximately 10 minutes later, it was still on
the bank, but we did not see the worm. At the time observations
were made, the weather was cool (air temp. = 11°C) and rainy,
and both L. terrestris and O. cyaneum were abundant on the soil
surface and in leaf packs along the stream.
On 21 February 2016, between 2100 and 2400 hours, we ob-
served a larval (SVL = ca. 70 mm) and a gilled adult (SVL 90
mm) D. tenebrosus feeding on L. terrestris that had washed into
a small stream (Fig. 2). Both Dicamptodon were in shallow (15
cm), slower-moving portions of the stream. At the time obser-
vations were made, the weather was cool (air temp. = 7°C) and
rainy, and L. terrestris were abundant on the soil surface and in
leaf packs along the stream.
To the best of our knowledge, this report is the first account
of larval Dicamptodon feeding out of water. Interestingly, both
species of worms described here are not native to North America,
but are introductions from Europe. Most existing studies do not
mention lumbricid worms as items of Dicamptodon diet, but this
may be an artifact of the studies’ timing or location. An indica-
tion that this phenomenon may be increasing and/or underes-
timated is the work of Parker (1994. Copeia 1994:705–718), who
studied the diet of larval Dicamptodon in a stream in northwest
California. Parker reported that, on two occasions following
heavy rains, “a large proportion of salamanders collected had
consumed terrestrial earthworms (Lumbricus sp.) that had been
washed into the stream and dominated total diet volume”.
We owe special thanks to Bill Fender of McMinnville, Oregon,
for identifying O. cyaneum.
CHRIS ROMBOUGH, Rombough Biological, PO Box 365, Aurora, Or-
egon 97002, USA (e-mail:; LAURA TRUNK, Jack-
son Bottom Wetlands Preserve, 2600 SW Hillsboro Hwy, Hillsboro, Oregon
97123, USA.
Fig. 1. Larval Dicamptodon tenebrosus eating Octolasion cyaneum on
the rocky bank of a small stream.
Fig. 2. Neotenic Dicamptodon tenebrosus eating Lumbricus terrestris
in the shallow margins of a stream. (The salamander pictured com-
pletely swallowed the Lumbricus within ca. 5 min.)
Herpetological Review 48(3), 2017
PLETHODON CINEREUS (Eastern Red-backed Salamander).
HABITAT. Plethodon cinereus is a woodland salamander primar-
ily found in wooded areas and forests of northeastern United
States and southeastern Canada (Petranka 1998. Salamanders
of the United States and Canada. Smithsonian Institution Press,
Washington, D.C. 587 pp.). As wooded areas and forests become
more fragmented, understanding whether P. cinereus can use or
traverse non-wooded habitats may become more important in
determining the ability of this species to respond to the increased
isolation of wooded fragments. Previous research has shown
that P. cinereus can move limited distances across non-wooded
habitats. For example, Marsh et al. (2004. Ecology 85:3396–3405)
found that P. cinereus displaced up to 55 m into a field habitat
were able to return to a forest habitat, and that P. cinereus would
cross open field habitat to reach islands of wooded habitat up to
25 m from a forested edge.
On 2 October 2016, we observed an adult P. cinereus (SVL
ca. 4–5 cm) under an artificial cover board (61 m x 61 m x 2 cm)
in a restored prairie habitat at the Granville Schools Land Lab,
Granville, Licking County, Ohio, USA (40.08888°N, 82.54083°W;
WGS84). The cover board had been originally placed in Spring
2016 as part of a larger grid of cover boards used to monitor snake
and small mammal distributions on the Granville Schools Land
Lab, a 15-ha restored prairie/wetland system that was reclaimed
from agricultural field (soybean and corn rotation) beginning in
the summer of 2014. The two nearest wooded habitats are 300 m
and 500 m from the cover board, suggesting the possibility the
salamander moved that far from the nearest suitable habitat. The
longest dispersal movement of P. cinereus observed to date is 143
m through a forested habitat (Sterrett et al. 2015. Herpetol. Rev.
46:71). In addition to the distance, the salamander would have
crossed at least one two-lane road or school driveway to reach
the cover board site from the nearest woodlands. Roads are of-
ten significant barriers to the movements of P. cinereus (Marsh et
al. 2005. Conserv. Biol. 19:2004–2008, Marsh et al. 2008. Conserv.
Genet. 9:603–613). We did not subsequently observe any sala-
manders under any cover boards, suggesting this salamander
may have been transiting the area, perhaps dispersing among
woodlands or engaging in a seasonal migration (e.g., Woolbright
and Martin 2014. J. Herpetol. 48:546–551) and/or was not suc-
cessful at colonizing the prairie habitat.
We thank J. Reding for his driving the Land Lab restoration
project, which was funded by the U.S. Fish and Wildlife Service’s
Partners for Wildlife Program. We also thank J. Rettig and L. Smith
for help in placing the cover boards.
GEOFFREY R. SMITH, Department of Biology, Denison University,
Granville, Ohio 43203, USA (e-mail:; WESLEY O.
SMITH, Granville High School, 248 New Burg St. Granville, Ohio 43023, USA.
PLETHODON PETRAEUS (Pigeon Mountain Salamander).
ARBOREAL BEHAVIOR. Plethodon petraeus is a species of
conservation concern because it is only known to occur along
approximately 17 km of the southeastern slope of Pigeon
Mountain in northwest Georgia. Described as a crevice-
dwelling salamander, climbing behavior in in this species is well
documented on rocky outcrops, cliff faces, and cave entrances
(Wynn et al. 1988. Herpetologica 44:135–143; Jensen et al. 2002.
Southeast. Nat. 1:3–16). Despite morphological adaptations for
climbing and ample evidence of utilizing vertical exposed rock,
there have been no reports of individuals on trees.
At 2200 h on 21 April 2016 within Crockford-Pigeon Mountain
Wildlife Management Area in Walker County, Georgia, USA
(precise locality withheld due to conservation concerns) a
female adult P. petraeus was observed 45 cm above the ground
on the side of a moss-covered hardwood tree (Fig. 1). The tree
was within 1 m of a rock outcrop. During this encounter, the
individual was seen consuming an ant, a large component of
this species’ diet (Jensen and Whiles 2000. J. Elisha Mitchell
Sci. Soc. 116:245–250). Although it was not raining at the time
of the observation, the location had received rainfall within the
previous 24 h. This observation adds to the knowledge of this
species’ foraging habits beyond the rock crevices they are closely
associated with. It also adds to the increasing documentation of
facultative use of vegetation by plethodontid salamanders for
foraging (McEntire 2016. Copeia 104:124–131).
KATE C. DONLON, 2021 Coey Rd, Columbus, Ohio 43210, USA
(e-mail:; THOMAS C. McELROY, 1000 Chastain Rd,
Kennesaw, Georgia 30144, USA (e-mail:
BEHAVIOR. Plethodon wehrlei is a little-studied species with
a distribution largely within the Central Appalachian Plateau.
Although Green and Pauley (1987. Amphibians and Reptiles
of West Virginia. University of Pittsburgh Press, Pittsburgh,
Pennsylvania. 241 pp.) note that P. wehrlei may occasionally
ascend vegetation, a recent review on the arboreal ecology of
Plethodontidae (McEntire 2016. Copeia 104:124–131) did not
report arboreal behavior in P. wehrlei. Observations of arboreal
behavior should be noted, as it may have potential implications
Fig. 1. Plethodon petraeus foraging on the side of a tree, Crockford-
Pigeon Mountain Wildlife Management Area, Walker County,
Georgia, USA.
Herpetological Review 48(3), 2017
for plethodontid research and conservation that occurs within
the range of P. wehrlei as traditional sampling methods are often
biased towards terrestrial behavior.
At 2200 h on 8 August 2013, numerous (> 5) P. wehrlei were
noted climbing woody vegetation (e.g., Rhododendron maxi-
mum, Great Laurel, and Betula alleghaniensis, Yellow Birch)
alongside a forest road in Monongahela National Forest, Ran-
dolph County, West Virginia, USA (38.63000°N, 79.88000°W, WGS
84, elev. 1065 m) following a rain event. Immediately after, a 1 x
20 m transect was searched for approximately 30 min. The tran-
sect was heavily vegetated with Yellow Birch and Great Laurel
and had minimal herbaceous ground cover. Air temperature was
approximately 20°C , humidity > 90%, and foliage and leaf litter
were wet. Twenty-seven salamanders were recorded, including
18 P. wehrlei, two P. glutinosus, five P. cinereus, and two Desmog-
nathus ochrophaeus. Twelve of the 18 (66%) P. wehrlei were found
on woody vegetation (primarily branches of great laurel; Fig. 1) at
a height of 0.5–1.5 m. The only other salamander found exhibit-
ing arboreal behavior was a single D. ochrophaeus.
This observation highlights that vertical structure may be
more important to P. wehrlei than previously noted and efforts
should be made to adequately sample and address this habitat
in future studies.
MICHAEL GRAZIANO, 210 Kottman Hall, School of Environment and
Natural Resources, The Ohio State University, Columbus, Ohio 43210, USA;
PLETHODON YONAHLOSSEE (Yonahlossee Salamander). AR-
BOREAL BEHAVIOR. At 2314 h on 27 August 2016, we observed
an adult male Plethodon yonahlossee approximately one m off
the ground on a small, dead hemlock tree in Smyth County, Vir-
ginia, USA (36.71450°N, 81.51010°W; WGS 84). The salamander
had a large mental gland and sat vertically along the trunk with
its head lifted just off the tree (Fig. 1). A review of climbing and
arboreal behavior of plethodontid salamanders (McEntire 2016.
Copeia 104:124–131) reported many instances of other Plethod-
on climbing trees, but to the best of our knowledge, this is the
first documentation of this behavior in P. yonahlossee.
TODD W. PIERSON, Department of Ecology and Evolutionary Biol-
ogy, University of Tennessee, Knoxville, Tennessee 37996, USA (e-mail: tpi-; WILL LATTEA (e-mail:, EVIN
T. CARTER, Department of Ecology and Evolutionary Biology, University
of Tennessee, Knoxville, Tennessee 37996, USA (e-mail: ecarte19@vols.; LINDSEY HAYTER, Admiral Veterinary Hospital, 204 Watt Road,
Knoxville, Tennessee 37934, USA (e-mail:
Taricha granulosa is well-known for its toxicity to most verte-
brates (Petranka 1998. Salamanders of the United States and
Canada. Smithsonian Institution Press, Washington, DC. 587
pp.). Here, I relate an account of predation on an adult T. gran-
ulosa by Anas platyrhynchos (Mallard Duck). At 1100 h on 20
January 2014, I saw a mixed flock of ducks which included 26 A.
platyrhynchos feeding on a small farm pond in Marion County,
Oregon, USA (45.2322°N, 122.7894°W, WGS 84; elev. 55 m). I
watched the ducks from approximately 18 m away, using a pair
of 10 x 25 binoculars. As I watched, a female A. platyrhynchos
dipped her head underwater and caught an adult T. granulosa
(ca. 75 mm SVL). The duck held the T. granulosa in its bill, with a
grip just anterior to the hind legs. The bright orange venter of the
T. granulosa was clearly visible, even without binoculars.
The duck spent the next five minutes manipulating the
squirming T. granulosa: shaking it, dipping it in the water, toss-
ing it in the air and catching it by a different part of the body. The
latter movement appeared to be an attempt to re-position the
T. granulosa in its bill. After five minutes, the duck managed to
secure a grip on the T. granulosa’s head. At this point, the duck
swallowed the T. granulosa head-first, a process that required al-
most a minute and involved much head-tossing and thrusting
of its bill into the air. During this entire process, the duck con-
tinued swimming around on the pond. Using my binoculars, I
watched the duck very closely for the next two hours. It spent this
time swimming and preening in the same manner as the other A.
platyrhynchos present. It showed no apparent ill effects, nor did
it exhibit any behavior that I could distinguish as different from
Fig. 1. Yonahlossee Salamander (Plethodon yonahlossee) climbing a
tree, Smyth County, Virginia, USA.
Fig. 1. Wehrle’s Salamander (Plethodon wehrlei) exhibiting arboreal
Herpetological Review 48(3), 2017
that of the other ducks in the flock. After two hours, all the ducks
flew away.
Unfortunately, I do not know the fate of the duck in this observa-
tion. Nonetheless, it is interesting that the duck was able to consume
the newt, and that it did not exhibit symptoms of poisoning for at
least two hours. Birds, including A. platyrhynchos, are reported to
be highly susceptible to the toxin of T. granulosa (Storm 1948. Her-
petology of Benton Co., Oregon. Ph.D. Thesis, Oregon State Univ.,
Corvallis. 280 pp.; Brodie 1968. Copeia 1968:307–313; Mobley and
Stidham 2000. Wilson Bull. 112:563–564). Indeed, the only existing
account of duck predation on T. granulosa relates an observation of
a female A. platyrhynchos found dead with an adult male T. granu-
losa in its crop (Storm, op. cit.). Existing data suggest that, were the
duck affected by the newt’s poison, 15 min. is a sufficient interval
in which to exhibit symptoms, particularly given the rapid meta-
bolic rate of ducks and the relatively short time required to induce
symptoms in much larger animals (Brodie, op. cit.; Bradley and
Klika 1981. J. Am. Med. Assoc. 246:247; Rombough 2008. Herpetol.
Rev. 39:336). Although regional variation in newt toxicity has been
documented (Brodie and Brodie 1990. Evolution 44:651–659; Brodie
and Brodie 1991. Evolution 45:221–224), the reported susceptibil-
ity of birds suggests that the A. platyrhynchos would be expected to
show some symptoms of poisoning, even at relatively low levels of
toxicity. Why this one did not is puzzling. It is also puzzling that the
newt’s noxious skin secretions did not repel the duck. It may be that
dipping the T. granulosa in the water was the ducks attempt to rinse
off these skin secretions, though this is speculation on my part.
This observation adds to a growing body of literature that
suggests amphibians might be seasonally important food sourc-
es for ducks, particularly during periods of high protein need
(Wells 2007. Ecology and Behavior of Amphibians. University of
Chicago Press, Chicago, Illinois. 1400 pp.; Rombough and Brad-
ley 2010. Herpetol. Rev. 41:203), an aspect of the birds’ life history
that might have been overlooked because of the timing and loca-
tion of many previous studies of A. platyrhynchos diet.
I thank Michael O’Loughlin for reviewing this note.
CHRIS ROMBOUGH, Rombough Biological, P.O. Box 365, Aurora,
Oregon 97002, USA; e-mail:
ADENOMERA HYLAEDACTYLA (Napo Tropical Bullfrog).
PREDATION. There are many published accounts of Giant
Fishing Spiders (Ancylometes rufus) and other ctenid spiders
consuming frogs (Prado and Borgo 2003. Herpetol. Rev. 34:238–
239; Toledo 2005. Herpetol. Rev. 36:395–400; Melo-Sampaio
et al. 2012. Herpetol. Rev. 43:636–637). At 2311 h on 5 August
2011 an adult Adenomera hylaedactyla was found being preyed
upon by an Ancylometes rufus in leaf litter (Fig. 1). Five minutes
later, another spider (Ctenus sp.) was observed preying upon a
juvenile A. hylaedactyla (Fig. 2). Both predation events occurred
in a small forest fragment in Rio Branco, Acre, Brazil (9.911880°S,
67.767382°W, WGS 84; 173 m elev.). At the time of the observation,
the spiders were immobilizing the frogs by biting on either side
of the body with their chelicerae and injecting venom into the
lateral portion near the groin. Our observations lasted 30 minutes
using white headlamps. We did not observe any resistance or
fight behavior by the frogs, possibly due to the quick effect of the
spiders’ toxins.
PAULO ROBERTO MELO-SAMPAIO, Departamento de Vertebrados,
Museu Nacional, Quinta da Boa Vista – São Cristóvão – CEP: 20140-040, Rio
de Janeiro, Rio de Janeiro, Brazil (e-mail:; LO-
RENA CORINA BEZERRA DE LIMA, Laboratório de Ecologia e Evolução,
Instituto Butantan, Av. Dr. Vital Brazil, 1500 – Butantã – CEP: 05503-900 São
Paulo, São Paulo, Brazil (e-mail:;
CAMILA MONTEIRO BRAGA DE OLIVEIRA, (e-mail: camila.mbo@gmail.
com); RAELLEN DA SILVA MOURA, União Educacional do Norte, BR 364
Km 02 – Alameda Hungria, 200 – Jardim Europa II – CEP: 69.915-497, Rio
Branco, Acre, Brazil (e-mail:
AMPLEXUS. Amphibian reproduction is linked to the commu-
nication strategies and the acoustic, visual, and chemical signals
produced by breeding pairs (Bowcock et al. 2008. Anim. Behav.
75:1571–1579; Wells 2010. The Ecology and Behavior of Amphib-
ians. University of Chicago Press, Chicago, Illinois. 1400 pp.).
However, signals do not always guarantee intraspecific mating
(Wells 2010, op. cit.). Here, we report the first observation of in-
terspecific amplexus between Atelopus carrikeri and A. laetis-
simus, two amphibian species endemic to the Sierra Nevada de
Santa Marta, Colombia (SNSM). The SNSM is an isolated moun-
tain range located on the northeastern Caribbean coast of Co-
lombia. The SNSM National Park boasts extremely high levels of
biodiversity and endemism and was recently ranked the world’s
most irreplaceable protected site (Le Saout et al. 2013. Science
Fig. 1. Ancylometes rufus preying on an adult Adenomera hylaedactyla.
Fig. 2. Ctenus sp. preying on a juvenile Adenomera hylaedactyla.
Herpetological Review 48(3), 2017
Atelopus carrikeri inhabits paramo ecosystems at high eleva-
tions between 2900 and 4800 m elev. (Rueda-Solano 2012. Her-
petotropicos 8:61–66; Rueda-Solano et al. 2016. J. Therm. Biol.
58:91–98), whereas A. laetissimus inhabits cloud forests and is
usually found at lower elevations of 1500–2800 m elev. (Ruiz-Car-
ranza et al. 1994. Revista Acad. Colomb. Ci. Exact. 19:153–163).
The species are listed as Critically Endangered and Endan-
gered, respectively (
UK.2014-3.RLTS.T54519A3015811.en; 27 April 2017).
We recorded interspecific amplexus between a male A. car-
rikeri and female A. laetissimus (Fig. 1) at 2130 h on 20 June 2016,
in cloud forest at a location locally known as the Pascual stream,
in San Pedro de la Sierra, in the Cebolletas mountain range on
the western slope of the SNSM, Cienaga, Colombia (10.91944°N,
73.93056°W, WGS 84; 2000 m elev.). During our two-day field
expedition, we recorded a reproductive event of A. laetissimus,
with more than 20 observations of intraspecific A. laetissimus
amplexus (Fig. 2). We easily recognized the male A. carrikeri due
to its smooth skin with bright coloring and big size with sturdy
limbs, and darkened iris; which is very different than male A. la-
We are uncertain how this amplexus occurred since they are
usually allopatric, utilizing different ecosystems in the SNSM.
However, a plausible hypothesis is that the Pascual stream begins
in the paramo and crosses a large part of the Cebolletas moun-
tain range until it reaches the Sevilla river. This path creates zones
of contact between the paramo and forested ecosystems, which
would allow for A. carrikeri individuals to disperse into the distri-
butional limits of A. laetissimus. This potential corridor may allow
for the occurrences of interspecific amplexus between these two
Atelopus species. The result of this amplexus may be the hybrid-
ization or introgression of genes between the two species, similar
to reported hybridizations between Rhinella atacamensis and
R. arunco (Correa et al. 2012. J. Herpetol. 46:568–577). However,
we are not certain hybridization will happen. Our new finding of
the amplexus between interspecific Bufonids accompanies other
similar records within the family (Haddad et al. 1990. Rev. Bras.
Biol. 50:739–744; Machado and Bernarde 2011. Herpetol. Notes
4:167–169; Correa et al. 2012, op. cit.; Flores-Hernández and Mar-
tínez-Coronel 2014. Acta Zool. Mex. 30:395–398; Sodré et al. 2014.
Herpetol. Notes. 7:287–288; Costa-Campos et al. 2016. Acta Zool.
Mex. 32:385–386). Nonetheless, this finding could have important
conservation implications for Atelopus populations in the SNSM.
Fig. 2. Multiple intraspecific amplexus between conspecific Atelopus laetissimus encountered in the Pascual stream, in San Pedro de la Sierra,
during 20–21 June 2016, in the Sierra Nevada de Santa Marta, Colombia.
Fig. 1. Interspecific amplexus between a male Atelopus carrikeri (top)
and a female Atelopus laetissimus (bottom) from Pascual stream,
2200 m cloud forest, during the reproductive season in June 2016, in
the Sierra Nevada de Santa Marta, Colombia.
Herpetological Review 48(3), 2017
JOSÉ LUIS PÉREZ GONZÁLEZ, Grupo de Investigación en Ecología
Aplicada, Facultad de Ciencias Básicas, University del Magdalena, Santa
Marta, Colombia (e-mail:; NICO-
LETTE ROACH, Department of Wildlife, Fisheries and Ecological Sciences,
Texas A&M University, College Station, Texas 77843, USA; Global Wildlife
Conservation, PO Box 129, Austin, Texas 78767, USA (e-mail: nroach@; LUIS ALBERTO RUEDA SOLANO, Grupo de Investigación en
Ecología Aplicada, Facultad de Ciencias Básicas, University of Magdalena,
Santa Marta, Colombia; Grupo Biomics, Facultad de Ciencias, University de
los Andes, Bogotá, Colombia (e-mail:
like the iconic Central American Atelopus species, little is known
about the behavior of Amazonian Atelopus species. During field
work in Amazonia I was able to observe and film, to my knowl-
edge, previously undescribed behavior of A. spumarius sensu
lato (Lötters et al. 2002. Salamandra 38:95–104) in three different
individuals of two different populations, one on each side of the
Amapari River near the village of Serra do Navio, Amapá, Brazil
(A: at 1120 h on 6 April 2016 at 0.9426000°N, 51.9437667°W, 108 m
elev.; B: at 1340 h on 7 April 2016 at 0.88876°N, 52.02463°W, 91 m
elev.; C: at 1120 h on 9 April 2016 at 0.88916°N, 52.02416°W, 91 m
elev., WGS84). The animals were completely undisturbed when
filmed in their natural habitat from a minimum distance of ca. 4
m. The observed individuals were all adult males (A. SVL = 3.01
cm, 1.47 g; B. SVL = 2.77 cm, 1.57 g; C. SVL = 2.85 cm, 1.41 g) that
were calling at the time of encounter or started calling during the
encounter. All three individuals were roaming on a branch or a
tree trunk (Fig. 1) and eventually, when sitting, shortly rubbed
their venters on the wood in a shaky or swaying movement. The
behavior was initiated by slightly lifting the posterior, followed
by moving the venter to the left and right in a swaying manner
(video footage of this behavior can be viewed at: https://tinyurl.
com/hq7j4zs). In terms of duration it took ca. 4–5 s from begin-
ning to termination of this distinct behavior. In all incidents
it was displayed in close proximity to the used calling spot on
the same branch or tree trunk. The behavior was observed be-
fore as well as after calling activities. Further, in the footage of
the third observation one can clearly recognize liquid excretion
during the behavior (for video footage see:
y9ftpgv2). Since the same individual was filmed urinating some
minutes prior to the excretion event (also visible in the footage),
I have reason to assume a function other than bladder emptying,
especially as the origin of the liquid cannot be stated with cer-
tainty. Further observations and examination of the secretion as
well as the nature and texture of the skin in the involved area will
help to understand whether the observed behavior could be a
manner of territorial marking.
DANIELA C. RÖSSLER, Trier University, Department of Biogeography,
Am Universitätsring 15, 54296 Trier, Germany; e-mail: roesslerdaniela@aol.
los is a monotypic genus containing only B. ternetzi (Craugasto-
ridae) and is endemic to the Cerrado biome (Valdujo et al. 2012.
S. Am. J. Herpetol. 7:63–78). This species is commonly found in
riverine forest litter and permanent streams with a rocky bed in
the Cerrado and gallery forests (Bastos et al. 2003. Anfíbios da Flo-
resta Nacional de Silvânia, Estado de Goiás. Stylo Gráfica e Editora.
Goiânia, Goiás. 29 pp.; Araújo et al. 2007. Check List 3:153–155).
At 1910 h on 24 May 2016, we found an individual Ancylo-
etes concolor feeding on an adult male B. ternetzi (total length
= 1.5 cm; municipality of João Pinheiro, Minas Gerais, Brazil;
17.41121°S, 45.66604°W, WGS84; elev. 691). At the moment of the
observation, the air temp. was 23°C and the water temp. 22°C.
The predator was found with its prey already dead in the stream
margin on stony substrate.
Spiders from the genus Ancylometes are normally found as-
sociated with water bodies and can easily paddle on the water
surface, diving for prey and away from predators (Höfer and
Brescovit 2000. Insect Syst. Evol. 31:323–360). Besides preying
upon invertebrates, they also prey on small vertebrates such as
fish (Gasnier et al. 2009. In Fonseca et al. [eds.], A Fauna de Ar-
trópodes da Reserva Florestal Ducke: Estado Atual do Conheci-
mento Taxonômico e Biológico, pp. 223–230. Instituto Nacional
de Pesquisas da Amazônia–INPA, Manaus) and anurans (Maffei
et al. 2010. Herpetol. Notes 3:167–170; Bocchiglieri et al. 2010.
Herpetol. Rev. 41:325; Moura and Azevedo 2011. Biota Neotr.
11:1–3). To our knowledge, this is the first report of predation on
B. ternetzi by A. concolor. After the observation, the frog and spi-
der were collected and deposited in the Museu de Zoologia João
Moojen, at the Universidade Federal de Viçosa, Viçosa, Minas
Gerais, Brazil (voucher number MZUFV 17150).
We thank Antonio D. Brescovit for helping with the identifica-
tion of the spider species, Katie Lempke for English review, Den-
drus Projetos Ambientais e Florestais for financial support, and
Sean Graham for editorial suggestions.
Fig. 1. Typical position of a male Atelopus spumarius sensu lato on a
branch, in which the described behavior was observed.
Fig. 1. Barycholos ternetzi being preyed upon by an Ancylometes con-
color in the municipality of João Pinheiro, Minas Gerais, Brazil.
Herpetological Review 48(3), 2017
J. M. GUEDES (e-mail:, and RENATO N. FEIO,
Museu de Zoologia João Moojen, Departamento de Biologia Animal, Uni-
versidade Federal de Viçosa, CEP 36570-000, Viçosa, MG, Brazil (e-mail:
CALLIMEDUSA TOMOPTERNA (Tiger-striped Leaf Frog)
and DENDROPSOPHUS MINUTUS (Lesser Treefrog).
INTERSPECIFIC AMPLEXUS. Availability of reproductive sites,
such as calling and oviposition sites, shared by anuran species
might lead to interspecific matings, particularly when males
perform active searches for females, use satellite behavior, or
have limited capacity to discriminate between sexes, all of which
are common behaviors in many Neotropical frogs (Sodré et al.
2014. Herpetol. Notes 7:287–288; Rocha et al. 2015. Herpetol.
Notes 8:213–215; Ceron and Zocche 2016. Herpetol. Rev.
47:120). Herein we report two cases of interspecific amplexus
between individuals from two different families: Hylidae and
Phyllomedusidae. At 2041 h on 18 January 2016, we found many
reproducing anuran species at a temporary pond, in Reserva
Extrativista Arapixi, Boca do Acre, Amazonas, northern Brazil
(8.993683°S, 67.831433°W, WGS 84; 137 m elev.). These species
were: Callimedusa tomopterna, Dendropsophus leucophyllatus,
D. marmoratus, D. minutus, D. sarayacuensis, Phyllomedusa
bicolor, P. camba, Pithecopus palliatus, and Scinax garbei.
We observed two pairs of male Dendropsophus minutus
(Lesser Treefrog, Ranita Amarilla Común) amplexing male Cal-
limedusa tomopterna (Tiger-striped Leaf Frog) on vegetation
around the pond. The pairs remained in amplexus and did
minimal movements while being observed for 30 minutes. The
first pair was found in inguinal amplexus (Fig. 1) while the sec-
ond pair was backwards with the mouth of the male D. minutus
mouth directed just above the cloaca of C. tomopterna (Fig. 2).
In the same pond we observed conspecific amplexus of adult D.
minutus (Fig. 3), and P. camba. No females of C. tomopterna were
found. Although interspecific amplexus has been documented
in Amazonian species such as bufonids (Machado and Bernarde
2011. Herpetol. Notes 4:167–169) and hylids (Aichinger 1987.
Salamandra 23:269–276), amplexus between different families is
rarely observed in nature (Carvalho and Nascimento 2012. Her-
petol. Rev. 43:461).
This study was conducted with SISBIO permit #51748-1.
PAULO ROBERTO MELO-SAMPAIO, Programa de Pós-graduação em
Zoologia, Museu Nacional, Quinta da Boa Vista, Rio de Janeiro, Rio de Janei-
ro, CEP: 20940-040, Brazil (e-mail:; JOSIMAR
COSTA DA SILVA, Instituto Desenvolver Amazonas - Av. Amazonas, 1728.
Bairro Macaxeiral, Boca do Acre, Amazonas, CEP: 69850-000, Brazil.
CRAUGASTOR SABRINUS (Long-legged Stream Frog). DIET.
Craugastor sabrinus (Craugastoridae) ranges from eastern Gua-
temala to the Maya Mountains in southern Belize (Crawford and
Smith 2005. Mol. Phylogenet. Evol. 35:546–555). It is found in wet
and moist tropical forests along rivers and streams. Listed as near
threatened in the IUCN Red List, its population decline is mainly
due to habitat degradation and loss, especially in Guatemala.
Species from the genus Craugastor are opportunistic feeders and
have been documented feeding on arthropods (Lieberman 1986.
Acta Zool. Mex. 15:1–72). Despite being considered common in
Belize, little is known about the natural history and diet of C. sab-
rinus. Here we report a vertebrate prey item for this species.
At 0900 h on 7 Mar 2016, we observed a Scincella cherriei
(Brown Forest skink; SVL ca. 60 mm) crossing the path on the
Gibnut Trail in the Cockscomb Basin Wildlife Sanctuary in south-
ern Belize (16.78540°N 88.45860°W; WGS 84). The S. cherriei was
disturbed by our movement and ran through leaf litter away from
Fig. 1. Interspecific amplectant pair of males of Dendropsophus
minutus and Callimedusa tomopterna.
Fig. 2. Another interspecific amplectant pair of males of Dendropso-
phus minutus and Callimedusa tomopterna with unusual position.
Fig. 3. Syntopic conspecific amplectant pair of Dendropsophus
Herpetological Review 48(3), 2017
the path. We then observed the S. cherriei encounter a C. sabrinus
(SVL ca. 80 mm), which promptly proceeded to feed on the skink,
with only the tail of the skink remaining outside the mouth of the
frog when first observed by us (Fig. 1). We observed the S. cher-
riei moving within the body cavity of the C. sabrinus for approxi-
mately five minutes. Although we did not stay to observe the frog
completely ingest the skink, we consider S. cherriei to be a prey
item of the C. sabrinus.
MARY-RUTH LOW, Wildlife Reserves Singapore, 80 Mandai Lake Rd,
Singapore 729628 (e-mail:; BRADLEY NISSEN,
2305 Shelby Dr, Charlottesville, Virginia 22901, USA; GLISELLE MARIN,
5619 Moguel & Lizarraga Ave, Belize City, Belize, Central America; JORDI
JANSSEN, Dijkstraat 1, 6701 CH Wageningen, Netherlands.
PREDATION. Eleutherodactylus planirostris is a small direct-
developing frog native to Cuba, Isla de Juventud, the Cayman Is-
lands, and the Caicos Islands (Frost 2016. Amphibian Species of
the World: an Online Reference. Version 6.0, http://research.amnh.
org/herpetology/amphibia/index.html) [Accessed 2017.03.20].
American Museum of Natural History, New York) and introduced
to the southeastern USA, Hawaii, Hong Kong, Guam, Philippines,
Jamaica, Honduras, Surinam, Isla de Guanaja (Honduras), and Ve-
racruz, Mexico (Frost, op. cit.; Olson et al., 2011. Pacific Sci. 66:255–
270). Predators of E. planirostris in its native range include inverte-
brates, frogs, mammals, birds, lizards, and snakes (Henderson and
Powell 1999. In Crother [ed.] Caribbean Amphibians and Reptiles,
pp. 223–268. Academic Press, San Diego, California), including
three species of Cubophis (Henderson and Powell 2009. Natural
History of West Indian Reptiles and Amphibians. University Press
of Florida, Gainesville. 495 pp.). In the introduced range, preda-
tion is less well-documented. Diadophis punctatus (Ring-necked
Snake) is the only predator native to the USA known to consume
E. planirostris (Meshaka 2011. Herpetol. Conserv. Biol. 6:1–101).
Although Ernst and Ernst (2003. Snakes of the United States and
Canada. Smithsonian Institution Press, Washington D.C. 661
pp.) list E. planirostris as a prey item of Rhadinaea flavilata (Pine
Woods Snake), they provide no documentation and none of the
references cited therein contain additional information. Here we
document predation on E. planirostris by R. flavilata.
At 1856 h on 11 October 2016, we captured a juvenile R. fla-
vilata (HM 163103) that had eaten an adult E. planirostris (HM
163012) at a private residence in Alachua County, Florida, USA
(29.658926°N, 82.379202ºW; WGS84). Rhadinaea flavilata eat
primarily amphibians, especially hylid frogs (Ernst and Ernst,
op. cit.). Other members of the genus Rhadinaea feed on eleu-
therodactylid frogs throughout Central and South America
(Myers 1974. Bull. Am. Mus. Nat. Hist. 153:1–268), as well as on
lizards, salamanders, amphibian eggs, and other anurans, in-
cluding some highly toxic dendrobatids (Lenger et al. 2014. Her-
petol. Notes 7:83–84). Other predators of E. planirostris in parts
of its introduced range include Osteopilus septentrionalis (Cuban
Treefrog; Meshaka, op. cit.), which is also a predator in the na-
tive range (Henderson and Powell, op. cit.), and Boiga irregularis
(Brown Treesnake) on Guam (Mathies et al. 2012. Herpetol. Rev.
ANDREW M. DURSO, Department of Biology, Utah State University,
Logan, Utah 84322, USA (e-mail:; LUKE SMITH,
3716 NW 7th Place, Gainesville, Florida 32607, USA (e-mail: smithsqrd@
GASTROPHRYNE OLIVACEA (Western Narrow-mouthed Toad).
PREDATION. On 12 November 2016, 11.5 km SE of Valentine, Jeff
Davis County, Texas, USA (30.5217°N, 104.4020°W; WGS 84), we
found a dead Gastrophryne olivacea impaled on a barbed wire
fence (Fig. 1) at a known Loggerhead Shrike (Lanius ludovicia-
nus) larder. Documented predators of G. olivacea include snakes
(Agkistrodon contortrix, Thamnophis spp.) and other frogs (Litho-
bates spp.). Gastrophryne olivacea possess toxic and slippery skin
secretions that may afford them some protection from predation
(Dodd 2013. Frogs of the United States and Canada. Volume 1. The
John Hopkins University Press, Baltimore, Maryland. 460 pp.). To
our knowledge, this is the first record of shrike predation upon G.
olivacea (Clark 2011. Son. Herpetol. 24:20–22; Dodd 2013, op. cit.).
The specimen is deposited in the James F. Scudday vertebrate col-
lection at Sul Ross State University (SRSU 6928).
Fig. 1. Gastrophryne olivacea fatally impaled by a Loggerhead Shrike.
Fig. 1. Craugastor sabrinus ingesting a Scincella cherriei.
Herpetological Review 48(3), 2017
GAN SEILER (e-mail:, and SEAN P. GRAHAM,
Department of Biology, Geology, and Physical Sciences, Sul Ross State Uni-
versity, Alpine, Texas, 79832, USA (e-mail:
DATION. Here we report opportunistic predation of Hypsiboas
marginatus by Attila rufus (Gray-hooded Attila; Tyrannidae), a
bird endemic to the Atlantic Forest of eastern Brazil (Sick 1997.
Ornitologia Brasileira. Editora Nova Fronteira. Rio de Janeiro.
912 pp.). At 1030 h on 30 September 2008 in Bairro Guaraú,
municipality of Peruíbe, state of São Paulo, Brazil (24.358850°S,
47.018913°W; WGS 84), an adult A. rufus was observed captur-
ing a frog in a pond on a dirt road. After that, the bird flew and
perched in a tree, where it was photographed with a H. albomar-
ginatus in its bill (Fig. 1A). The bird beat the frog against a branch
several times. The frog gave a distress call and inflated its body.
After subjugating the frog, the bird swallowed it posterior-first.
At 1145 h on 8 September 2012, at Opashaus Lodge, mu-
nicipality of Domingo Martins, state of Espírito Santo, Brazil
(20.348391°S, 40.675957°W, WGS 84), an adult A. rufus was ob-
served swallowing an individual of H. albomarginatus (Fig. 1B).
It took 40 min. to the complete ingestion (also posterior-first).
This is the first record of predation of H. albomarginatus by A.
rufus. Although there are no Neotropical birds that are entirely
frog specialists, frogs seem to be frequent opportunistic prey for
avian taxa in this region (Toledo et al. 2007. J. Zool. 271:170–177).
We thank Almir Almeida and Mario Candeias for sharing their
observations and photographs
JOÃO PAULO GAVA JUST, Programa de Pós-Graduação em Biologia
Animal (PPG-BA), Departamento de Ecologia, Zoologia e Genética, Univer-
sidade Federal de Pelotas (UFPel), Capão do Leão, Rio Grande do Sul, Brazil
(e-mail:; JAIRO JOSÉ ZOCCHE, Programa de
Pós-Graduação em Ciências Ambientais (PPG-CA), Laboratório de Ecologia
de Paisagem e de Vertebrados, Universidade do Extremo Sul Catarinense
(UNESC), Ave. Universitária, 1105, Criciúma, Santa Catarina, Brazil (e-mail:
about the natural history of Leptodactylus caatingae, a species
that inhabits the Caatinga of northeastern Brazil. This species is
recognized as an explosive breeder that reproduces at temporary
ponds (Heyer and Juncá 2003. Proc. Biol. Soc. Washington
116:317–329; Magalhães et al. 2013. S. Am. J. Herpetol. 8:203–210)
and can be found within highly human-disturbed areas (Vieira
et al. 2012. Bol. Mus. Para Emílio Goeldi. Cienc. Nat. 7:153–156).
Here we present the first report on its predators.
At 2000 h on 24 February 2016, we found a juvenile Lepto-
dactylus fuscus (SVL = 31.2 mm; head width = 11 mm) feeding
on a juvenile of L. caatingae (SVL = 24.2 mm; head width = 8.2
mm), near a small temporary pond formed after heavy rains, at
vacant land inside urban perimeter in the municipality of Jequié,
Bahia, Brazil (13.840533°S, 40.072654°W, WGS 84; elev. 194 m). At
the moment we found them, the juvenile L. fuscus had already
captured the juvenile L. caatingae and had swallowed its left
leg almost entirely (Fig. 1). The prey struggled for about 20 min.
to disentangle from the mouth of the predator. When it finally
managed to liberate itself, its left foot was already completely di-
gested. It is possible that our presence with lamps and camera
disturbed the predator and hence helped the juvenile L. caatin-
gae to get free. Or it is possible that the prey was too big to be
swallowed completely (the prey made up 77.6% of the predators
length). We collected both frogs and deposited them in the her-
petological collection of Universidade Estadual do Sudoeste da
Bahia (MHNJCH 1079 and MHNJCH 1080 for L. caatingae and L.
fuscus, respectively).
Although it is known that L. fuscus is an opportunistic preda-
tor (Sugai et al. 2012. Biota Neotrop. 12:99–104) and that anuro-
phagy is common in the family Leptodactylidae (Measey et al.
2015. PeerJ 3:e1204, DOI 10.7717/peerj.1204), this is the first re-
port of L. fuscus preying on other frogs.
We thank Nathana Pereira for helping with fieldwork. The
frogs were collected under ICMBio/SISBIO permit #35068.
GABRIEL NOVAES-E-FAGUNDES, Programa de pós-graduação em
Zoologia, Universidade Estadual de Santa Cruz, Rodovia Jorge Amado, km
16, CEP 45662-900, Ilhéus, Bahia, Brazil (e-mail: gnovaesefagundes@gmail.
com); JULIANA ZINA, Departamento de Ciências Biológicas, Universidade
Estadual do Sudoeste da Bahia, Avenida José Moreira Sobrinho, Jequiez-
inho, CEP 45208-091, Jequié, Bahia, Brazil.
FENSIVE BEHAVIOR. Leptodactylus insularum is a common
terrestrial inhabitant of lower elevations of southern Central
and northern South America. Until recently the northern popu-
lations of the species were regarded as a synonym of L. bolivi-
anus (Savage 2002. The Amphibians and Reptiles of Costa Rica: A
Fig. 1. Hypsiboas albomarginatus being preyed upon by Gray-hood-
ed Attila (Attila rufus) in southeast Brazil. A) Predation event from
Bairro Guaraú, state of São Paulo. B) Predation event from Opashaus
Lodge, state of Espírito Santo.
Fig. 1. Leptodactylus fuscus ingesting left leg of a juvenile of Lepto-
dactylus caatingae.
Herpetological Review 48(3), 2017
Herpetofauna between Two Continents, between Two Seas. Uni-
versity of Chicago Press, Chicago, Illinois. 934 pp.; Heyer and de
Sá 2011. Contrib. Zool. 635:1–58).
On 12 March 2015, I caught an adult specimen of L. insularum
ca. 300 m NW of La Gamba Tropical Research Station, approx. 10
km NW Golfito, Provincia de Puntarenas, Costa Rica (8.70312°S,
83.20456°W, WGS 84; elev. 120 m). The individual was placed in
a bag overnight and handled for a photo session the next morn-
ing. During manipulation the frog inflated its body and raised its
legs and body well above the ground, displaying the aposematic
coloration of the posterior thighs (Fig. 1). This behavior lasted for
a short while (< 20 sec) and could be repeated several times by
tapping the frog gently on its head.
Defensive behaviors of frogs were recently reviewed by Toledo
et al. (2011. Ethol. Ecol. Evol. 23:1–25). Accordingly, a behavior as
described above is classified as full body-raising with legs vertically
stretched, showing aposematic colors. Within the genus, full
body-raising has been described from L. labyrinthicus, L. laticeps,
L. latrans, and L. mystacinus, but apparently not from members
of the L. bolivianus group (Toledo et al., op. cit.). According to
Toledo et al. (op. cit.), the behavior is common in species with
noxious skin secretions. Toxic skin secretions are known from a
number of species of the genus (e.g., Ryan et al. 2010. Herpetol.
Rev. 41:337–338; Toledo et al. 2011, op. cit.; Haddad et al. 2013.
Guia dos Anfíbios da Mata Atlântica: Diversidade e Biologia.
Anolisbooks, São Paulo. 544 pp.), but to my knowledge not yet
from L. insularum, but they are likely to occur.
Many thanks to Jörg Scheidung and Jeff Schreiner for their
help in the field and to Mark Scherz for his help in the office.
MICHAEL FRANZEN, Zoologische Staatssammlung München (ZSM-
SNSB), Münchhausenstrasse 21, 81247 München, Germany; e-mail:
PREDATION. Limnonectes palavanensis is a nocturnal leaf-
litter frog found in the western and northern part of Borneo and
Palawan Island of the Philippines (Das 2007. A Pocket Guide:
Amphibians and Reptiles of Brunei. Natural History Publications
[Borneo], Kota Kinabalu, Sabah, Malaysia. 208 pp.). This small
frog (snout–urostyle length: males = 21–30 cm, females = 28–33
cm) inhabits the forest floor of primary and secondary mixed-
dipterocarp forest, where males sporadically call to attract
females. Females lay eggs in the leaf litter and the eggs are
guarded by the males until hatching. The male then transports
the newly hatched tadpoles to small bodies of water (Inger and
Voris 1988. Copeia 1988:1060–1061; Goyes Vallejos 2016. Doctoral
Dissertation. University of Connecticut, Storrs, Connecticut).
Instances of predation have never been reported for Limnonectes
palavanensis. Here, we report a predation event by the Rough-
backed Snake (Xenodermus javanicus) of an adult female of L.
palavanensis. Seemingly a frog specialist, X. javanicus is active at
night and it has been observed foraging under dead leaves on the
forest floor, prime L. palavanensis habitat (Stuebing et al. 2014. A
Field Guide to the Snakes of Borneo. Natural History Publications
[Borneo], 2nd edition, Kota Kinabalu, Sabah, Malaysia. 310 pp.).
This observation took place at the Kuala Belalong Field Studies
Centre (KBFSC), located within the Ulu Temburong National
Park (Brunei Darussalam; 4.546°N, 115.157°E; WGS 84). On the
night of 16 October 2014, we observed an adult X. javanicus with
its stomach engorged, close to an artificial pool sporadically used
by males of L. palavanensis to deposit tadpoles. The individual
was taken to the KBFSC Laboratory to be photographed and
released the following night (Fig. 1A). At 0900 h on 17 October
2014, a regurgitated individual of L. palavanensis was found in
the terrarium where the individual X. javanicus was held. Upon
examination, we found that it was an adult female (Fig. 1B).
This observation constitutes the first record of predation of L.
Fig. 1. Leptodactylus insularum (San Miguel Island Frog) exhibiting
defensive behavior.
Fig. 1. A) Rough-backed Snake (Xenodermus javanicus) after
regurgitation occurred. B) Regurgitated individual of Limnonectes
Herpetological Review 48(3), 2017
JOHANA GOYES VALLEJOS, Department of Ecology and
Evolutionary Biology, University of Connecticut, Storrs, Connecticut, USA
(e-mail:; HANYROL H. AHMAD SAH, Faculty
of Science, Universiti Brunei Darussalam, Tungku Link, Gadong BE 1410,
Brunei Darussalam.
and AMBYSTOMA MACULATUM (Spotted Salamander).
INTERSPECIFIC AMPLEXUS. Interspecific amplexus has been
described in a variety of both native and invasive anurans,
especially in those species that have explosive reproductive
events (Wells 2007. The Ecology and Behavior of Amphibians.
University of Chicago Press, Chicago, Illinois. 1400 pp.). When
multiple anuran species are breeding in aggregations there may
be misidentification during the mating process. The literature
reports males attempting to mate with inanimate objects, other
males, dead conspecifics, and individuals of other species
(Bedoya et al. 2014. Herpetol. Notes. 7:515–516; Rocha et al.
2015. Herpetol. Notes. 8:213–215). The literature usually reports
incidences of interspecies amplexus between organisms of the
same order (generally between anurans). However, multiple
instances of interspecific amplexus between anurans and
caudates have been described. Here I report the interspecific
amplexus between an anuran, Lithobates sphenocephalus, and a
caudate, Ambystoma maculatum.
During the spring amphibian migrations in Harford County,
Maryland, USA (39.2555°N, 76.15480°W, WGS84; elev. 6.5 m),
multiple species of amphibians breed simultaneously in the
same ephemeral wetlands. At 2144 h on 25 March 2015, an adult
L. sphenocephalus was found in amplexus with an adult A. macu-
latum (Fig. 1) in an ephemeral roadside ditch less than 1 m from
the edge of a road. The area is part of a large wetland complex
with breeding Anaxyrus americanus, L. sylvaticus, Pseudacris
crucifer, Acris crepitans, and L. palustris all present within the
same wetland complex that night. After photos were taken, the
animals were released back into the roadside ditch. The A. macu-
latum immediately swam away and started struggling violently
to remove the L. sphenocephalus, but was unable to accomplish
this in the 20 min. they were observed. There has been no other
observed interspecific amplexus at the site within the last two
I thank Richard Seigel for his review and comments on this
HUNTER J. HOWELL, Department of Biological Sciences, Towson
University, 8000 York Road, Towson, Maryland 21252-0001, USA; e-mail:
AL NIGHT SHELTER. The Guayana highlands of South America,
also known as Pantepui, constitute a singular biogeographical
province recognized by the uniqueness of its biota, biodiver-
sity and endemism. This region is characterized by sandstone
tabletop mountains (tepuis) that often emerge as high-elevation
“islands” (1500–3000 m elev.) in the overall landscape (Huber
1995. In Steyermark et al. [eds.], Flora of the Venezuelan Guay-
ana, pp. 1–61. Timber Press, Portland, Oregon). Bufonids of the
genus Oreophrynella are endemic to the summits and slopes of
the eastern tepuis in southern Venezuela and adjacent Guyana
and Brazil (ca. 1067–2800 m elev.). This genus currently contains
nine species. They are clearly distinguished from other bufonids
by their small size, thick skin between digits, and opposable toes
(Kok 2009. Zootaxa 2071:35–49). Most of the species are diurnal
rock dwellers with terrestrial habits (McDiarmid and Gorzula
1989. Copeia 1989:445–451; Señaris et al. 1994. Publ. Asoc. Ami-
gos de Doñana 3:1–37; Señaris 1995. Mem. Soc. Cienc. Nat. La
Salle 140:177–182; Señaris et al. 2005. Pap. Avul. Zool. 45:61–67).
Oreophrynella highland species are found active mainly on bare
sandstone in open areas on the tepuis or resting beneath rocks,
but occasionally they can be observed sitting < 1 m above ground
on leaves of Stegolepis guianensis, Lomaria sp., and Brocchinia
hechtioides (McDiarmid and Gorzula 1989, op. cit.). Although
McDiarmid and Gorzula (1989, op. cit.) have studied at least four
species of Oreophrynella (including O. quelchii) in four tepuis,
it is not clear what species they reported climbing bushes, and
importantly, why these rock dwellers may engage in such climb-
ing. The two upland species O. dendronastes and O. macconnelli
have arboreal habits and are primarily found associated to veg-
etation (Lathrop and MacCulloch 2007. Herpetologica 63:87–93;
Kok 2009, op. cit.).
Oreophrynella quelchii is a small toad (SVL = 18–23 mm),
endemic to the summit of Roraima and Wei-Assipu tepuis, near
the borders of Venezuela, Guyana and Brazil, between 1700–
2800 m elev. (MacCulloch et al. 2007. Herpetol. Rev. 38:24–30).
Oreophrynella quelchii is diurnal and terrestrial, and is usu-
ally found on bare sandstone or open bare rocky surfaces, but
also in peat patches along or near small streams or temporary
puddles. This species is listed as Vulnerable (Hoogmoed and
Señaris 2004.
Fig. 1. Interspecific amplexus by Lithobates sphenocephalus on
Ambystoma maculatum.
Fig. 1. Oreophrynella quelchii resting in a tangle of lichen attached
to a Bonnetia roraimae bush at the summit of Mount Roraima, Ven-
Herpetological Review 48(3), 2017
T54853A11216183.en; 28 Mar 2017) because it is only known
from two localities. Although this species is common on the
summit of the Roraima tepui and its population is apparently
stable, their behavior and ecology remained virtually unknown.
Here, we report on an opportunistic observation on the use
of a bush as a night shelter by O. quelchii. This record was made
at the summit of Roraima tepui (5.161255°N, 60.781025°W, WGS
84; 2277 m elev.) on 19 November 2015 at 1936 h. We observed an
adult O. quelchii lodged within a small tangle of lichen attached
to a small Bonnetia roraimae bush, at approximately 50 cm above
the ground (Fig. 1). The toad remained immobile during the 20
min we stayed observing it.
Our observation confirms the ability of O. quelchii to climb
in vegetation. While the species inhabiting the sandstone sum-
mits of the tepuis are primarily rock dwellers (McDiarmid and
Gorzula 1989, op. cit.), we believe that the capacity of O. quelchii
to climb bushes at night could be a way to reduce predation by
terrestrial spiders (Theraphosidae). Theraphosid spiders are the
only known potential predator for O. quelchii so far (McDiarmid
and Gorzula 1989, op. cit.). This predator is nocturnal and may
easily predate O. quelchii in the terrestrial environment, both on
top of or beneath the sandstone rocks where this species is gen-
erally found. This is the first record of diurnal O. quelchii climb-
ing bushes to refuge at night probably to protect themselves
from predation by nocturnal, terrestrial theraphosid spiders at
the ground level in open bare rocky areas.
We are grateful to Crystal Kelehear for the constructive criti-
cisms that have improved this manuscript.
NANDEZ, Programa de Pós-graduação em Biodiversidade e Conservação,
Laboratório de Zoologia, Faculdade de Ciências Biológicas, Universidade
Federal do Pará, Rua José Porfírio, 2515, Esplanada do Xingu, 68.372-040,
Altamira, Pará, Brazil (e-mail:; J. CELSA SEÑARIS, Insti-
tuto Venezolano de Investigaciones Cientícas, Centro de Ecología, Labora-
torio de Ecología y Genética de Poblaciones, Altos de Pipe, Apartado Postal
21827, Caracas 1020-A, Venezuela (e-mail:
PHYLLOMEDUSA VENUSTA (Lovely Leaf Frog). DIET. Phyllome-
dusa venusta is an arboreal frog found in northern Colombia, the
valley of Magdalena, the Darién on both sides of the Colombia-
Panamá border, and western Venezuela. The species is common,
but populations are decreasing due to deforestation by agricul-
tural and livestock activities, illegal plantations, human estab-
lishments, and use of agrochemicals (Rodríguez-Mahecha et al.
2008. Guía Ilustrada de Fauna del Santuario de Vida Silvestre Los
Besotes,Valledupar, Cesar, Colombia. Editorial Panamericana,
Formas e Impresos, Bogotá, Colombia. 574 pp.). The food habits
and many other aspects of its biology and ecology are unknown.
Herein we describe the diet of P. venusta in the dry tropical
forest of Colombia at three sites: 1) The Natural Reserve of Civil
Society Campoalegre, Municipality Los Cordobas, Department
of Cordoba (8.48502°N, 76.19520°W, WGS84; elev. 120 m); 2)
Finca Los Mameyales, Municipality Piojó, Department of Atlán-
tico (10.74480°N, 75.09279°W, WGS84; elev. 206 m); 3) Las Deli-
cias farm, Municipality Aracataca, Department of Magdalena
(10.58694°N, 74.14224°W, WGS84; elev. 197 m).
We examined 28 stomachs of P. venusta collected during
0800–1200 h and 1600–1800 h within forests and disturbed ar-
eas. Samples were obtained during 2007 in the dry season (Janu-
ary–March), first rains (April–June), and heavy rains (Septem-
ber–December). SUL (mm), and maximum mouth width (mm)
were recorded for each individual. We identified prey to lowest
taxonomic level possible (family and genus), and their length
and width were measured (complete prey only) using a digital
caliper (nearest 0.1 mm). The individual volume of each prey
item and the number of prey items per stomach for each prey
category were recorded. Volume of each prey item was estimated
using the formula of a prolate spheroid.
Of the captured frogs, six were females and 22 were males
(mean SUL = 67.60 ± 8.76 mm; mean mouth width = 21.76 ± 2.10
mm). The diet consisted of 16 types of prey and was dominated
in volume and frequency by orthopterans. Acarina showed the
highest numerical contribution (Table 1).
It has been suggested that acariphagia occurs in small an-
urans in terrestrial habits. However, P. venusta is large and arbo-
real, suggesting that acariphagia is a trophic phenomenon not
limited to the species defined by Simon and Toft (1991. Oikos
61:263–278). The large numbers of orthopterans and blattarians
consumed are congruent with that reported for other Phyllom-
edusa spp. (Parmelee 1999. Sci. Pap. Nat. His. Mus. Univ. Kansas
11:1–59; Vaz-Silva et al. 2004. Herpetol. Rev. 35:160; Freitas et al.
2008. Biota Neotrop. 8:101–110). Considering the type and prey
proportion, P. venusta appears to be a generalist predator with a
sit-and-wait foraging strategy.
We are grateful to Colciencias, Universidad Nacional de Co-
lombia, Universidad del Atlántico, people at our field sites, Tropi-
cal Organism Biology Group of the Biology Department of the
Universidad Nacional de Colombia, J. O. Combita, M. C. Franco,
and N. Vanegas.
ARGELINA BLANCO-TORRES, Tropical Organism Biology Group,
Biology Department, Universidad Nacional de Colombia, Carrera 45 No.
26-85, Building 421, Laboratory 224, Bogota, Colombia, 111321 (e-mail: ar-; MARTA DURÉ, Centro de Ecología Aplicada del Lito-
table 1. Composition of arachnid prey in the diet of Phyllomedusa
venusta in tropical dry forest of northern Colombia. Volume in mm3.
L = larvae.
Prey Number Volume Frequency of
(%) (%) occurrence
Acarina 5 (31.25) 0.61 (0.02) 0.07
Archegozetes 4 (25) 0.55 (0.02) 0.04
Euzetes 1 (6.25) 0.06 0.04
Opiliones 1 (6.25) 11.76 (3.98) 0.04
Phrynidae 1 (6.25) 111.76 (3.98) 0.04
Orthoptera 4 (25) 1694.67 (60.30) 0.14
Acrididae sp1 1 (6.25) 47.09 (1.68) 0.04
Tettigonidae sp1 1 (6.25) 1255.03 (44.66) 0.04
Tettigonidae sp1 1 (6.25) 342,.00 (12.17) 0.04
Gryllidae sp1 1 (6.25) 50.54 (1.80) 0.04
Blattodea 2 (12.5) 864.60 (30.76) 0.07
Blattellidae sp1 1 (6.25) 370.38 (13.18) 0.04
Blattellidae sp2 1 (6.25) 494.21 (17.59) 0.04
Coleoptera 1 (6.25) 27.52 (0.98) 0.04
L.Coleoptera sp1 1 (6.25) 27.52 (0.98) 0.04
Lepidoptera 1 (6.25) 110.88 (3.95) 0.04
L.Lepidoptera sp1 1 (6.25) 110.88 (3.95) 0.04
Hymenoptera 2 (12.5) 0.32 (0.01) 0.04
Solenopsis 2 (12.5) 0.32 (0.01) 0.04
Herpetological Review 48(3), 2017
ral – Consejo Nacional de Investigaciones Cientícas y Técnicas, Ruta 5, Km
2.5, Corrientes, Argentina, 3400; MARIA ARGENIS BONILLA, Organism Bi-
ology Group, Biology Department, Universidad Nacional de Colombia, Car-
rera 45 No. 26-85, Building 421, Laboratory 224, Bogotá, Colombia, 111321.
PROCERATOPHRYS SCHIRCHI (Sapo-de-chifres; Smooth
Horned Frog). ANTIPREDATOR BEHAVIOR. Proceratophrys
schirchi is a medium (SVL = 39–50 mm) frog endemic to the At-
lantic Forest, occurring in Rio de Janeiro, Espírito Santo, Minas
Gerais and Bahia states in eastern Brazil (Haddad et al. 2013.
Guia de anfíbios da Mata Atlântica: Diversidade e Biologia.
Editora Anolis Books, São Paulo, São Paulo. 544 pp.). Species of
the genus Proceratophrys usually inhabit the leaf-litter after the
metamorphosis (Giaretta et al. 2000. J. Herpetol. 34:173–178;
Kwet and Faivovich 2001. Copeia 2001:203–215). Few studies deal
with aspects of the natural history, ecology, and behavior of Pro-
ceratophrys species.
At 1930 h on 21 May 2016, we captured a male P. schirchi at
Reserva Biológica Augusto Ruschi, Santa Teresa, Espírito Santo,
southeastern Brazil (19.9103°S, 40.5502°W, WGS 84; 650 m elev.).
The frog was sitting on top of the leaves (Fig. 1A), and remained
motionless until we hand-captured it, it displayed the behavior
of thanatosis or death feigning in hand (Fig. 1B). When placed on
the ground the individual remained in this position for ca. one
minute. After returning it to leaf-litter, the frog displayed stiff-
legged behavior (Fig. 1C). This latter behavior was reported for
P. moehringi (Weygoldt 1986. Zool. Jahrb. Syst. 113:429454); for
P. appendiculata (Sazima 1978. Biotropica 10:158); for P. renalis
(Peixoto et al. 2013. Herpetol. Notes 6:479430); for P. boiei and
P. melanopogon (Toledo et al. 2010. J. Nat. Hist. 44:19791988).
Once captured, the individual was transported in a wet plas-
tic bag to the laboratory. On site, it displayed the behavior of puff-
ing-up the body (Fig. 1D). The same behavior was reported for P.
cristiceps (Mângia and Garda 2015. Herpetol. Notes 8:1114). The
similarity in behavior between the congeners may be indicative
of convergence among leaf-litter anurans (Sazima 1978, op. cit.;
Garcia 1999. Herpetol. Rev. 30:224).
We report for the first time a detailed repertoire of antipreda-
tor mechanisms of P. schirchi, contributing to the knowledge on
behavioral ecology of this species.
The specimen is deposited in the Zoological Collection of In-
stituto Nacional da Mata Atlântica (MBML 9677; Museu de Biolo-
gia Mello Leitão), Santa Teresa, Espírito Santo, Brazil.
We thank Instituto Chico Mendes de Conservação da Biodi-
versidade for the field work’s license (n° 49.871-1). ATM thanks
Coordenação de Aperfeiçoamento Pessoal de Nível Superior and
DAK and ECC thank Fundação de Amparo a Pesquisa no Espírito
Santo (FAPES) for scholarships. RBG Clemente-Carvalho is grate-
ful to the Universidade Vila Velha and FAPES, which sponsored
the research of the Laboratório de Ecologia de Anfíbios e Répteis
(#44/2014 and #0611/2015, respectively).
ALEXANDER T. MÔNICO,Universidade Vila Velha, Laboratório de
Ecologia de Anfíbios e Répteis, Vila Velha 29102-770, Espírito Santo, Brazil
Instituto Federal do Espírito Santo, Laboratório de Genética, Santa Teresa
29650-000, Espírito Santo, Brazil; TATIANE DE MELLO DO CARMO, Centro
Universitário do Norte do Espírito Santo, São Mateus, Espírito Santo, Brazil;
CLEMENTE-CARVALHO, Universidade Vila Velha, Laboratório de Ecologia
de Anfíbios e Répteis, Vila Velha, Espírito Santo, Brazil (e-mail: rutebeatriz@
the genus Pseudis are known to occur in Argentina (Vaira et al.
2012. Cuad. Herpetol. 26:131–159). Tadpoles are large, reach-
ing total lengths of 170 mm (Guzmán and Raffo, 2011. Guía de
los anfibios del Parque Nacional y la Reserva Natural El Palmar
Otamendi. Administración de Parques Nacionales, Buenos Aires.
104 pp.). In Argentina, P. platensis is distributed across Buenos
Aires, Chaco, Corrientes, Entre Ríos, Formosa, Santa Fe, Santiago
del Estero and Salta Provinces (Vaira et al. 2012, op. cit.). Pseudis
platensis hosts the nematodes Gyrinicola sp. (Kehr and Hamann
2003. Herpetol. Rev. 34:336–341), Spiroxys sp. (González and
Hamann 2010. Brazil. J. Biol. 71:1089–1092), and Brevimulticae-
cum sp. (González and Hamann 2013. Brazil. J. Biol. 73:451–452)
from Corrientes Province, Argentina and Cosmocerca podicipi-
nus, Rhabdias sp., Brevimulticaecum sp., and Physocephalus sp.
(Campião et al. 2010. Parasitol. Res. 106:747–751; Campião et al.
2016. Comp. Parasitol. 83:92–100) from Corumbá, Mato Grosso
do Sul, Brazil.
In this note we provide a new host record of Gyrinicola cha-
baudi occurring in P. platensis. Six tadpoles of P. platensis (mean
body length = 101.1 mm ± 9.0 SD) were collected from Bañado
de Viñalito (24.406639°S, 63.02925°W, WGS 84; 218 m elev.), Salta
Province, Argentina and deposited in the herpetology collection
of the Universidad Nacional de San Juan, San Juan, Argentina as
UNSJ 3000. The body cavity was opened by a mid-ventral inci-
sion, the digestive tract was removed and its contents examined
for helminths using a dissecting microscope. Fifty-three nema-
todes (21 males, 32 females) were removed and identified as
G. chabaudi. Infection prevalence (number tadpoles infected/
number tadpoles examined x 100) was 100%; mean intensity
(mean number of nematodes per infected tadpole) was 8.83 ±
4.45 SD, range = 5–16. All of the nematodes were deposited in the
Helminthological Collection, Fundación Miguel Lillo as (CH-N-
FML 07710).
Gyrinicola chabaudi was described from specimens recov-
ered from the gut of Leptodactylus ocellatus tadpoles from Santo
Amaro, São Paulo, Brazil (Araujo and Artigas 1982. Mem. Inst.
Butantan 44/45:383–390) and Scinax nasicus from Corrientes,
Argentina (González and Hamann 2005. Facena 21:145–148).
The males were found later from the intestine of tadpoles of S.
Fig. 1. Antipredator postures of Proceratophrys schirchi (MBML9677):
A) natural posture; B) thanatosis; C) stiff-legged behavior; and D)
puffing-up the body.
Herpetological Review 48(3), 2017
ruber and Rhinella crucifer (Souza-Júnior et al. 1991. Rev. Brasil.
Biol. 51:585–588). The specimens of G. chabaudi (males) identi-
fied herein possess the diagnostic characters of this species, es-
pecially three pairs of genital papillae: one preanal pair, another
postanal, laterally projecting and a third ventral pair located in
a short, subulated and coiled tail. In this note, the distribution
of G. chabaudi is expanded and P. platensis is a new host record.
We are grateful to Marissa Fabrezi (Instituto de Biología y
Geociencias del NOA-Salta) for identifying the tadpoles.
GABRIEL CASTILLO, Universidad Nacional de San Juan Argentina. Di-
versidad y Biología de Vertebrados del Árido, Departamento de Biología,
San Juan, Argentina (e-mail:; GERALDINE RA-
MALLO, Instituto de Invertebrados, Fundación Miguel Lillo, San Miguel
de Tucumán, Argentina (e-mail:; CHARLES R.
BURSEY, Pennsylvania State University, Department of Biology, Shenango
Campus, Sharon, Pennsylvania 16146, USA (e-mail:; STE-
PHEN R. GOLDBERG, Whittier College, Department of Biology, Whittier,
California 90608, USA (e-mail:; JUAN CARLOS
ACOSTA, Universidad Nacional de San Juan Argentina. Diversidad y Bi-
ología de Vertebrados del Árido, Departamento de Biología, San Juan, Ar-
gentina (e-mail:
portant prey for numerous arthropod taxa, including ground
beetles (Toledo 2005. Herpetol. Rev. 36:395–399; Bernard and
Samolg 2014. Entomol. Fennica 25:157–160). Previous studies
have shown that Epomis larvae feed exclusively on amphibians
and display a unique luring behavior in order to attract their prey
(Wizen and Gasith 2011. PLoS ONE 6:e25161). Moreover, the lar-
val mandibles are characterized by two curved “hooks,” a modi-
fication for grasping onto the amphibian skin (Brandmayr et al.
2010. Zootaxa 2388:49–58). Published observations of Epomis
beetles attacking amphibians are scarce, and the majority of our
knowledge comes from reports originating in Japan (Crossland
et al. 2016. Herpetol. Rev. 47:107–108) or the Middle East (Wizen
and Gasith 2011, op. cit.). To the best of our knowledge, the only
record from India of amphibian predation by Epomis reports of
a ground-dwelling toad Duttaphrynus scaber carrying the beetle
larva (Barve and Chaboo 2011. Herpetol. Rev. 42:83–84).
Pseudophilautus amboli is a small endemic frog distributed in
the Western Ghats of India. It is known from a few localities only
in Maharashtra and Karnataka (
orinae/Pseudophilautus/Pseudophilautus-amboli; 20 Feb 2017).
This species is classified as Critically Endangered due to its narrow
distribution range, and is threatened by habitat loss and fragmen-
tation (; 2 Jun 2017).
Here we report predation of P. amboli by Epomis larvae in India.
At 2300 h on 22 October 2016, we performed an amphibian
survey at Amboli forest, a hilly location on the Northern West-
ern Ghats ridge in Sindhudurg District of Maharashtra, India
(15.964681°N, 74.003616°E, WGS 84; 690 m elev.). During our visit
we observed several juveniles of P. amboli, active on broad leaves
in the forest, approximately 20 cm above the ground surface.
Upon close inspection, we noticed that five of these specimens
(SVL ca. 40 mm) had small beetle larvae attached to their bod-
ies (Fig. 1). GW identified the larvae as Epomis sp. based on his
work with this genus and its interactions with amphibians. All
larvae observed on P. amboli were first-instars attached to the
throat area, and some had their head embedded deep inside the
amphibian’s flesh (Fig. 1B). Nevertheless, the frogs were still alive
and did not show any sign of struggling. They seemed to behave
normally and moved about in the vegetation without problems.
The amphibians and larvae were not collected.
The infected P. amboli may have encountered the Epomis
larvae on vegetation above the ground surface, similarly to what
is reported for E. nigricans larvae attacking tree frogs in Japan
(Tachikawa 1994. In Amazing Life of Insects, Atlas 48th Special
Exhibition. Otaru Museum, Otaru. 20 pp.). The location of the
larvae on the amphibians’ bodies suggests that they enticed the
frogs to approach by displaying their characteristic luring be-
havior (summary in Wizen and Gasith 2011, op. cit.). Moreover,
because Epomis larvae feed exclusively on amphibians in a para-
sitic manner, the interaction is usually fatal to the amphibian.
Our observations serve as evidence for the existence of a stable
breeding population of Epomis beetles in the area that relies on
the frogs as its main food source. This calls for further research
to monitor and evaluate the impact of the beetles on the popula-
tion of the Critically Endangered amphibian.
We thank Nirman Chowdhury and Gargi VR for assistance in
the field.
GIL WIZEN, 602-52 Park St. E, Mississauga, Ontario L5G 1M1, Can-
ada (e-mail:; ANISH PARDESHI (e-mail: anish- and KAKA BHISE, Malabar Nature Conservation
Club, 591 Amboli Bazar, Amboli, Sawantwadi Taluka, Sindhudurg District,
Maharashtra, India (e-mail:
RANA BOYLII (Foothill Yellow-legged Frog). PREDATION. Rana
boylii lives and breeds primarily in perennial stream habitats of
Fig. 1. Epomis sp. larvae preying on Pseudophilautus amboli in Am-
boli forest, Sindhudurg District of Maharashtra, India. A) A juvenile P.
amboli metamorph active on the vegetation with an Epomis sp. larva
attached to its throat. B) Epomis sp. larva with its head embedded in
the flesh of P. amboli.
Herpetological Review 48(3), 2017
the Pacific Coast region of western North America, from cen-
tral Oregon to Baja California (Zweifel 1955. Univ. California
Publ. Zool. 54:207–292; Hayes et al. 2016. Gen. Tech. Rpt. PSW-
GTR-248). Rana boylii is a Species of Special Concern in Califor-
nia and population declines have been primarily attributed to
altered flow and temperature regimes resulting from the con-
struction of dams along many of their native streams (Kupfer-
burg 1996. Ecol. Appl. 6:1332–1344). Chytrid fungus (Batracho-
chytrium dendrobatidis) may have played an important role in
the decline of R. boylii in California (Padgett-Flohr and Hopkins
2009. Dis. Aquat. Org. 83:1–9), and so might the introduction of
non-native species such as the American Bullfrog (Lithobates
catesbeianus; Kupferberg 1997. Ecology 78:1736–1751). Little is
known about the impact that predation might have had on R.
boylii populations, but predation of adult R. boylii by L. catesbe-
ianus has been reported in at least two stream systems in north-
ern California (Crayon 1998. Herpetol. Rev. 29:232; Hothem et al.
2009. J. Herpetol. 43:275–283). Here, I report on the predation of
a R. boylii larva by L. catesbeianus.
Copeland Creek is the primary drainage of the 255.8-ha Mit-
sui Ranch, located at the top of Sonoma Mountain in Sonoma
County, California, USA (38.335834°N, 121.579652°W; WGS 84).
The creek drains a landscape comprised primarily of annual
grasslands with isolated stands of Umbellularia californica (Cali-
fornia Bay Laurel), Quercus lobata (Valley Oak), and Q. agrifiolia
(Coast Live Oak). On 25 May 2015, while surveying the creek, I
encountered and collected three L. catesbeianus in a small pool.
The stomach of one small L. catesbeianus (SVL 70 mm; gape 27
mm) contained one snail (Physidae), one beetle (Hydrophylli-
dae), one water strider (Gerridae), and one R. boylii larva at de-
velopmental stage 28 (Gosner 1960. Herpetologica 16:183–190).
Rana draytonii and L. catesbeianus do not breed in this section
of Copeland Creek but they use it for dispersal, foraging, and
as a moist corridor to move between ponds when flows are low
enough to permit these activities. Thus, I assume predation of
R. boylii larvae by L. catesbeianus is opportunistic rather than a
constant pressure.
I thank the Sonoma Mountain Ranch Preservation Founda-
tion for allowing access and providing opportunity.
JEFFERY T. WILCOX, Sonoma Mountain Ranch Preservation Founda-
tion, 3124 Sonoma Mountain Road, Petaluma, California 94594, USA; e-
Tadpole aggregation behaviors have been observed in many
North American anuran families, including Bufonidae, Hylidae,
Leptodactylidae, Microhylidae, Ranidae, Rhinophrynidae, and
Scaphiopodidae (Wells 2007. The Ecology and Behavior of Am-
phibians. The University of Chicago Press, Chicago, Illinois. 1148
pp.). Benefits of aggregations include increased foraging oppor-
tunities, increased resource availability, physiological benefits,
and decreased predation. However, this could come at the cost
of increased competition, cannibalism, and predation risk (Mc-
Diarmid and Altig 1999. Tadpoles: The Biology of Anuran Larvae.
The University of Chicago Press, Chicago, Illinois. 444 pp.; Wells,
op. cit.). Although the exact triggers influencing the occurrence
of aggregation in tadpoles are not fully understood, it is likely
strongly influenced by environmental and ecological parameters
and a balance between the benefits and costs of this behavior.
Observations of various forms of aggregation in spadefoot
(family Scaphiopodidae) tadpoles have resulted in the identifi-
cation of three major types of aggregation: feeding aggregations,
premetamorphic protective aggregations, and metamorphic
aggregations (Bragg 1965. Gnomes of the Night: The Spadefoot
Toads. University of Pennsylvania Press, Philadelphia, Penn-
sylvania. 127 pp.; Black 1973.. Ph.D. dissertation, University of
Oklahoma, Norman, Oklahoma. 221 pp.). Within this family, the
majority of studies have detailed aggregation behaviors in Plains
Spadefoot (Spea bombifrons) tadpoles (Bragg and King 1960. Was-
mann J. Biol. 18:273–289; Bragg 1964. Wasmann J. Biol. 22:299–
305; Bragg 1965, op. cit.; Black 1968. Proc. Oklahoma Acad. Sci.
49:13–14). Similarly, several studies have documented aggrega-
tion behaviors in tadpoles of the Eastern Spadefoot (Scaphiopus
holbrookii; Abbott 1884. Am. Nat. 18:1075–1080; Ball 1936. Trans.
Connecticut Acad. Arts Sci. 32:351–379; Richmond 1947. Ecology
28:53–67), Hurter’s Spadefoot (S. hurteri; Bragg 1956. Herpeto-
logica 12: 201–204; Bragg 1959. Wasmann J. Biol. 17:23–42; Bragg
1968. Wasmann J. Biol. 26:11–16), and Mexican Spadefoot (Spea
multiplicata; Dodd 2013. Frogs of the United States and Canada,
Volume 2. The Johns Hopkins University Press, Baltimore, Mary-
land. 982 pp.). Compared to these species, considerably less is
known about Scaphiopus couchii. Black (1973, op. cit.) experi-
mentally demonstrated aggregation behaviors in S. couchii tad-
poles under laboratory conditions, and similar to Bragg (1965,
op. cit.), provided no account of this behavior in the field. Here,
we report an observation of aggregation behavior in S. couchii
tadpoles from the Chihuahuan Desert of west Texas.
On 3 July 2014, two large aggregations of Scaphiopus couchii
tadpoles were found in a shallow, ephemeral pool along a dirt road
on C. E. Miller Ranch, Jeff Davis County, Texas, USA (30.60548°N,
104.64239°W; WGS 84; Fig. 1). Each aggregation consisted of ap-
proximately 700 tadpoles that were visible at the surface of the
water. However, turbid water and the presence of individuals at
the bottom of the pool prevented a more accurate assessment of
the number of tadpoles in each aggregation. Surrounding these
large aggregations were smaller aggregations of 8–20 tadpoles. In
sum, the total size of each of these aggregations likely exceeded
1000 tadpoles. The majority of individuals were feeding at the
water surface in a vertical position, as described by Black (1973,
op. cit.). A group of 16 individuals (15 fluid preserved and one
preserved as tissue sample) were taken as vouchers to confirm
identification (following Altig and McDiarmid 2015. Handbook
of Larval Amphibians of the United States and Canada. Cornell
University Press, Ithaca, New York. 345 pp.) and deposited at
the Biodiversity Collections at the University of Texas at Austin
(TNHC 91983 [TJL 2691]). Although aggregations have been ob-
served in other Scaphiopus (see Bragg 1965, op. cit.), to the best
of our knowledge, this appears to be the first detailed description
of aggregation behavior in the field for S. couchii.
We thank the Miller family for their continued hospitality and
support of our herpetological research program, and J. Farkas for
reviewing earlier drafts of this manuscript. Specimens were col-
lected under a Texas Parks and Wildlife Department Scientific
Collecting Permit (SPR-1097-912) issued to TJL.
DREW R. DAVIS, Department of Biology, University of South Dakota,
414 East Clark Street, Vermillion, South Dakota 57069, USA (e-mail: drew.; TRAVIS J. LaDUC, Texas Natural History Collections, De-
partment of Integrative Biology, 10100 Burnet Rd, PRC 176–R4000, The Uni-
versity of Texas at Austin, Austin, Texas 78758-4445, USA (e-mail: travieso@
SCINAX NASICUS (Lesser Snouted Treefrog). PREDATION BY
cephalus typhonius is a large hylid with generalist feeding habits,
Herpetological Review 48(3), 2017
mostly feeding on insects and arachnids (Vaz-Silva et al. 2004.
Herpetol. Rev. 35:160; Dure and Kehr 2006. Herpetol. Rev. 16:109).
Records of occasional adult anuran prey include Hypopachus
variolosus (Dundee and Liner 1985. Herpetol. Rev. 16:109), Den-
dropsophus soaresi (Loebmann 2013. Herpetol. Notes 6: 275–
276), and Scinax ruber (Cintra et al. 2013 Herpetol. Rev. 44:500).
Here, we report a predation event by a male T. typhonius on an
individual of Scinax nasicus.
At 2345 h on 15 December 2016 at Alto Farm, municipality
of Aquidauana, Mato Grosso do Sul state, Brazil (19.57349°N,
56.15404°W; WGS 84), we found a male T. typhonius with most
the body of an S. nasicus already swallowed, bitten on its inguinal
region (Fig. 1). Swallowing was assisted by pushing the prey in
using the forelimbs. Scinax nasicus is a small hylid frog and has
been reported to be predated by a snake (Leptophis ahaetulla;
Lopez et al. 2003. Herpetol. Rev. 34:68–69) and birds ( Toledo et al.
2005 Herpetol. Bull. 92: 31–32; Ávila 2005. Herpetol. Rev. 29:169).
This is the first report of S. nasicus as prey of an anuran, and first
record of this species in the diet of T. typhonius.
We thank Marina and Lucas Leuzinger for logistical support
during our research at Barranco Alto farm. LSMS acknowledges
grant #2015/25316-6 from the São Paulo Research Foundation
(FAPESP) and for a Rufford Small Grant from The Rufford Foun-
dation. TSFS acknowledges productivity grant #310144/2015-9
from the National Council of Technological and Scientific Devel-
opment (CNPq).
LARISSA SAYURI MOREIRA SUGAI, Universidade Estadual Paulista
(UNESP), Instituto de Biociências, Rio Claro, and Programa de Pós-Gradua-
ção em Ecologia e Biodiversidade, 13506-900, São Paulo, Brazil (e-mail: (lar-; GABRIEL NAKAMURA DE SOUZA, Programa de Pós
Graduação em Ecologia, Universidade Federal do Rio Grande do Sul, Porto
Alegre, 90040-060, RS, Brazil; JOSÉ LUIZ MASSAO MOREIRA SUGAI, Cen-
tro de Ciências Biológicas e da Saúde, Universidade Federal de Mato Grosso
do Sul, Campo Grande, 79070-900, MS, Brazil; THIAGO SANNA FREIRE
SILVA, Universidade Estadual Paulista (UNESP), Instituto de Geociências e
Ciências Exatas, Rio Claro, 13506-900, SP, Brazil.
SCINAX RUBER (Red-snouted Treefrog). PREDATION. Birds
make up 15% of the main groups of vertebrate predators of post-
metamorphic frogs, and Toledo et al. (2007. J. Zool. 271:170–177)
recorded 38 species of anurans as prey for 27 species of birds in
the neotropical region. Herein we report a predation event on S.
ruber by Monasa nigrifrons (Black-fronted Nunbird; Fig. 1), a bird
that feeds on small vertebrates and large insects, mostly arthro-
pods, small lizards, and amphibians (Sherry and McDade 1982.
Ecology 63:1016–1028; Melo and Marini 1999. Ararajuba 7:13–15).
The event occurred on the morning of 1 October 2016 in a terra-
firme forest with Guadua bamboo located in the Zoobotanical
Park belonging to the Federal University of Acre, Brazil (9.9572°S,
67.8736°W, WGS 84, 165 m elev.). The attack method used by M.
nigrifrons was sally-strike (Remsen and Robinson 1990. Stud.
Avian Biol.13:144–160). The bird was perched on a bamboo stalk
at a height of 6 m and quickly flew down and captured the am-
phibian that was on a stalk about one meter from the ground.
After catching it, the bird returned to the same place and began
to manipulate the prey until it swallowed it (Fig. 1). This is the
first record of predation by M. nigrifrons on Scinax ruber.
JAILINI DA SILVA ARAÚJO, Laboratório de Herpetologia, Universidade
Federal do Acre CEP 69915-900 Rio Branco – Acre, Brazil (e-mail: jaillini@; EDSON GUILHERME, Laboratório de Ornitologia, Centro de
Fig. 1. Trachycephalus typhonius preying upon Scinax nasicus.
Fig. 1. Monasa nigrifrons preying on Scinax ruber.
Herpetological Review 48(3), 2017
Ciências Biológicas e da Natureza – CCBN, Universidade Federal do Acre
CEP 69915-900 Rio Branco – Acre, Brazil (e-mail:
phibians are an important part of the diet of many predators. For-
ty-five percent of predation events on amphibians, particularly
anurans, are by snakes (Toledo et al. 2007. J. Zool. 271:170–177).
Here we document a new predator-prey interaction between a
neonate Mexican Dusky Rattlesnake (Crotalus triseriatus) and a
Spea multiplicata.
At 1530 h on 24 May 2016 in San Gaspar Tlahuelilpan, Mete-
pec, Estado de México, México (19.244665°N, 99.559191°W, WGS
84, 2583 m elev.), a local gave us a living newborn rattlesnake (C.
triseriatus; total length = 205.5 mm; 9.2 g) in a plastic bottle. A
few minutes later the snake regurgitated a mostly undigested S.
multiplicata (total length = 70.7 mm; 4.8 g). The toad represented
34.4% and 52.1% of the snake’s total length and body weight, re-
spectively (Fig. 1). This record is consistent with reports of prey
consumption that represent 50%, or more, of snake body mass
in vipers (Mociño-Deloya et al. 2014. Rev. Mex. Biodiv. 85:1289–
1291; Rebón-Gallardo et al. 2015. Rev. Mex. Biodiv. 86:550–552).
After the toad was regurgitated, the snake had a slight ab-
dominal distension, as has been reported in neonates of similar
size (Mociño-Deloya et al. 2014, op. cit.; Rebón-Gallardo et al.
2015, op. cit.). Crotalus triseriatus is endemic to Central México
and considered a sit-wait generalist predator, eating some inver-
tebrates, such as arthropods, but mainly vertebrates such as sal-
amanders (Pseudoeurycea spp.), frogs, lizards (Sceloporus bican-
talis, S. grammicus, S. scalaris, S. torquatus), rodents (Microtus
mexicanus, Neotomodon alstoni, Peromyscus spp.), rabbits (Syl-
vilagus floridanus), and individuals of its own species (Mociño-
Deloya et al. 2014, op. cit.).
Tadpoles of S. multiplicata are preyed on by aquatic larvae
of scavenging beetles (Hydrophilus sp.), larvae of salamanders
(Ambystoma tigrinum), turtles (Kinosternon flavescens), grack-
les (Quiscalus sp.), and skunks (Spilogale putorius; Wright and
Wright 1949. Handbook of Frogs and Toads of the United States
and Canada. Comstock Publishing Associates, Ithaca, New York.
640 pp.). However, until this observation, known predators of
adults included only Thamnophis marcianus (Woodward and
Mitchell 1990. Southwest. Nat. 35:449–450).
JAVIER MANJARREZ, Laboratorio de Biología Evolutiva, Centro de inves-
tigación en Recursos Bióticos. Facultad de Ciencias, Universidad Autónoma
del Estado de México, Km 14.5 carretera Toluca – Ixtlahuaca, San Cayetano,
Toluca, Estado de México 50200, México (e-mail:
On 12 November 2016, 11.5 km SE of Valentine, Jeff Davis County,
Texas, USA (30.52166°N, 104.40198°W; WGS 84), we found two
Spea multiplicata impaled on a barbed wire fence at a known
Loggerhead Shrike (Lanius ludovicianus) larder. Gartersnakes
(Thamnophis sirtalis) are currently the only reported predator
of S. multiplicata (Woodward and Mitchell 1990. Southwest. Nat.
35:449–450; Dodd 2013. Frogs of the United States and Canada.
The Johns Hopkins University Press, Baltimore, Maryland. 982
pp.). To our knowledge, this is the first record of predation by L.
ludovicianus upon S. multiplicata (Clark 2011. Sonoran Herpetol.
24:20–22; Dodd 2013, op. cit.). The specimens of S. multiplicata
were preserved in the Sul Ross State University James F. Scudday
vertebrate collections as SRSU 6929–6930.
HAM, Department of Biology, Geology, and Physical Sciences, Sul Ross
State University, Alpine, Texas 79832, USA (e-mail: sean.graham@sulross.
edu); CRYSTAL KELEHEAR, Smithsonian Tropical Research Institute,
Apartado 0843-03092, Balboa, Ancon, Panama (e-mail: crystal.kelehear@
TION. When threatened, species of the genus Trachycepha-
lus release a sticky secretion presumably as a strategy to deter
predators (Delfino et al. 2002. J. Morphol. 253:176–186). Despite
Fig. 1. Partially digested Mexican Spadefoot (Spea multiplicata), top,
and the neonate Mexican Dusky Rattlesnake (Crotalus triseriatus)
that regurgitated it, bottom. Fig. 1. Trachycephalus typhonius being consumed by Leptophis
Herpetological Review 48(3), 2017
this defense, amphibians (Langdref-Filho et al. 2012. Herpetol.
Rev. 43:472) and snakes (De Freitas and Lima 2012. Herpetol.
Rev. 43:472) have been reported to feed upon Trachycephalus.
Among the snakes, Leptophis ahaetulla (Albuquerque and Di-
Bernardo 2005. Herpetol. Rev. 36:325), Liophis poecilogyrus (Silva
et al. 2003. Herpetol. Rev. 34:68), and Clelia bicolor (Prado 2003.
Herpetol. Rev. 34:231–232) are known predators of T. venulosus.
On the night of 10 December 1999, along the Sucuri River, Mato
Grosso do Sul state, Brazil (21.254479°S, 56.570091°W; WGS 84),
we observed a Leptophis ahaetulla preying on a female T. typho-
nius (Fig. 1). The behavior was recorded on the trunk of a tree,
about two meters from the ground. To our knowledge this is the
first report of T. typhonius as prey of L. ahaetulla.
NELSON RODRIGUES DA SILVA, Laboratório de Herpetologia,
Universidade Federal de Alagoas, 57072-970, Maceió, Alagoas, Brazil
SOUZA, Laboratório de Prática de Ensino de Biologia, Universidade
Federal de Mato Grosso do Sul, 79.070-900, Campo Grande, Mato Grosso
do Sul, Brazil (e-mail:
DERMATONOTUS MUELLERI (Mueller’s Termite Frog).
interspecific amplexus are found between living and dead
individuals of many anuran species (Bedoya et al. 2014. Herpetol.
Notes 7:515–516; Ribeiro et al. 2014. Herpetol. Rev. 45:479–480;
Sodré et al. 2014. Herpetol. Notes 7:287–288), though this behavior
is more common among species with explosive breeding (Wells
2007. The Ecology and Behavior of Amphibians. University of
Chicago Press, Chicago, Illinois. 1148 pp.). During breeding
events necrophilic interactions may occur, often when males
drown females, but continue to perform reproductive behaviors
after her death (Izzo et al. 2012. J. Nat. Hist. 46:2961–2967). In this
work, we report two cases of interspecific amplexus and one of
necrophilia, involving Trachycephalus typhonius (Hylidae) and
Dermatonotus muelleri (Microhylidade).
Our observations were made at night on 20–21 January 2016, in
a temporary lagoon in Aratuba, Ceará, northeast Brazil (4.4116°S,
39.0479°W, WGS 84; 841 m elev.). At 2350 h on 20 January 2016, an
adult T. typhonius was observed amplexing an adult D. muelleri
in the water (Fig. 1A). At 2355 h on the same date, we observed
another adult individual T. typhonius amplexing a female D.
muelleri out of the water (Fig. 1 B). At 2203 h on 21 January 2016,
a case of necrophilia was observed between an adult T. typhonius
and a male D. muelleri (with blackened vocal sac) inside the
water (we moved the pair onto the ground for photography; Fig.
1C). Each observation lasted about four minutes.
A case of necrophilia has already been recorded among con-
specific T. typhonius (de Moura and Loebmann 2014. Herpetol.
Bras. 3:60–61). To our knowledge, the present work reports the
first record of interspecific amplexus and necrophilia between T.
typhonius and D. muelleri and indeed, among the families Mi-
crohylidae and Hylidae.
FREDE LIMA-ARAUJO, Programa de Pós-Graduação em Ecologia
e Recursos Naturais, Universidade Federal do Ceará, Campus Pici, CEP
60455-760, Fortaleza, Ceará, Brazil (e-mail:; ANA
CAROLINA BRASILEIRO, Programa de Pós-Graduação em Ecologia
e Recursos Naturais, Universidade Federal do Ceará, Campus Pici, CEP
60455-760, Fortaleza, Ceará, Brazil (e-mail:;
SANJAY VEIGA MENDONÇA, Pós-Graduação em Ciências Veterinárias,
Universidade Estadual do Ceará, Campus do Itaperi, CEP 60740-00,
Fortaleza, Ceará, Brazil.
Batagur trivittata is a large (carapace length [CL] to 580 mm),
critically endangered (< 10 adult females survive in the wild),
Fig. 1. Interspecific amplexus in the water (A), interspecific amplexus
on land (B) and necrophilia (C) between Trachycephalus typhonius
and Dermatonotus muelleri.
Herpetological Review 48(3), 2017
aquatic turtle endemic to the major rivers of Myanmar (Ernst
and Barbour 1989. Turtles of the World. Smithsonian Institution
Press, Washington, D.C. 313 pp.; Rhodin et al. 2011. Turtles in
Trouble: The Worlds’ 25+ Most Endangered Tortoises and Fresh-
water Turtles – 2011. IUCN Tortoise and Freshwater Turtle Spe-
cialist Group, Lunenburg, Massachusetts. 54 pp.). Adult B. trivit-
tata exhibit pronounced sexual dichromatism; mature males
have an olive-green carapace with a black vertebral stripe and
two flanking dark lateral stripes, the plastron is ivory white, the
neck is yellowish, and the head is bright yellow-green with a
prominent black stripe extending backwards from the nostrils.
In contrast, the carapace, plastron, and head-neck of the much
larger mature females are uniformly dark brown to gray-black
(Theobald 1867–68. J. Linn. Soc. Zool. 1868:4–64; Smith 1931.
The Fauna of British India, including Ceylon and Burma. Vol. 1.
Loricata and Testudines. Taylor and Francis, London. 185 pp.).
Hatchlings and juveniles of both sexes have a dark greenish-
brown carapace with a pale yellow plastron. Although sexual
dichromatism has long been recognized in B. trivittata (e.g.,
Theobald, op. cit.), the age and body size at which these differ-
ences become apparent has hitherto gone unreported. We here
describe the development of sexually dimorphic coloration in
two cohorts of B. trivittata being reared in captivity as part of
a head-starting conservation program (Platt et al. 2014. Turtle
Survival 2014:45–48).
The cohorts of young turtles on which our observations are
based were hatched from eggs deposited by wild female B. trivit-
tata during February–March 2012 and 2013 in sandbanks along
the Chindwin River in Sagaing Region, Myanmar. We collected
the eggs shortly after deposition (< 24 h) and incubated them
under natural conditions at a secure sandbank near Limpha Vil-
lage (25.80570°N; 95.52200°E; India-Bangladesh datum). Upon
hatching (late May through early June), the neonates were trans-
ferred to 946-liter fiberglass tanks and maintained for three years
(stocking density of 20–25 hatchlings/tank during the first year,
reduced to 10–12 juveniles/tank in subsequent years). Young
turtles were reared on a diet composed primarily of chopped wa-
ter spinach (Ipomoea aquatica; provided on alternate days) and
commercial cat kibble (provided once per week). At three years
of age, each cohort was transferred to a concrete rearing pond
(9.1 × 12.1 m and 12.1 × 12.1 m; both ponds ca. 1.5 m deep) and
watermelon and figs (Ficus glomeratus) were added to the diet
when available. We weighed, measured, and recorded the shell
and head coloration of each turtle during February or March
2013–17. The 2012 and 2013 cohorts consist of 85 and 135 turtles,
Differences in coloration among individuals in each cohort
were not apparent during the first two years post-hatching;
plastrons remained white to yellow, carapaces were a uniform
greenish-brown, and head coloration a dull green. At age three,
a slight darkening of the plastron was evident on a few turtles,
but otherwise plastron, carapace, and head coloration appeared
unchanged from previous years. Sexual dichromatism first
became evident at age four in both cohorts. At this age, females
began to display areas of dark pigmentation on the plastron,
plastral bridge, and undersurface of the marginal scutes. The
degree of pigmentation varied greatly among individuals with
some displaying extensive, well-defined dark patches while in
others pigmentation was faint although discernible, and confined
to small areas of each scute. On the plastron and underside of
the marginal scutes, darkening appears to begin in the posterior
distal-most corner of each scute and radiates upwards and
outwards towards the mid-line. Darkening of the plastral bridge
followed a similar pattern except the pigmentation extends
upwards and outwards, away from the mid-line. In four-year-old
males, the plastron becomes bright cream-white, the black head-
stripe is obvious, and the head assumes a subdued greenish
hue. The carapace is gray-brown and the three black carapacial
stripes typical of adult males are not yet visible. A small number
(< 10%) of individuals in each cohort retained juvenile coloration
at age four and could not be reliably sexed. The mean (± 1 SD) CL
of four-year-old B. trivittata was 210 ± 38 mm (N = 220; range =
113–280 mm). The mean CL of males (CL = 212 ± 31 mm; range =
113–260 mm; N = 71) was slightly larger than females (CL = 208 ±
41 mm (range = 116–280 mm; N = 137), although this difference
was not significant (two-tailed t-test with unequal variances; t =
-0.67; DF = 176; P = 0.50).
At age five (2012 cohort only), dichromatic coloration was
more pronounced. Among many females, dark pigmentation of
the plastral bridge and underside of the marginal scutes is near-
complete or complete (Fig. 1A). Pigmentation of the plastron
in most females was extensive, although as with four-year-old
turtles, considerable individual variation was noted (Fig. 1B).
Fig. 1. Sexual dichromatism
among a five-year-old cohort of
Batagur trivittata hatched from
eggs collected along the Chind-
win River in Myanmar. Heav-
ily pigmented plastron, plastral
bridge, and marginal scutes of
female (1A). Less extensive plas-
tral pigmentation in this female
illustrates the wide range of in-
dividual variation present within
the same cohort (1B). Uniform
grey-black coloration of the head
and neck of female (1C). Head
coloration in male (1D). Plastron
and thickened tail of male (1E).
Herpetological Review 48(3), 2017
The carapace and head-neck of females are grey-black (Fig.
1C). Among males, the yellow-green head coloration becomes
brighter and more defined, and faint striping is evident on the
carapace (Fig. 1D). These color changes in males are accompa-
nied by a pronounced thickening of the tail (Fig. 1E). The mean
(± 1 SD) CL of five-year-old turtles was 264 ± 18 mm (range =
224–303 mm). Five-year old females (CL = 270 ± 18 mm; range =
224–303 mm; N = 56) were larger than males (CL = 250 ± 11 mm;
range = range = 230–274 mm; N = 29) and this difference was
significant (two-tailed t-test with unequal variances; t = 6.05, DF
= 80, P < 0.0001).
In conclusion, we found that sexually dichromatic coloration
becomes apparent among B. trivittata when CL exceeds about
120 mm between the age of four and five years, at least under
captive conditions. We expect dichromatism to develop at a later
age among wild turtles, which presumably grow at a slower rate.
Whether there is a body size × age interaction as demonstrated
in some reptiles (e.g., Alligator mississippiensis; Joanen and Mc-
Nease 1987. In Webb et al. [eds.], Wildlife Management: Croco-
diles and Alligators, pp. 329–340. Surrey Beatty & Sons, Pty., Ltd.,
Chipping Norton, NSW) remains unknown. Furthermore, our
observations indicate dichromatism first becomes evident at
about the same age the sexes begin to diverge in body size. Shine
and Iverson (1995. Oikos 72:343–348) determined that females of
most turtle species attain sexual maturity at about 70% of maxi-
mum body size. According to Smith (op. cit.), female B. trivittata
may reach a CL of 580 mm. Based on this value female B. trivit-
tata should thus become sexually mature when CL reaches 406
mm or about twice the current (2017) mean CL of the 2012 co-
hort. Our observations therefore suggest that sexual dichroma-
tism among B. trivittata becomes evident well before females
reach sexual maturity.
We thank the Ministry of Environmental Conservation and
Forestry for granting us permission to conduct research in Myan-
mar. Fieldwork in Myanmar was made possible by generous
grants from Andrew Sabin and the Andrew Sabin Family Founda-
tion, Panaphil Foundation, Helmsley Charitable Trust, Margaret
A. Cargill Foundation, and United States Fish and Wildlife Ser-
vice. The field assistance of Tun Win Zaw and Moe Aung Thu was
critical to the success of our project. We also thank Deb Levinson
and Ruth Elsey for assistance with obtaining literature, and Lewis
Medlock for insightful comments on a draft of this manuscript.
This paper represents technical contribution number 6564 of the
Clemson University Experimental Station.
STEVEN G. PLATT (e-mail:, TINT LWIN (e-mail:, and MYO MIN WIN, Wildlife Conservation Soci-
ety - Myanmar Program, No. 12, Nanrattaw St., Kamayut Township, Yangon,
Myanmar (e-mail:; KALYAR PLATT, Turtle Survival Al-
liance - Myanmar Program, No. 12, Nanrattaw St., Kamayut Township, Yan-
gon, Myanmar (e-mail:; THOMAS R. RAINWATER,
Tom Yawkey Wildlife Center & Belle W. Baruch Institute of Coastal Ecology
and Forest Science, Clemson University, P.O. Box 596, Georgetown, South
Carolina 29442, USA (e-mail:
CHELONIA MYDAS (Eastern Pacific Green Sea Turtle). DIET.
Chelonia mydas is considered an opportunistic omnivore in all
stages of its development (Amorocho and Reina 2008. J. Exp. Mar.
Biol. Ecol. 360:117–124). Novel diet items have been reported,
including Ptilosarcus undulatus (Sea Pen) (Seminoff et al. 2002.
Copeia 2002:266–268); Pleuroncodes planipes (Pelagic Red Crabs)
(Lopez-Mendilaharsu et al. 2005. Aquat. Conserv. Mar. Freshw.
Ecosyst. 15:259–269); tunicates and crustaceans (Amorocho and
Reina 2007. Endang. Species Res. 3:43–51); hydrozoans, scypho-
zoans, nematodes, annelids, mollusks (Carrion-Cortez et al.
2010. J. Mar. Biol. Assoc. U.K. 90[5]:1005–1013); squids, octopus
(Riosmena-Rodriguez and Lara-Uc 2015. Herpetol. Rev. 46:617)
and sea urchins (Reséndiz et al. 2016. Herpetol. Rev. 47:282).
It has been suggested that this dietary diversity is a response
to the energetic requirements of these animals in the early life
stages, facilitating nutritional gains for development and matu-
ration (Bjorndal 1985. Copeia 1985[3]:736–751), and optimiz-
ing digestion time (Amorocho and Reina 2008. J. Exp. Mar. Biol.
Ecol. 360:117–124). It has also been noted that diet in C. mydas
is influenced by resource availability (Balazs 1980. NOAA Tech.
Memo. NOAA-TM-NMFS-SWFS-7; Garnett et al. 1985. Wildl. Res.
12:103–112) and that diet selection is linked to the composition
and capacity of their hind-gut microflora, which may change as
the turtles grow and/or occupy different habitats (Bjorndal 1980.
Mar. Biol. 56:147–154).
In 2016 we collected food samples from the esophagi of 39 ju-
venile and subadult C. mydas (mean body mass 39.30 ± 27.78 kg)
during three field forays, and recorded straight carapace length
(mean 74.42 ± 27.78 cm). Collection sites were “el Espinazo del
diablo” (27.919490°N, 114.194480°W), “el Dátil” (27.777139°N,
114.171358°W) and “la Choya” (27.645778°N, 114.091806°W) at
Ojo de Liebre Lagoon, Baja California Sur, Mexico. In all samples,
a combination of the red algae Polysiphonia sp., Spyridia sp,. and
green algae Codium sp. were present and comprised 75% of the
total volume. Turtle mean body condition index (BCI) was 1.48
(range = 1.2–1.8), similar to the values reported for previous stud-
ies (Koch et al. 2007. Mar. Biol. 153[1]:35– 46; Seminoff et al. 2003.
J. Mar. Biol. Assoc. U.K. 83:1355–1362), which suggests that the
animals were in good nutritional status and had the capacity for
future favorable reproductive performance.
This is the first report of targeted Polysiphonia sp., Spyridia
sp. and Codium sp. consumption by C. mydas in Ojo de Liebre
Lagoon and in Baja California Sur. It has been established that
marine algae provide minerals, vitamins, carbohydrates, and
protein (Lourenço et al. 2002. Phycol. Res. 50:233–241); they also
stimulate the metabolism and immune system, and can reduce
heavy metals absorption (Ohta et al. 2009. Biol. Pharm. Bull.
32[5]:892–898). In addition, the ingestion of algae provide an im-
portant source of energy in sea turtles (Bjorndal 1997. In P. L. Lutz
and J. A. Musick (eds.), The Biology of Sea Turtles, pp. 199–231.
CRC Press, Boca Raton, Florida), increasing rates of growth and a
faster attainment of sexual maturity (Amorocho and Reina 2008.
J. Exp. Mar. Biol. Ecol. 360:117–124). The fact that Polysiphonia
sp., Spyridia sp. and Codium sp. accounted for 75% of the total
diet of C. mydas corroborates that these red and green algae are a
significant food resource for the turtles and the fact that the ani-
mals were in good nutritional status suggests the capacity to as-
similate nutrients from those algae species (Bjorndal 1990. Bull.
Mar. Sci. 47[2]:567–570). It is also important to note that these
algae species were not previously reported as components of C.
mydas diet in Ojo de Liebre lagoon or in Baja California Sur, sug-
gesting that turtles are able to adapt by shifting their food source.
Ojo de Liebre lagoon is an important feeding and development
area for C. mydas, and in these inshore foraging habitats, turtles
demonstrate high site fidelity (Balazs and Chaloupka 2004. Mar.
Biol. 145:1043–1059). For this reason, understanding feeding
ecology and diet of sea turtles is essential for their conservation
in these areas by identifying important food resources and allow-
ing informed decisions on the management of endangered pop-
ulations (Bjorndal 1999. In K. L. Eckert et al. [eds.], Priorities for
Herpetological Review 48(3), 2017
Research in Foraging Habitats, pp. 12–18. Research and Manage-
ment Techniques for the Conservation of Sea Turtles. IUCN/SSC
Marine Turtle Specialist Group Publication No. 4). It is desirable
to continue monitoring the different foraging areas in the Ojo
de Liebre Lagoon throughout the year to identify any seasonal
variation in the diet of sea turtles in relation to their age classes
and the presence of these algae and others nutrients.
DE SAN IGNACIO” Programa de Conservación de Especies en
Riesgo (PROCER) de la Comisión Nacional de Areas Naturales
Protegidas (CONANP) for funding this research and for their
assistance during field work. Thanks to Everardo Mariano-Me-
lendez and Oscar Javier Salazar-Mendez (Reserva de la Biosfera
el Vizcaíno) for their support and guidance during the develop-
ment of this research. Thanks to Aarón Sanchez, Fabian Castillo,
Joaquín Rivera and Antonio Zaragoza from Exportadora de Sal
S.A. (ESSA) for their assistance with fieldwork logistics and to
Carmen Méndez Trejo (Programa de investigación en Botanica
marina UABCS).
EDUARDO RESÉNDIZ, Alianza Keloni A. C. Antonio Rosales 698, col.
Centro, C.P. 23000, La Paz B.C.S. México and Universidad Autónoma de Baja
California Sur, Carretera al Sur KM 5.5., Apartado Postal 19-B, C.P. 23080,
La Paz, B.C.S. México (e-mail:; YOALLI HERNÁNDEZ-
Universidad Autónoma de Baja California Sur, Carretera al Sur KM 5.5.,
Apartado Postal 19-B, C.P. 23080, La Paz, B.C.S., México; MARÍA MÓNICA
LARA-UC, Alianza Keloni A.C. Antonio Rosales 698, col. Centro C.P. 23000,
La Paz, B.C.S., México and Universidad Autónoma de Baja California Sur,
Carretera al Sur KM 5.5., Apartado Postal 19-B, C.P. 23080, La Paz, B.C.S.,
México (e-mail:
FLOOD RESPONSE. Snapping Turtles are ranked as S3 in Mani-
toba due to the relatively few known populations, and are list-
ed as Special Concern nationally (COSEWIC 2008. vii + 47pp.).
There is little known about population sizes, survivorship, clutch
sizes, habitat use, movement, and nest predation in Manitoba
(Norris-Elye 1949. Can. Field Nat. 63:145–147). Here we report
on habitat use and movement of Chelydra serpentina in south-
western Manitoba during non-flood (2010 and 2012) and flood
(2011) years. The 2011 flood in the Assiniboine River watershed
remained in flood stage for 100 days beginning in early May and
remained high through early August (
ing/2011/index.html; 1 October 2013). To date, there have been
few studies on the impacts of changes in flood plains on animal
spatial ecology (Hanke et al. 2015. Wetl. Ecol. Manag. 23:215–226;
Zeng et al. 2015. PLoS ONE 10:e0127387), and even fewer that
have investigated the impacts of flooding on reptiles (Bateman
et al. 2008. J. Arid Environ. 72:1613–1619; Yagi and Litzgus 2012.
Copeia 2012:179–190).
This study took place in the lower Little Saskatchewan Riv-
er, in southwestern Manitoba, Canada (49.8872°N, 100.1211°W;
WGS 84). The headwaters of the Little Saskatchewan are in Rid-
ing Mountain National Park and the river meanders southward
for approximately 150 km where it flows into the Assiniboine Riv-
er, approximately 15 km W of Brandon. The main study area was
located between Glenorky (north) and Kirkhams (south) bridges.
The straight-line distance between the most northern and south-
ern locations of the study area was 10 km; the distance along the
Little Saskatchewan River between these same two points was
12.5 km. The surrounding area is primarily agricultural and pas-
toral land. The vegetation community beyond the riparian habi-
tat is comprised of shrubs, such as Silver Sage (Elaeagnus com-
mutate), and trees including Aspen (Populus tremuloides) and
Burr Oak (Quercus macrocarpa) with occasional gaps in which
the grazed land reaches the riparian habitat of the river.
Nine Snapping Turtles were located and captured by hand or
net from14 May to 15 June 2010; one additional turtle was cap-
tured on 9 June 2011. Nine out of ten of the turtles were captured
in May 2010 or June 2011 in the same marsh (approximately 7.5
ha in size), located on the west side of the Little Saskatchewan
River and approximately 1.5 km S of the Glenorky Bridge (Fig.
1). One turtle was captured in a nearby rivulet in June 2010. We
recorded weight (kg), carapace width and length (cm), plastron
length (cm), pre-cloacal tail length (cm), post-cloacal tail length
(cm), water temperature (°C), and air temperature (°C) for each
turtle (Table 1). We determined sex by either the presence of
eggs and/or the ratio of the ratio of posterior plastron lobe to
pre-cloacal tail length (Mosimann and Bider 1960. Can. J. Zool.
38:19–38). All individuals were adults, and there were six males,
two females, and two of unknown sex. Each turtle was also given
a unique notch combination on the marginal scutes of the cara-
pace with a triangle file.
All turtles were restrained by placing their heads into a buck-
et, as described by (Galbraith and Brooks 1983. Herpetol. Rev.
14:115). A VHF radio transmitter (Advanced Telemetry Systems,
transmitter model R1930, 24 g, ATS, Isanti, Minnesota, USA) was
attached to the lower right side of the carapace. The area was first
sanded and cleaned, and the radio transmitter was adhered to
the shell with waterproof epoxy gel and Mighty Putty™ (Brown
et al. 1990. Can. J. Zool. 68:1659–1663). The transmitters have a
battery life of approximately three years. We tracked animals by
foot and canoe once a week using an ATS R410 VHF receiver (ATS,
Isanti, Minnesota, USA) during the summers of 2010 (14 May–21
October), 2011 (19 April–September 11), and 2012 (22 March–6
We calculated home range sizes (ha) using the minimum
convex polygon (MCP for 95% of points) in R v3.2.1 (adehabi-
tatHR, MCP; Table 2). We assumed that the transmitter had been
shed when we recorded 30 consecutive locations in the same
spot for an individual, and excluded these locations from the
home range calculations. Home range sizes were larger in the
flood year (2011) compared to non-flood years (2010 and 2012;
Table 2), although we were unable to complete statistical analy-
ses because of our small samples sizes.
When turtles were located we recorded the date, time, loca-
tion (UTM within 5 m), habitat, and behavior of each located
turtle (when observed). We categorized turtle locations into 6
types: river (main channel of the Little Saskatchewan River or
Assiniboine River), riverbank, splay channel (small flow of water
that was separate from the main river channel, but joined to the
main river channel at both ends), marsh (marshy areas typically
within 25–100 m of the river), field (flooded area that had been
cultivated), and rock (exposed rock in the river). We also catego-
rized the behavior of individuals (where we had a visual record)
into 3 categories: basking in the open, moving through the water,
or resting on the bottom. We were unable to evaluate difference
between the sexes because we only tracked two females.
The majority of our locations during tracking were in the river
(75%, N = 202), a smaller percentage were found in the marsh
(22%, N = 60), a handful of locations were in splay channels (3%,
N = 7), and a single location was in a field (0.3%, N = 1). They
Herpetological Review 48(3), 2017
also were more often found in the river during the summer (July
1st onwards; 86%, N = 140 out of 162 summer locations), than
in the spring (May and June; 57%, N = 62 out of 108 spring lo-
cations). In the spring, there were more locations in the marsh
(47%, N = 41 out of 108) compared to the summer (12%, N = 19
out of 162 summer locations). During tracking, animals were not
typically visible (20%, N = 56), and were less often visible during
the flood year (2011; 7%, N = 7) compared to the non-flood years
(2010: 29%, N = 30; 2012: 24%, N = 19). Visible animals were most
commonly found in the marsh (66%, N = 37) or on the riverbank
(18%, N = 10); they were less frequently found in the river (11%, N
= 6), in the splay channel (4%, N = 2) or on rocks in the river (2%,
N = 1). The Little Saskatchewan River is very turbid (especially in
the spring) so visibility in the river is limited to the near-shore,
shallow areas.
Water in the marsh, or splay channels was clear, and therefore
animals located in these areas were easy to see. Animals basking
on the edge of the marsh, riverbank, or on rocks in the river were
also easy to see. Because of this visibility bias, we only compared
the locations of visible animals for the marsh. More of the visible
animals in the marsh were seen in the water column (47%, N =
17), with fewer animals resting on the bottom (36%, N = 13) or
basking on the edge (17%, N = 6). Atmospheric basking behavior
(animal is out of the water) was highest in the marsh (10%, N = 6),
compared to the river (5%, N = 11), or splay channels (0%).
Snapping Turtles observed in our study in southwestern
Manitoba appeared to have larger home ranges during the flood
year (2011) than in non-flood years (2010 and 2012). This in-
crease in home range size during flood events has been shown
in other turtle species. For example, flooded peatlands due to
beaver dams created new aquatic habitat (of high thermal qual-
ity) resulting in larger home range sizes in Spotted Turtles (Clem-
mys gutatta) (Yagi and Litzgus 2012. Copeia 2012:179–190). The
2011 flood on the Little Saskatchewan River did not appear to
Fig. 1. Radio telemetry locations for 10 Chelydra serpentina tracked in southwestern Manitoba in 2010–2012. The area flooded in 2011, but
returned to non-flood conditions in 2012.
table 1. Morphology, water temperature, and air temperature capture data for female (N = 2) and
male (N = 6) Common Snapping Turtles captured in the Little Saskatchewan River in southwestern
Manitoba in 2010–2011. Morphological measures included: mass, carapace length (CL), carapace
width (CW), and plastron length (PL).
Female Male
Mean ± SE Range Mean ± SE Range
Mass (kg) 7.2 ± 0.35 6.8 – 7.5 10.9 ± 1.49 7.2 – 17.3
CL (cm) 29.2 ± 0.40 28.8 – 29.6 35.1 ± 1.65 30.0 – 41.9
CW (cm) 26.4 ± 1.50 24.9 – 27.9 31.5 ± 1.73 27.3 – 38.9
PL (cm) 22.5 ± 0.35 22.5 – 23.2 25.8 ± 0.92 23.1 – 28.1
Water Temp. (°C) 17.0 ± 1.50 15.5 – 18.5 21.2 ± 1.89 16.0 – 25.5
Air Temp. (°C) 20.8 ± 4.75 16.0 – 25.5 22.6 ± 2.21 16.0 – 29.0
Herpetological Review 48(3), 2017
permanently create more aquatic habitat, because in 2012 they
had home range sizes comparable to pre-flood levels in 2010.
That said, there were differences in where they were spending
their time in the non-flood years. In 2010, they were all found
in the northern section of the river (Fig. 1), whereas in 2011 and
2012 some animals occupied the southern portions of the river,
where it intersected with the Assiniboine River.
Possibly, individuals were swept downstream in 2011, result-
ing in larger home ranges. There were more turtles located down-
stream during 2011 compared to 2010 (Fig. 1). Currents were not
measured along this stretch of the river during the flood event,
but were significantly higher along the nearby Assiniboine River
during the same time frame. On May 9, 2011, the Assiniboine
River in Brandon recorded a peak flow of 1280 m3/s, and a crest
at 364 m above sea level (
html; 1 October 2013). The southern edge of our study site in-
cludes the junction of the Little Saskatchewan and Assiniboine
Rivers (Fig. 1) so high water levels on the Assiniboine River were
directly affecting the Little Saskatchewan River, in addition to
the high water volume on the Little Saskatchewan River itself.
Water levels remained high on both the Little Saskatchewan and
Assiniboine Rivers until August of 2011. Individuals may have
been swept downstream, and then chose to hibernate in these
locations and remained there during 2012. None of our tracked
turtles were in the southern portion of the Little Saskatchewan
River in 2010, but some were there in 2012 after the flood (Fig. 1).
It is also possible that females were moving downstream to
seek out nesting habitat. In this system, we typically found nests
and nesting females on sandy shorelines. During a flood event
there will likely be less nesting habitat available, because the san-
dy shorelines were flooded and no longer be available. This may
have caused the increase in female home range sizes. The two
females that we tracked both increased their home range sizes,
although our small samples sizes do not allow us to statistically
test for differences between males and females. Given that both
males and females increased their home range size, it is likely
that the differences we saw during flooding were a combination
of high flow rates and lack of nesting habitat. Given that climate
change is likely to increase the frequency and severity of flood
events, it is imperative that we further study these events so that
we can have a better understanding of how these events impact
animals both in the short- and long-term.
Thanks to N. Cairns, D. Hoysak, A. Hoysak, and P. Malcolm for
their assistance with data collection. Funding and in-kind sup-
port were provided by Brandon University Research Committee,
Brandon University Student Union, Department of Zoology/Bi-
ology, and Manitoba Conservation. All protocols were done with
the approval of the Brandon University Animal Care Committee
(2009-R05) and all necessary permits for field study were ob-
tained (Manitoba Conservation: WB11022, WB12410).
PAMELA L. RUTHERFORD, Department of Biology, Brandon Universi-
ty, 270 18th St., Brandon, Manitoba, Canada R7A 6A9 (e-mail: rutherfordp@; CHRISTOPHER D. MALCOLM, Department of Geography,
Brandon University, 270 18th St., Brandon, Manitoba, Canada R7A 6A9 (e-
Detailed information on the ecology of neonatal emydine turtles
is scarce (Costanzo et al. 2008. J. Exp. Zool. 309A:297–379). In-
deed, because of their small body size, and their high susceptibil-
ity to predation, field studies on neonatal emydids are logistically
complex. Accordingly, the foraging ecology, and thus the precise
composition of the diet of emerging young emydids is virtually
unknown. Despite this lack of detailed information, it is usually
assumed that neonatal emydine turtles rely on residual yolk un-
til nest emergence and that; after emergence their diet is com-
posed of gastropods and insects based on information gathered
on larger juvenile individuals (Ottonello et al. 2005. Amphibia-
Reptilia 26:562–565).
We studied Emys orbicularis, a typical emydine turtle species,
in “Brenne” one of the largest wetlands of central France. These
field studies include protection of nests, and subsequent moni-
toring of the emergence of neonates. At the end of the emergence
table 2. Home range sizes (ha) calculated for minimum convex polygons (MCP for 95% of points) for
10 turtles tracked in the Little Saskatchewan River in southwestern Manitoba from May 2010 to August
2012. For each individual, the number of times the turtle was located is indicated in brackets. For mean
values the numbers of individuals is indicated in brackets.
2010 2011 2012
144 M 3.98 (13) 40.10 (11) NA
160 M NA 16.88 (10) 48.09 (7)
182 M 14.12 (14) 125.27 (11) NA
223 M 68.00 (11) 160.64 (8) 67.15 (12)
286 M 11.68 (11) 4.36 (11) 19.45 (15)
307 M 3.95 (16) NA NA
240 F 11.25 (13) 320.85 (13) NA
323 F 66.43 (10) 162.27 (11) 112.73 (13)
203 NA 10.07 (11) NA NA
261 NA NA NA 21.93 (9)
Mn ± SE 23.69 ± 9.59 (8) 118.62 ± 42.03 (7) 53.87 ± 17.15 (5)
Range 3.95 – 68.0 4.36 – 320.85 19.45 – 112.73
Mn ± SE (F) 38.84 ± 27.59 (2) 241.56 ± 79.29 (2) NA
Mn ± SE (M) 20.35 ± 12.09 (5) 69.45 ± 31.06 (5) 44.90 ± 13.86 (3)
Herpetological Review 48(3), 2017
period, nests are excavated to assess the emerging success (num-
ber of viable young produced) as well as the presence and the
number of dead individuals, dead embryos, or undeveloped
At the end of May 2017, three weeks after the usual latest
emergence date, we excavated a nest that had produced two
turtles. The nest also contained one dead individual, three dead
embryos, and three undeveloped eggs. Surprisingly, we found an
additional living neonate at the bottom of the nest. Upon cap-
ture, this individual defecated, suggesting that it had ingested
solid food after birth while still being within the nest. Examina-
tion of the fecal pellet revealed that it was composed of the cara-
pace scutes of other neonatal turtles.
This observation indicates that neonatal E. orbicularis can
feed on solid food sources while still being within the nest. Al-
though we do not know if the turtle consumed was alive, this
seems unlikely, and to our knowledge, it is the first reported case
of intra-nest necro-cannibalism in an emydine turtle species. We
do not know whether the young had ingested parts of a (dead)
sibling because it was unable to leave the nest or if it remained in
the nest because it found food resources there. Future examina-
tions of nest contents may allow us to quantify the frequency of
such unusual behavior.
FREDERIC BEAU, RNN de Chérine, Maison de la Nature et de la
Réserve, 36290 Saint-Michel-en-Brenne (e-mail: rncherine.frederic@or-; FRANCOIS BRISCHOUX, CEBC UMR7372 CNRS-ULR, 79360 Vil-
liers en Bois, France (e-mail:
Intraspecific basking has been observed in a number of North
American turtle species (e.g., Weber and Layzer 2014. Herpetol.
Rev. 45:117; Jones and Cochran 2014. Herpetol. Rev. 45:311–312;
Hartzell et al. 2015. Herpetol. Rev. 46:621; Hartzell and Hartzell
2016. Herpetol. Rev. 47:453). On 28 June 2017 at 1330 h, I observed
and photographed interspecific basking of Glyptemys insculpta
with two Graptemys geographica (Fig. 1) within the North Branch
of the Susquehanna River, Columbia County, Pennsylvania, USA.
Identification of each species was confirmed with binoculars.
All three turtles were observed basking on a fallen tree emerging
from the river approximately 30 m from the riverbank within a
section of the river approximately 280 m wide and several meters
in depth. Each turtle was situated approximately 1 m away from
the other turtles and all turtles remained in the same positions
during approximately 10 minutes of observation.
Throughout their range, G. insculpta are known to occupy lo-
tic habitats ranging from small streams to large rivers; however,
some differences in typical lotic habitat use have been noted be-
tween populations (Harding and Bloomer 1979. HERP, Bull. New
York Herpetol. Soc. 15:9–26; Ernst and Lovich 2009. Turtles of
the United States and Canada, 2nd ed. Johns Hopkins University
Press, Baltimore, Maryland. 827 pp.). G. insculpta populations
associated with the Great Lakes region typically occupy larger
streams and rivers, thus more often sharing habitat and bask-
ing opportunities with G. geographica (Harding 1990. In Beaman
et al. [eds.], Proceedings of the 1st International Symposium on
Turtles and Tortoises: Conservation and Captive Husbandry, pp.
31–35. California Turtle and Tortoise Club, Van Nuys, California;
J. Harding, pers. comm.). However, eastern populations of G. ins-
culpta more typically occupy smaller streams and overall appear
to spend more time away from water seasonally than western
populations of this species (e.g., Harding and Bloomer 1979, op.
cit.; Kaufman 1992. J. Herpetol. 26:315–321; Ernst 2001. Chelon.
Conserv. Biol. 4:94–99; Hartzell, pers. observ.). Because G. geo-
graphica typically inhabit large bodies of water (e.g., lakes, riv-
ers; Pluto and Bellis 1986. J. Herpetol. 20:22–31; Ernst and Lovich
2009, op. cit.), this observation of interspecific basking with G.
geographica appears to be unusual for G. insculpta populations
in the eastern portion of their range.
SEAN M. HARTZELL, Department of Biological and Allied Health Sci-
ences, Bloomsburg University of Pennsylvania, Bloomsburg, Pennsylvania
17815, USA; e-mail:
PODOCNEMIS EXPANSA (Giant South American River Turtle).
JUVENILE MOVEMENT. The Turtle Conservation Program of the
Lower Negro River was founded in 2014, and is focused on a pri-
ority area for chelonian conservation in the Amazon (Fagundes
et al. 2015. Divers. Distrib. 1–13). The program meets a demand
from the Brazilian riverine communities of three protected ar-
eas (PAs: Jaú National Park, Rio Unini Extractive Reserve, and
Rio Negro State Park North Section) to protect turtle species that
are an important food resource. One of the program’s principal
aims is to promote the conservation and management of nesting
Fig. 1. Glyptemys insculpta (arrow) basking with Graptemys
Fig. 1. Post-hatching Podocnemis expansa recaptured in the Jaú Riv-
er, Amazonas State, Brazil. The figure shows the individual marked
with the phalanx amputation method.
Herpetological Review 48(3), 2017
sites for four different turtle species in the above mentioned PAs
through participatory monitoring.
In December 2015, we marked 510 Podocnemis expansa
hatchlings in the Jaú River, Amazonas State, Brazil (2.2535°S,
62.6510°W; WGS 84). We used the phalanx amputation method to
remove a toe from all hatchlings to indicate their birth year (Bal-
estra et al. 2016. Biodiv. Bras. 6:130). We then randomly selected
a sample of 30 marked hatchlings for taking biometric measures.
Hatchlings had a mean straight-line carapace length of 57 ± 3.60
mm (51.75–64.00, N = 30). Hatchlings remained in captivity for
one month before being released in Supiá Lake (01.904639°S,
61.465639°W; WGS 84). Nine months after their release we re-
captured one marked juvenile P. expansa (Fig. 1) on Cuxiaú Lake
(01.85660°S, 61.60416°W; WGS 84), 38.47 km upstream from Su-
piá Lake (Fig. 2). The recaptured individual had a straight-line
carapace length of 108 mm. We estimated that the juvenile trav-
eled an average of 4.27 km/month to reach Cuxiaú Lake.
The movements of Podocnemis species are strongly related
to the hydrological variation of Amazonian rivers (Alho and
Pádua 1982. Acta Amazon. 12:323–326; Fachín-Terán et al. 2006.
Chelon. Conserv. Biol. 5:18–24). Adults of P. expansa can migrate
hundreds of kilometers searching for nesting sites or feeding
areas during the dry and flooded periods, respectively. On the
other hand, knowledge about hatchling and juvenile movements
is scarce. A recent study suggests that P. expansa hatchlings may
follow adults to feeding areas immediately after leaving nesting
sites, as a post-hatchling parental care strategy (Ferrara et al.
2013. J. Comp. Psychol. 127:24–32). Conversely, adult females
tracked by satellite telemetry system in the Xingu River showed
different dispersion routes after nesting and only one female
stayed close to the nesting beach (Carneiro and Pezzuti 2015.
Herpetol. Rev. 46:244–245). Our findings provide the first record
of the movement of a P. expansa juvenile. We could not determine
if the juvenile migrated with other individuals.
Actions aimed at reversing the population decline of P. e x -
pansa should concentrate on understanding and protecting ju-
veniles and adults (Mogollones et al. 2010. Chelon. Conserv. Biol.
9:79–89) and our finding is a first step in achieving this goal.
FAGUNDES (e-mail:, CAMILA R. FERRARA (e-mail:, THAIS Q. MORCATTY, Wildlife Conservation Society
Brasil - Universidade Federal do Amazonas, Av. General Rodrigo Octávio,
6200, 69080-900, Manaus, Amazonas, Brazil (e-mail: tatamorcatty@gmail.
com); VIRGÍNIA C. D. BERNARDES, Centro de Estudos de Quelônios da
Amazônia, Instituto Nacional de Pesquisas da Amazônia, Av. André Araújo,
2936, 69067-375, Manaus, Amazonas, Brazil (e-mail: virginiacdbernardes@; RACHEL K. ACOSTA (e-mail:
br), MARIANA M. LEITÃO (e-mail: mariana.leitã, ANA
L. C. B. FIGUEIREDO (e-mail: , ANA F.
C. Z. TINTO, Instituto Chico Mendes – Rua Antenor Carlos Frederico, 69,
69750-000, Novo Airão, Amazonas, Brazil (e-mail:
br); POLLYANA F. LEMOS, Instituto de Pesquisas Ecológicas – Rod. Dom
Pedro I, Km 47, Nazaré Paulista, São Paulo, Brazil (e-mail:;
PAULO C. M. ANDRADE, Laboratório de Animais Silvestres, Universidade
Federal do Amazonas - Mini campus, Av. General Rodrigo Octávio, 6200,
69080-900, Manaus, Amazonas, Brazil (e-mail:
br); JAIME G. NERY-JÚNIOR, Departamento de Mudanças Climáticas e
Unidades de Conservação, Secretaria do Meio Ambiente – Rua Raimundo
Marques de Medeiros, 84, 69730-000, Novo Airão, Amazonas, Brazil (e-mail:; MARIA C. G. PEREIRA, Departamento de
Mudanças Climáticas e Unidades de Conservação, Secretaria do Meio
Ambiente – Av. Mário Ipiranga Monteiro, 3280, 69050-030, Manaus,
Amazonas, Brazil (e-mail:
DISTANCE MOVEMENT AND HOMING. Several turtle species
in riverine ecosystems move relatively long distances (Moll
and Moll 2004. The Ecology, Exploitation, and Conservation of
River Turtles. Oxford University Press, Oxford, UK; Ernst and
Lovich 2009. Turtles of the United States and Canada, 2nd ed.,
Johns Hopkins University Press, Baltimore, Maryland. 827 pp.).
Podocnemis expansa exhibits the longest known movements
of any riverine turtle, migrating 45–422 km between foraging
areas and nesting beaches in Amazon basin rivers such as the
Río Caquetá and Río Trombetas (von Hildebrand et al. 1997.
Aspectos de la Biología Reproductiva y Técnicas para su Manejo.
Disloque Editores, Santa Fe de Bogotá, Colombia; Vogt 2008.
Amazon Turtles. Biblios, Lima, Peru). In Malaysia, a marked
female Batagur baska was recaptured at a nesting site 80 km
upstream from its initial capture site in the Perak River (Moll
1980. Malaysian J. Sci. 6:23–62). In North America, MacLaren et al.
(2017. Herpetol. Rev. 48:180–181) reported a Pseudemys gorzugi
that traveled 35.5 km in 33 days from Dolan Falls Preserve in
Devil’s River, Texas, USA, downstream to its confluence with the
Rio Grande. They noted this might be the longest movement by
any freshwater turtle in North America. Pseudemys suwanniensis
is another freshwater turtle capable of moving long distances
(Carr. 1952. Handbook of Turtles. Cornell University Press,
Ithaca, New York). Six individuals of this species moved 4 km
in the Santa Fe River (SFR), the largest Florida tributary of the
Suwannee River; the longest distance was 5.9 km (Kornilev et
al. 2010. Chelon. Conserv. Biol. 9:196–204). Munscher et al.
(2015. Herpetol. Rev. 47:127) confirmed long range movement
of a juvenile P. suwanniensis that travelled 14.8 km downriver
between two springs that feed the lower Suwannee River, Florida,
USA. Here we add an additional observation of P. suwanniensis
in the Suwannee River drainage that extends the longest known
movement distance by a North American freshwater turtle.
On 5 May 2011, GRJ captured a subadult female P. suwan-
niensis (241 mm straight midline plastron length [PL]) by hand
while conducting a snorkeling survey of the turtle assemblage in
the SFR (Johnston et al. 2016. Bull. Florida Mus. Nat. Hist. 54:69–
103). The exact location of capture is unknown, but it was within a
1-km reach between Rum Island (29.8327°N, 82.6778°W; WGS84)
Fig. 2. Probable movement track of the juvenile of Podocnemis ex-
pansa between its capture in December 2015 and recapture in Au-
gust 2016, in the Jaú National Park, Amazonas State, Brazil.
Herpetological Review 48(3), 2017
and Ginnie Springs (29.8336°N, 82.6879°W), Gilchrist County,
Florida. The turtle was individually marked by drilling a unique
combination of holes in the marginal scutes and released at the
1-km reach where it was captured initially. On 14 July 2013, TT
and ES recaptured this individual (288 mm PL) in a baited hoop
trap in the Suwannee River estuary (29.3261°N, 83.1076°W; Dixie
County, Florida) while conducting a survey of the Suwannee Al-
ligator Snapping Turtle (Macrochelys suwanniensis) population
(Thomas 2013. M.S. thesis, University of Florida, Gainesville).
Our conservative estimate of the distance between her original
capture location and recapture at the mouth of the Suwannee
River is 130 km. She was released at the trap location and not
observed again until her recapture (293 mm PL) by GRJ on 1 July
2015 in the same 1-km reach of the SFR near Rum Island where
she was originally captured and marked. Sometime between July
2013 and July 2015 this subadult female travelled 130 km against
the river current from the Suwannee River estuary back to her
original capture location at Rum Island in the SFR, a minimum
roundtrip of 260 km.
We hypothesize that she was displaced from her normal
home range by strong currents that followed Tropical Storm Deb-
by on 26 June 2012, and then returned home as currents returned
to normal. Our ability to determine the frequency of this putative
response to flooding is limited by the small number (N = 29) of
P. suwanniensis opportunistically captured in baited hoop traps
in the Suwannee River operated by the Florida Fish and Wildlife
Conservation Commission during 2011–2013. During the six
years prior to the M. suwanniensis survey, we captured and indi-
vidually marked 1226 P. suwanniensis in the SFR. Of these, only
one was captured in the Suwannee River. We have no data to esti-
mate how many individuals or which demographic groups move
between the SFR and Suwannee River estuary, and whether they
occur routinely or only under certain environmental conditions.
Our observation, coupled with the capture of 13 unmarked P.
suwanniensis at the confluence of the Suwannee River with the
Gulf of Mexico, also confirms observations by Carr (1940. A Con-
tribution to the Herpetology of Florida. Univ. Florida Publ., Biol.
Sci. Ser. 3:1–118) that this species regularly forages in the Suwan-
nee River estuary.
We thank Ken Dodd, John Iverson, Vivian Páez, Anders Rho-
din, and Dick Vogt for their help with literature.
GERALD R. JOHNSTON, Department of Natural Sciences, Santa Fe
College, Gainesville, Florida 32606, USA (e-mail: jerry.johnston@sfcollege.
edu); JOSEPH C. MITCHELL, Florida Museum of Natural History, University
of Florida, Gainesville, Florida 32611, USA (e-mail: dr.joe.mitchell@gmail.
com); ERIC SUAREZ, Fish and Wildlife Research Institute, Florida Fish and
Wildlife Conservation Commission, 1105 S.W. Williston Road, Gainesville,
Florida 32601, USA (e-mail:; TRAVIS M. THOM-
AS, Nature Coast Biological Station, University of Florida, Cedar Key, Florida
32625, USA (e-mail: Travis.thomas@u.edu).
ENTANGLEMENT. Entanglement in human-made materials is
well documented in snakes (Barton and Kinkead. 2005. Soil Wa-
ter Conserv. Soc. 60:33A–35A; Kapfer and Paloski. 2011. Herpetol.
Conserv. Biol. 6:1–9), but is less frequently reported for terrestrial
At 1250 h on 15 April 2017, two living and one deceased Ter -
rapene carolina triunguis were observed in close proximity to a
roll of discarded woven-wire fencing in Cleveland County, Okla-
homa, USA (35.2446°N, 97.4002°W; WGS 84). One of the living T.
carolina triunguis had its claw and part of its foot entangled in
the fencing (Fig. 1). A second living individual was not entangled.
A third deceased individual was present, and while the cause of
death cannot be determined, it is plausible that it too may have
been entangled in the discarded fencing and succumbed. The
entangled T. carolina triunguis was released, and a portion of
the fencing was removed from the area. Photos of this event have
been deposited at HerpMapper (HM 185273).
Brisbin, Jr. et al. (2003. In Mitchell et al. [eds.], Urban Her-
petology, pp. 367–380. Society for the Study of Amphibians &
Reptiles, Salt Lake City, Utah) reported a single case of wire-fence
entanglement and mortality in Terrapene carolina carolina, Here
we provided further evidence of this threat to Terrapene spp.
CHRISTOPHER E. SMITH, Wildlife Research & Consulting Services,
LLC, PO Box 803, Lakeland, Minnesota 55043, USA (e-mail: christopher.
smith@; MARK W. PARKER, 126 E. Mosier St. Nor-
man, Oklahoma 73069, USA (e-mail:
MUM SIZE. During a snorkeling survey of the freshwater turtle
assemblage in the Ichetucknee River in Ichetucknee Springs
State Park, Columbia County, Florida, USA (29.965669°N,
82.761917°W; NAD 84) on 22 March 2015, we hand-captured an
unmarked female Trachemys scripta scripta that measured 336
mm straight midline carapace length (CLmid), 344 mm maximum
carapace length (CLmax), 317 mm straight midline plastron length
(PLmid), 326 mm maximum plastron length (PLmax), 156 mm shell
height (HT), 245 mm shell width (SW), and 6400 g mass (Fig. 1).
The Ichetucknee River is an 8.4 km spring-fed tributary of the
Santa Fe River in northern Florida with clear, thermally stable
(19.9–23.6°C) water that supports an abundance of submerged
aquatic macrophytes (Heffernan and Cohen 2010. Limnol.
Oceanogr. 55:677–688; Chapin and Meylan 2011. Herpetol. Con-
serv. Biol. 6:51–60). During 2014–2017, we captured and marked
103 T. s. scripta at this site; the second largest female in our
sample measured 242 mm CLmid and 228 mm PLmid. On 2 June
2001, JCM captured a large female T. s. scripta by hand in Dare
County, North Carolina (35.785906°N, 75.755947°W, NAD 84)
measuring 335 mm CLmax, 318 mm PLmax, 138 mm HT, 229 mm
SW, and 4500 g mass. The only other female caught at this lo-
cation was 218 mm CLmax, 205 mm PLmax, 89 mm HT, and 183
mm SW. Habitat in this part of North Carolina is pocosin wet-
lands in deep peat deposits (Weakley and Schafale 1991. Wet-
lands 11 [Supplement]:355–375). Most aquatic habitats are deep,
Fig. 1. Terrapene carolina triunguis with its hind foot entangled in
woven-wire fencing.
Herpetological Review 48(3), 2017
dark water, human-dug ditches along access roads. A 309 mm
CLmax and 294 mm PLmax female T. s. scripta came from the Great
Dismal Swamp National Wildlife Refuge in Virginia, also a region
with extensively-ditched pocosin wetlands (Mitchell 1994. The
Reptiles of Virginia, Smithsonian Institution Press, Washington,
D.C. 352 pp.). The large turtles we report here exceed previously
known maximum sizes for T. s. elegans (Platt and Rainwater 2003.
Herpetol. Rev. 34:242; Powell et al. 2016. Peterson Field Guide to
Reptiles and Amphibians of Eastern and Central North America.
Houghton Mifflin Co., Boston, Massachusetts. 494 pp.) and T. s .
scripta (Mitchell, op. cit.).
Very large female sliders may exist in other habitats and pop-
ulations, but some of the largest known until now have been in
ponds on barrier islands off the coast of South Carolina (Gibbons
et al. 1979. Georgia J. Sci. 37:155–159; DeGregorio et al. 2102.
Herpetol. Conserv. Biol. 7:307–312). These authors hypothesized
that large Alligator mississippiensis in the ponds ate the smaller
T. s. scripta and contributed to the rapid growth of large females.
Large alligators do not occur in pocosin wetland ditches and
rarely occur in spring runs (Johnston et al. 2016. Bull. Florida
Mus. Nat. Hist. 54:69–103; ECM, pers. obs.). Other selective forc-
es may have contributed to the exceptional growth of the large
females we report here, and we cannot rule out the possibility
that growth in freshwater turtles is highly individualistic (Cong-
don and van Loben Sels 1993. J. Evol. Biol. 6:547–557). We could
not ascertain whether these two large females are extremely old
individuals or had exceptional growth histories or both. Because
they were substantially larger than all the other individuals in
their respective populations, we caution against using data from
these turtles to infer an association between habitat type and
geographic location without additional information.
We thank Pete Butt, Georgia Shemitz, Tabitha Hootman, and
the many volunteers who help us conduct turtle surveys in the
Ichetucknee River. Bob Powell provided the source for the mea-
surement in the field guide. Our research in Ichetucknee Springs
State Park was facilitated by Ginger Morgan and Sam Cole un-
der Florida Department of Environmental Protection permit
number 03011712 to GRJ. JCM thanks the DoD Legacy Resource
Management Program for funding his research on Dare County
Bombing Range in North Carolina.
JOSEPH C. MITCHELL, Florida Museum of Natural History, University
of Florida, Gainesville, Florida 32611, USA (e-mail: dr.joe.mitchell@gmail.
com); GERALD R. JOHNSTON, Department of Natural Sciences, Santa Fe
College, Gainesville, Florida 32606, USA (e-mail: jerry.johnston@sfcollege.
edu); ERIC C. MUNSCHER, Department of Natural Resources, SWCA Envi-
ronmental Consultants, Houston, Texas 77040, USA (e-mail: emunscher@
DUCTION. During the 2015 alligator nesting season in southwest
Louisiana, alligator nests experienced flooding and egg loss. On
16 June 2016, Tropical Storm Bill made landfall along the west
Texas Gulf Coast, and on 18 June water levels rose to 67.7 cm
above marsh level in the coastal marshes of southwest Louisi-
ana. On 25 June, water levels had dropped to 54.8 cm, and on
29 June, had receded to 47.4 cm above mean marsh level. Using
these water levels and nest height measurements, an estimation
of the hatch rate was estimated for the 2015 season, near Grand
Chenier, Louisiana.
From studies conducted on Rockefeller Refuge (Grand Che-
nier, Louisiana, USA), nest construction and egg deposition
takes place over a 21-day period which typically extends from
early to late June (Joanen and McNease 1992. In Proceedings of
the 11th Working Meeting Crocodile Specialist Group, Volume I,
pp. 207–221). This study documented daily nesting activity and
related the nesting period to spring ambient temperatures. Nest-
ing occurred earliest when temperatures in March–May were the
highest (Joanen and McNease 1979. Int. Zoo Yearbook 19:61–66).
Using this information, egg deposition for the 2015 season was
predicted to commence on 5 June and end on 25 June, with peak
nesting near 14 June. On 8 June, approximately halfway through
the first week of egg deposition, the first ten nest containing eggs
were found. The age of the oldest eggs, determined from ban-
width measurements (Ferguson 1985. In Gans et al. [eds.], Biol-
ogy of the Reptilia: Development, pp. 329–491, John Wiley and
Sons, New York), was 2–3 days old, which put the date of deposi-
tion on 5–6 June, and thus the calculated laying dates correlated
very closely with the predicted date of 5 June as the onset of the
nesting season.
Using nest dimension data collected by Joanen (1969. Proc.
Southwest. Assoc. Game Fish Comm. Conf. 23:141–151), the bot-
tom of the egg cavity was found to be 35 cm below the top of
the nest. The results from this study also showed that the average
Fig. 1. Maximum known size Trachemys scripta female from Ichet-
ucknee Springs State Park, Florida, USA: A) carapace, B) plastron.
Herpetological Review 48(3), 2017
distance from the top of the nest to the top of the egg cavity was
17.4 cm, with an average egg cavity depth of 17.6 cm. Water lev-
els were recorded each week throughout the entire year in the
study area. Measured water levels for the month of June were
compared to the calculated dimensions of nest height and egg
cavity dimensions (Table 1). The average nest height of 10 nests
measured on 8 June was 91.4 cm above marsh level. The bottom
of the nest cavity was predicted to be 56.1 cm above marsh level.
With the bottom of the nest cavity at 56.1 cm and the measured
water levels at 50.0 cm, no eggs would be flooded during the first
week of egg deposition.
On 14 June, the midpoint of the nesting season, water levels
rose to 63.4 cm above marsh level. It was observed that females
tended to build their nests higher than the existing water levels.
Nest height was calculated to be 106.7 cm, with the bottom of the
egg cavities at 69.5 cm above the marsh floor. No eggs laid in the
second week of nesting would have been flooded; however, the
rise in water level during the week of 14 June would have inun-
dated the eggs laid during the first week.
On 22 June, water levels had begun to recede and were re-
corded at 61.6 cm. The tops of the nests constructed during this
period were predicted to have been 103.6 cm above marsh level,
with the bottoms of the egg cavities at 67.7 cm. Thus, no eggs laid
during the last week of nesting would have experienced flood-
ing. The water levels continued to fall, and were recorded to be
46.9 cm on 29 June in the study area. At this level, no eggs were
subjected to flooding, and water levels were below the egg cavi-
ties of the nests that were constructed during the first week of the
nesting period.
Studies conducted concerning the sequence of nesting and
clutch size indicated that the largest females that lay the largest
clutches (average 40 eggs/nest) construct nests during the first
week of the season. Approximately 25% of the total nesting ef-
fort (total eggs) is expended during the first week. The second
week comprises the bulk of the nesting effort (59%) and consists
of nests that average 38 eggs/clutch. During the third and final
week, smaller females construct nests that contain an average of
35 eggs/nest (Joanen and McNease 1992, op. cit.). The 2015 nest-
ing season was interesting in that water levels increased dramati-
cally during the second week of the season, forcing females to
build nests higher to accommodate the rising water. Nests that
were laid during the first week were flooded due to the increased
water levels, thus drowning the majority of the eggs and em-
bryos. Eggs laid during the second and third weeks of the season
were deposited in egg cavities constructed high enough above
the water levels to avoid losses to the rising water. Alligator eggs
are not affected by 2 h of submersion, but eggs flooded for 48 h
produce 100% mortality (Joanen and McNease1979, op. cit.).
Egg collections in the study area for the 2015 nesting season
revealed a 25% egg loss during the 2015 nesting season. A to-
tal of 21,435 eggs were collected from approximately 800 nests
within the study site. Apparently nest height, as demonstrated in
this study, is regulated by marsh water levels at the time of nest
construction and varies with water level fluctuations. The 19,077
hatchlings produced were incorporated into a Louisiana Depart-
ment of Wildlife-approved alligator ranching program which was
initiated in 1986, and allows farmers to collect eggs from private-
ly owned marshlands (Joanen et al. 1997. In C. H. Freese [ed.],
Harvesting Wild Species—Implications for Biodiversity, pp. 465–
506. The Johns Hopkins University Press, Baltimore, Maryland).
Our research was supported by the Miami Corporation. We
thank Chad Courville, Land Manager of Miami Corporation, for
help with collection of data and Darren Richard (Louisiana De-
partment of Wildlife and Fisheries) for supplying the water level
data at the study site. The authors would also like to thank Jerry
Savoie, the owner of Savoie’s alligator farm, for locating nests,
counting eggs, and allowing us to use these data.
TED JOANEN, 1455 Big Pasture Rd., Lake Charles, Louisiana 70607,
USA; MARK MERCHANT, Department of Chemistry, McNeese State Uni-
versity, 450 Beauregard Dr., 225 Kirkman Hall, Lake Charles, Louisiana
70609, USA (e-mail:
PLASIA. Alligator mississippiensis is known for its ability to resist
infection and survive with severe physical maladies (Merchant
et al. 2003. Comp. Biochem. Physiol. 136:505–513; Hayes 2016.
Alligators of Texas. Texas A&M University Press. College Station,
Texas. 227 pp.). Neoplastic diseases have been documented in
this species (Elsey and Nevarez 2011. Herpetol. Rev. 44: 503–504;
Elsey et al. 2013. Southeast. Nat. 12:31–34;) as well as in other
crocodilians (Janert 1998. J. Zoo Wildl. Med. 29:72–77; Huchzer-
meyer 2003. Crocodiles. Biology, Husbandry and Diseases. CABI
Publ., Wallingford, Oxon, UK. 337 pp.) and non-crocodilian rep-
tiles (Frye 1991. Biomedical and Surgical Aspects of Captive Rep-
tile Husbandry. Vol. II. Krieger Publ. Co. Malabar, Florida. 637 pp.;
Hes et al. 2007. Veterinarni Lekar 3:49–55).
We investigated a report of a male alligator (3.04 m, total
length) that was harvested on 15 September 2016 with an
abnormal “growth” on its front right forelimb (Fig. 1). This
individual was harvested on private property in Matagorda
County, Texas, USA, and sold in the commercial market trade;
therefore, histological or radiographic investigation was
not conducted. However, from photographic evidence we
concluded that the growth was most likely a large fibrosarcoma
or fibromyxoma, based on information from previous published
reports for A. mississippiensis. As indicated in previous reports
of neoplasia, palpation of the mass revealed that it was dense
rather than cystic or fluctuant. This individual did not seem to
be greatly affected by the mass in that it appeared healthy (i.e.,
not emaciated) and presumably able to feed and move normally.
Although determining exact age in crocodilians without
molecular techniques is considered subjective, this individual
appeared to be relatively old, based on total length, and had likely
persisted with the mass for some time, based on the relative size
of the mass. However, depending on the exact cause of the mass,
the growth could have been present from months to years.
The two previous reports of alligator neoplasia were from al-
ligators captured from the wild in coastal Louisiana. Based on a
table 1. Water levels at Superior Bridge in relation to predicted alliga-
tor nest dimensions.
Month June Water Bottom of egg Top of egg Top of
(day) levels (cm) cavity (cm) cavity (cm) nest (cm)
1 59.1
8 50.0 56.1 74.4 91.4
14 63.4 69.5 87.8 106.7
18 67.6 73.8 92.3 109.7
22 61.6 67.7 86.0 103.6
25 54.3 60.4 78.6 97.5
29 46.9 53.0 70.6 88.0
The marsh level is 30.1 cm
Herpetological Review 48(3), 2017
review of the available literature, there have been no reports of
neoplasia in A. mississippiensis from other portions of its range.
To our knowledge, this is the first report of a large neoplastic
growth in an alligator from Texas. This is contribution number
17-115 of the Caesar Kleberg Wildlife Research Institute.
CORD B. EVERSOLE (e-mail: and
SCOTT E. HENKE, Department of Animal, Rangeland, and Wildlife Sci-
ences, Caesar Kleberg Wildlife Research Institute (MSC 218), Texas A&M
University-Kingsville, Kingsville, Texas 78363, USA.
ligator mississippiensis is a generalist predator and scavenger,
consuming a wide variety of vertebrates and invertebrates (Ga-
brey 2010. Herpetol. Conserv. Biol. 5:241–250), including at least
40 species of birds (Gabrey and Elsey 2017. J. Louisiana Ornithol.
10:1–12). Here we document a new species of waterfowl in the
diet of American Alligator.
At 1215 h on 1 December 2016 (sunny, air temp. = 15.6°C),
using 10 x 50 mm binoculars, we observed an alligator (total
length ~ 1.4 m), 25 m from us, swimming with a dead adult male
Gadwall (Anas strepera) in its mouth, along the Maxent Canal,
Bayou Sauvage National Wildlife Refuge, New Orleans, Orleans
Parish, Louisiana, USA (30.05175°N, 89.87996°W, WGS 84; 0 m
elev.). After 2 min the alligator stopped swimming to float for 2
min. During this time a smaller alligator (total length ~ 1.1 m)
swam toward the larger alligator’s head and the duck carcass, ap-
proaching to within 0.6 m. The larger alligator then turned away
from the smaller alligator and resumed swimming. The alligator’s
location on the western border of the refuge suggests that it may
have either captured the duck or scavenged it. Although duck
hunting was not allowed on the refuge on that date, it was al-
lowed on adjacent private land. Reports of American Alligator
predation on ducks have been mostly from spring and sum-
mer on flightless juveniles and molting adults (Elsey et al. 2004.
Southeast. Nat. 3:381–390).
JENNIFER O. COULSON, Department of Ecology and Evolutionary Bi-
ology, Tulane University, New Orleans, Louisiana 70118, USA (e-mail: jcoul-; THOMAS D. COULSON, 64340 Fogg Lane, Pearl River,
Louisiana 70452, USA.
NON-NATIVE PREY. American Alligators are opportunistic
predators and their food habits have been well studied (Elsey et
al. 1992. Proc. Annu. Conf. Southeast. Assoc. Fish Wildl. Agencies
46:57–66, and references therein and below). Composition of
A. mississippiensis diet often varies due to regional limitation
in prey availability (Neill 1971. The Last of the Ruling Reptiles.
Alligators, Crocodiles, and their Kin. Columbia University
Press, New York. 486 pp.; Gabrey 2010. Herpetol. Conserv. Biol.
5:241–250). Additionally, differential digestion rates can lead
to over-representation of materials resistant to digestion or
under-representation of rapidly digested soft-bodied prey
items (Jackson et al. 1974. J. Herpetol. 8:378–381; Garnett 1985.
J. Herpetol. 19:303–304; Delany and Abercrombie 1986. J. Wildl.
Manage. 50:348–353; Platt et al. 1990. Northeast. Gulf Sci. 11:123–
130). We herein report on a novel prey item for A. mississippiensis,
which to our knowledge has not previously been reported.
During the sanctioned statewide autumn harvest of wild
A. mississippiensis one of us (EL) opportunistically observed
the presence of numerous non-native Pomacea maculata
(Giant Applesnail) in the stomach of a free-ranging adult A.
mississippiensis trapped in southeast Louisiana (Fig. 1). The TL
and sex of the A. mississippiensis specimen are unknown. The
A. mississippiensis was likely trapped on 4 September 2015; the
observation was made on 5 September 2015 when the carcass
was processed for the hide and meat markets. The processing
facility is located in Houma, Louisiana (Terrebonne Parish) and
A. mississippiensis processed there are generally trapped locally.
The stomach contained ca. 15–20 P. maculata opercula in addition
to Diamond-backed Water Snake (Nerodia rhombifer) remains.
Numerous (at least 20) other harvested A. mississippiensis
specimens had also consumed P. maculata; opercula were
commonly observed throughout the 2015 harvest season at the
same processing facility. When A. mississippiensis carcasses were
being unloaded from boats or trucks to be measured and placed
in cold storage, P. maculata opercula would be regurgitated from
the A. mississippiensis carcasses; numerous P. maculata were
typically expelled from each carcass. This observation prompted
EL to take the photograph (Fig. 1) of the actual stomach contents
of one specimen for documentation of this novel prey item. Of
interest, some local trappers likewise noted the presence of P.
maculata in harvested A. mississippiensis carcasses during the
2015 trapping season, which prompted them to use them as bait
during the 2016 autumn alligator harvest in sites in southeastern
Louisiana (Vacherie and Lake Des Allemands).
Native Pomacea paludosa (Florida Applesnail) have been ob-
served as prey of juvenile A. mississippiensis in Florida (Fogarty
and Albury 1967. Proc. Annual Conf. Southeast. Assoc. Game
Fish Comm. 21:220–222). In that study of 36 A. mississippiensis
caught in a single night in one canal, 24 stomachs contained
Fig. 1. Neoplastic mass on the front right forelimb of a wild American
Alligator (Alligator mississippiensis).
Herpetological Review 48(3), 2017
119 P. paludosa which comprised 65.8% of the stomach con-
tents (Fogarty and Albury, op. cit.). Other studies conducted in
Florida reported P. paludosa as important invertebrate prey for A.
mississippiensis (Delany and Abercrombie, op. cit.; Delany et al.
1986. Florida Field Nat. 16:90–96; Delany et al. 1999. Proc. Annu.
Conf. Southeast. Assoc. Fish Wildl. Agencies. 53:375–389). A re-
cent study conducted in Louisiana on food habits of 448 adult A.
mississippiensis reported unspecified “snails” were observed in
some stomach contents (Gabrey, op. cit.); it is unknown if these
were P. maculata.
Pomacea maculata is an invasive freshwater snail native
to South America, and has expanded and is now established
throughout the southeastern United States (Monette et al. 2016.
Southeast Nat. 15:689–696), including Louisiana (Byers et al.
2013. PLOS ONE 8[2]:e56812). This genus can pose risks to ag-
ricultural crops and human and wildlife health; laboratory stud-
ies demonstrated the neurotoxin linked to Avian Vacuolar My-
elinopathy (AVM) can be transferred by P. maculata to its avian
predators (Robertson 2012. Potential Threats of the Exotic Apple
Snail Pomacea insularum to Aquatic Ecosystems in Georgia and
Florida. Master of Science Thesis. University of Georgia, Athens,
Georgia. 74 pp.; Byers et. al., op. cit.). This could be a concern, as
A. mississippiensis prey on a variety of avian species (Gabrey and
Elsey J. Louisiana Ornithol. 10:1–10).
Other crocodilians have been adversely impacted by con-
sumption of non-native exotic species. Long-term monitoring of
Crocodylus johnstoni (Australian Freshwater Crocodile) popula-
tions shows a decline following invasion of the toxic prey species
Rhinella marina (Cane Toad; Letnic et al. 2008. Biol. Conserv.
141:1773–1782). However, the vulnerability to this deleterious
effect varies among populations of C. johnstoni (Somaweera et
al. 2012. Anim. Conserv. doi.10:1111/j.1469–1795.2012.00578x.;
Doody et al. 2014. Biol. Invasions 16:2303–2309), with some
populations having little or no mortality from ingestion of the
toads (Somaweera et al., op. cit.) while adverse impacts on other
populations may include local extirpation (Britton et al. 2013.
Wildl. Res. Observations on
another crocodilian, Crocodylus moreletii (Morelets Crocodile),
suggest it is able to tolerate consuming bufogenin from Marine
Toads (Platt and Rainwater 2007. Brenesia 67:79–81).
It would be of interest to determine to what extent A. mis-
sissippiensis consumes P. maculata; however it is important to
recognize the opercula are likely resistant to digestion and could
be over-represented in stomach content analyses (references
To our knowledge this prey species has not previously been
reported for A. mississippiensis despite how well studied food
habits are within the species. It may be beneficial to determine
if P. maculata are energetically advantageous as a prey item, per-
haps leading this taxa to become increasingly commonly con-
sumed by A. mississippiensis and other large adult crocodilians.
Whether consumption of possibly toxic P. maculata has adverse
effects on A. mississippiensis remains unknown; experiments to
evaluate acute toxicity are currently being conducted.
We thank Jeff Boundy for the snake identification and Gary
LaFleur, Jr. for assistance with suggesting photo documentation
of the P. maculata observation.
RUTH M. ELSEY, Louisiana Department of Wildlife and Fisheries, 5476
Grand Chenier Highway, Grand Chenier, Louisiana 70643 USA (e-mail:; ERIC LEDET, Louisiana Department of Wildlife and
Fisheries, 2000 Quail Drive, Baton Rouge, Louisiana 70898, USA; JACOBY
CARTER, U.S. Geological Survey, Wetland and Aquatic Research Center, La-
fayette, Louisiana 70506, USA.
ING AGGREGATION AND BEHAVIOR. Because crocodilians are
largely nocturnal, often wary, and turbidity may obscure under-
water activity, observing predation events in the wild is difficult
(Thorbjarnarson 1993. Copeia 1993:1166–1177; Platt et al. 2006.
Herpetol. J. 16:281–290), and hunting strategies are in general
poorly documented, even for some relatively well-studied spe-
cies (Dinets 2015. Ethol. Ecol. Evol. 27:244–250). On occasion,
feeding crocodilians aggregate into groups (defined here as 2
individuals) in which cooperative hunting may occur; however,
these behaviors remain under-reported in the literature and
consequently are not well understood (Doody et al. 2013. Ethol-
ogy 119:1–9; Dinets, op. cit.). Given the paucity of information on
feeding aggregations and cooperative hunting among crocodil-
ians, field observations are particularly noteworthy and essen-
tial for understanding these behaviors (Dinets, op. cit.). We here
describe a large feeding aggregation and associated behaviors of
A. mississippiensis as seen in a video clip and described in a per-
sonal communication provided by Thomas M. Braquet to RME.
The cellphone video (86 sec;
was filmed where Dupont Outflow Canal debouches into the Sa-
bine River in Orange County, Texas, USA, circa August 2009. The
surrounding habitat is freshwater marsh (sensu Chabreck 1972.
Bull. 664, Louisiana State Univ., Agric. Exp. Stat., Baton Rouge.
72 pp.), which in general supports high densities of A. mississip-
piensis (McNease et al. 1994. In Crocodiles: Proceedings of 12th
Working Meeting, Crocodile Specialist Group, pp. 108–120. IUCN
Publ., Gland, Switzerland). According to Braquet, who observed
Fig. 1. Viscera of harvested adult Alligator mississippiensis, with the
stomach opened to expose Pomacea maculata (Giant Applesnail)
Herpetological Review 48(3), 2017
the feeding aggregation for about 45 minutes before departing, a
large school of mullet (Mugil sp.) were attempting to swim into
the canal from the Sabine River. In the video, at least 50 alliga-
tors can be seen clustered around the mouth of the canal feeding
on the schooled fish. The aggregation appears to be composed
almost exclusively of adult alligators (total length 1.8 m). Four
egrets (probably Ardea alba) are also in view, perched in low
shrubs and along a fence, seemingly attracted by the schooling
fish. One alligator lunged out of the water in an unsuccessful at-
tempt to capture an egret perched in a shrub about 0.5 m above
the surface; the bird immediately took flight and then landed in
nearby shrub.
Braquet stated the foraging alligators approached the
schooled fish while swimming on the surface. The alligators
would then submerge, swim rapidly into the school, and resur-
face, often with a fish. Several alligators frequently approached
the school simultaneously, swimming side-by-side, before mak-
ing an underwater run at the fish, a behavior evident in the video
clip. Because mullet often school at or just below the surface, we
assume alligators were diving below the fish and then attempting
to capture them just before surfacing. According to Braquet, af-
ter consuming captured fish, individual alligators often repeated
this behavioral sequence several times before moving to the pe-
riphery of the feeding aggregation. Other alligators would then
swim forward and approach the school in the manner described
above. Numerous alligators can be seen in the video clip milling
about the periphery of the aggregation while others are actively
attempting to capture fish.
Also evident in the video are at least 19 (averaging one ev-
ery 4.5 seconds) instances of what Thorbjarnarson (op. cit.) de-
scribed as “leaping” behavior; alligators can be seen floating on
the surface and then suddenly propelling themselves upwards
and forward, partially emerging from the water with the body
arched, and rear- and fore-limbs clasped closely against the body,
which is then rotated to one side to avoid a mid-ventral (“belly”)
landing. Several leaping alligators appear to emerge from the be-
low the surface, perhaps the result of being propelled upwards
by their forward momentum when making an underwater rush
at schools of fish. In one instance, three or four mullet jump out
of the water immediately followed by an alligator leaping from
below the surface. Thorbjarnarson (op. cit.) characterized leap-
ing by foraging Caiman crocodilus as a high energy behavior with
a low success rate (0.8–1.0%).
Our report compliments a handful of previous accounts
that describe feeding aggregations of A. mississippiensis (Van
Doren 1928. Travels of William Bartram. Dover Publications,
New York. 414 pp.; Harper 1930. Sci. Monthly 31:56–67; Dinets
2010. Herpetol. Bull. 111:4–11; Dinets, op. cit.). As noted by
King et al. (1998. Cooperative feeding, a misinterpreted and
under-reported behavior of crocodilians. Available: http://www. several
commonalities emerge from these accounts, the foremost being
that aggregations assemble in response to dense concentrations
of fish. Second, feeding aggregations consist of unusually large
numbers of alligators concentrated within a relatively small
area, typically where one waterbody empties into another. Third,
intraspecific conflict over prey is rare to infrequent, probably
because the resource is so abundant that defending it from
conspecifics would prove energetically costly and yield few if
any benefits (e.g., Heinrich 1989. Ravens in Winter. Summit
Books, New York. 379 pp.). Fourth, individuals appear to rotate
in and out of the aggregation, i.e., when one alligator moves
away from the aggregation, another quickly takes its place. And
fifth, as seems to be the case in our video clip, feeding alligators
appear to disregard the presence of humans and other forms of
Feeding aggregations of crocodilians are frequently described
as cooperative behavior, an inclusive term encompassing
both coordinated (individuals relate in time and space to each
other) and more rarely, collaborative (individuals preforming
complimentary actions focused on same prey) hunting
strategies (Dinets 2015, op. cit.; Grigg and Kirshner 2015. Biology
and Evolution of Crocodylians. Cornell University Press, Ithaca,
New York. 649 pp.). As seen in the video clip, alligators appear
to be relating to each other, although nothing indicates that
complementary actions are being directed towards the same
prey. Therefore, our report could be described as an example of
coordinated hunting behavior. That said, Dinets (2015, op. cit.)
points out that evaluating putative examples of coordinated
or collaborative hunting behaviors is difficult because the
intentions of the participating animals can never be known with
certainty, echoing the earlier concerns of Gans (1989. Amer. Zool.
29:1051–1054) that such reports might describe nothing more
than accidental events rather than representative behaviors.
Indeed, Grigg and Kirshner (op. cit.) question whether feeding
aggregations should even be described as cooperative behavior
and called for additional research into these behaviors. Finally,
our report highlights the valuable role that citizen scientists
can play in recording rarely observed behaviors of crocodilians,
especially given the widespread availability of inexpensive digital
tools (Cohen 2008. Bioscience 58:192–197; O’Donnell and Durso
2014. Herpetol. Rev. 45:151–157).
We thank Thomas Braquet for providing the video clip and
describing his observations, and Deb Levinson, Thomas Rainwa-
ter, and Kent Vliet for obtaining literature.
STEVEN G. PLATT, Wildlife Conservation Society - Myanmar Pro-
gram, No. 12, Nanrattaw St., Kamayut Township, Yangon, Myanmar (e-mail:; RUTH M. ELSEY, Louisiana Department of Wildlife
and Fisheries, Rockefeller Wildlife Refuge, 5476 Grand Chenier Highway,
Grand Chenier, Louisiana 79643, USA (e-mail:
LISM. Cannibalism is a common phenomenon among farmed
and wild crocodilians that generally constitutes larger individu-
als preying upon smaller crocodilians (Somaweera et al. 2013.
Herpetol. Monogr. 27:23–51). Its significance as a mechanism
to regulate population density remains unclear given that can-
nibalism has not been thoroughly investigated in the wild, be-
cause of its difficulty to assess (Delany et al. 2011. Herpetologica
67:174–185). Yet, perhaps cannibalism is simply a response to the
lack of other food sources, or the consequence of intraspecific
competition for various resources among wild populations, such
as space (territory), food, or nesting habitat (Cedeño-Vázquez
et al. 2016. Mesoamer. Herpetol. 3:470–472). Competition for
resources could heighten the frequency of cannibalism among
conspecifics, especially during the breeding or nesting season.
Here, we report three occurrences of cannibalism observed in
wild Crocodylus moreletii during a field excursion in Chiquibul
Forest, Belize.
Chiquibul Forest is classified as lowland tropical broad-
leafed rainforest, and is the largest tropical rainforest remain-
ing in North America. It is home to a rich biodiversity of flora
and fauna, and is the largest protected area in Belize. In 2016,
co-managers of Chiquibul Forest Friends for Conservation and
Herpetological Review 48(3), 2017
Development (FCD) partnered with the Crocodile Research Co-
alition (CRC) to initiate research and a crocodile management
program for the population of C. moreletii in Chiquibul Forest.
During our initial nocturnal eyeshine survey of the Macal River
on the night of 23 June 2016, we observed a large crocodile (ca.
2.5 m in length) consuming a crocodile carcass at 1911 h. After
the crocodile discarded the carcass as we approached, we col-
lected the dead crocodile, a 1.8-m female. We observed a sec-
ond case of cannibalism 59 min later (2010 h), 3.7 km from the
first observation. The feeding crocodile (2.5 m in length) quickly
abandoned the carcass of a smaller crocodile upon our arrival
(ca. 5 m sighting distance). The dead crocodile was identified as
a 1.6-m female.
The third observation of cannibalism was observed at 2345 h
on 24 July 2016 on the Macal River during the capture and tagging
portion of the 2016 population survey in Chiquibul Forest. Inter-
estingly, this 1.7-m female was preyed upon in the same location
as the first observation the previous night. We confirmed that this
carcass was not the same individual from the night before as the
lower jaw of the crocodile was intact, whereas the lower jaw from
the crocodile the previous night was already removed. We sug-
gest that the larger crocodile observed predating on this carcass
was the same individual that consumed the smaller crocodile the
night before in the same location; the size appeared similar, and
the behavior upon our approach was comparable. Identification
of reproductive organs verified sex of all dead crocodiles, and we
approximated total length of dead crocodiles by measuring rear
of cranial plate–vent length (on the dorsal side). We returned all
3 crocodile carcasses immediately to the water once we recorded
GPS coordinates, length, and sex.
We thus report the first incidence of C. moreletii cannibalism
observed in Belize, as well as the second account of cannibal-
ism among wild C. moreletii (Cedeño-Vázquez et al., op. cit.).
Furthermore, most reports of cannibalism among wild croco-
diles witness a single observation. Here, we report three observa-
tions of cannibalism among C. moreletii in about a 29-h period.
In general, the magnitude and frequency of cannibalism among
crocodilians is unknown; however, we suggest that this type of
predation may be common among the population of C. moreletii
in Chiquibul, particularly in the Macal River during the breeding
season. Rangers of the FCD observed a fourth crocodile carcass
in the Macal River about three months prior to our survey. All
cannibalized females were of breeding size, and it would seem
unlikely that males would kill potential mating females. Fur-
thermore, good habitat for nesting sites along the Macal River is
limited. Thus, it is possible the cannibalized females were killed
by larger females as a consequence of territory and intraspecific
competition for quality nesting habitat, as our observations were
in close proximity to nests (ca. 238 m). Therefore, we suggest that
the larger males we witnessed preying upon the small-bodied
crocodiles were scavenging the remains of crocodiles killed by
females. Because we did not witness crocodiles killing other
crocodiles, it is also possible, but unlikely, that the carcasses were
crocodiles that died from some other cause (e.g., disease).
We thank the Belize Wildlife and Referral Clinic, Friends for
Conservation and Development, and the Belize Forest Depart-
ment for their support and assistance in this project. We also
thank the organizers of Canada CrocFest 2015 for funding this
field excursion and research.
MARISA TELLEZ, Crocodile Research Coalition, Maya Beach, Stann
Creek, Belize (e-mail:; SHAWN K. HEFLICK,
Crocodile Conservation International, Inc. (e-mail: she; MI-
RIAM BOUCHER, School of Forestry and Natural Resources, West Virginia
University, Morgantown, West Viriginia, USA; ANDREW AUSTIN, Houston
Community College, Houston, Texas, USA.
CROCODYLUS POROSUS (Saltwater Crocodile). DIET. Croco-
dylus porosus is the most widely distributed crocodilian species
in the world (Webb et al. 2010. In Manolis and Stevenson [eds.],
Crocodiles. Status Survey and Conservation Action Plan. Third
Edition, pp. 99–113. Crocodile Specialist Group, Darwin). In Sri
Lanka, it is found mainly in tidal rivers and marshlands outside
protected areas along the coastal wet zone and some parts of the
dry zone (Samarasinghe 2014. The Human-Crocodile Conflict in
Nilwala River, Matara [phase 1]. Young Zoologists’ Association
Publications, Sri Lanka. 118 pp.). Crocodylus porosus is an op-
portunistic ambush predator that is known to have the broadest
range of prey species in their diet (Ross and Garnet [eds.] 1989.
Crocodiles and Alligators. Facts On File, New York. 240 pp.). Juve-
niles hunt smaller prey such as insects, crustaceans, small fish,
amphibians, and reptiles. In adults, the significance of small in-
vertebrate prey fades in favor of vertebrates (Taylor 1979. Austr.
Wildl. Res. 6:347359). Adult C. porosus are known to feed on very
large prey, including buffalo, wild boar and monkeys. Here, we
report an adult C. porosus preying upon a previously unrecorded
prey, an Indian Porcupine (Hystrix indica).
The observation was made at 2130 h on 21 November 2015
in the Nilwala River in Matara District, Southern Province of Sri
Lanka, while conducting annual spotlight surveys on C. porosus.
The observation was made in mid-river, on a 4 m-long motor
boat using a 12 v, 400,000 candlelight glare free spot light, ca. 6.5
km upstream (5.96353°N, 80.5551°E). The observation lasted for
two minutes. An adult C. porosus, 2.6 m in total length, was spot-
ted in open water approximately 3.5 m away from the river bank.
The crocodile was swimming slowly along a straight line paral-
lel to the river bank. As we approached the animal to estimate
its size, we observed an adult Indian Porcupine in its jaws. Upon
encounter, the engine was turned off to reduce disturbance, and
the observation was made ca. 1.5 m away from the animal. We
did not detect any movement by the porcupine. The crocodile
had the neck and pectoral region of the porcupine secured in its
jaws, and the rest of the body was outside. We observed quills
intact in the body and tail region of the porcupine. After about
two minutes of observation, the crocodile dove underwater with
the carcass in its jaws.
Records of porcupine predation among crocodilians are rare.
The presence of quills and associated possibility of injury would
seem to make porcupines an unlikely prey item for crocodilians.
However, several crocodile species are known to prey on porcu-
pines: C. moreletii preying on Coendou bicolor (Platt et al. 2006.
Herpetol. J. 16:281–290), Paleosuchus trigonatus preying on Co-
endou mexicanus (Ortiz et al. 2013. Herpetol. Rev. 44:135), and
Crocodylus niloticus preying on Hystrix sp. (Piennar 1969. Koe-
doe 12:1). Magnusson et al. (1987. J. Herpetol. 21:85–95) observed
a specimen of P. trigonatus regularly feeding on porcupines, de-
spite having quills embedded in its jaws. Ortiz et al. (2013, op.
cit.) also reports a stressed P. trigonatus individual having quills
covering the head and jaws after a predation attempt on a porcu-
pine. Magnusson et al. (1987, op. cit.) considered porcupines to
be not hazardous to P. trigonatus. Because our observation was
brief, we were unable to conclude if the adult C. porosus we ob-
served was unharmed. However, we did not observe any quills
embedded around the head and jaws region of the crocodile.
Despite being one of the most well-studied crocodilians in the
Herpetological Review 48(3), 2017
world, ours is the first record of C. porosus preying on a porcu-
pine (Grigg and Kirshner 2015. Biology and Evolution of Croco-
dylians. CISRO Publishing, Clayton, Victoria, NSW. 672 pp.).
DINAL J. S. SAMARASINGHE, Environmental Foundation (Guaran-
teed Limited) 3A, 1st Lane Kirullapone, Colombo 5, Sri Lanka (e-mail: dinal.; DILIP ALWIS, University of Ruhuna, Tangalle Road,
Matara (e-mail:
ASITISM. The amphisbaenians are currently represented by
approximately 196 species (Uetz and Hošek 2016. http://www.; accessed 6 November 2016), belonging to
six families. The family Amphisbaenidae is the most diverse and
abundant in South America (Gans 2005. Bull. Amer. Mus. Natur.
Hist. 289:1–130; Vidal et al. 2008 Biol. Lett. 4:115–118; Uetz and
Hošek, op. cit.). Amphisbaena vermicularis is a medium-sized
amphisbaenid occurring from Brazil to Bolivia (Dirksen and De
La Viva 1999. Graellsia 55:199–215; Almeida et al. 2008. Braz. J.
Biol. 69:1183–1186). Here, we report on the helminth endopara-
sites in the gastrointestinal tracts of A. vermicularis in an area
of Caatinga vegetation in Exu, Pernambuco State, northeastern
In November 2013, a specimen of A. vermicularis (SVL: 16.2
cm; weight: 7.0 g) was collected at Colonia, Exu municipality
(7.5741°S, 39.7525°W, SAD 69; 456 m elev.), Pernambuco, north-
eastern Brazil. The animal was euthanized with 2% lidocaine hy-
drochloride. The gastrointestinal tract was removed for desicca-
tion and analysis of parasites using a stereomicroscope.
We found 137 endoparasites identified as Physaloptera sp., in
larval stage in the large intestine of A. vermicularis. Previously,
A. vermicularis had a record of infection by a single species of
Pentastomida (Raillietiella gigliolli, prevalence of 55.5%) (Al-
meida, et al., 2008. Braz. J. Biol. 69:1183–1186). Nematodes of
the genus Physaloptera Rudolphi, 1819 have been recorded in
amphibians, birds, mammals, and reptiles (Pereira et al. 2012. J.
Parasitol. 98:1227–1235). Physaloptera spp. have been reported
parasitizing the following lizards, Ameiva ameiva, Cercosaura
argulus, Cnemidophorus litoralis, C. ocellifer, Hemidactylus
mabouia, Hoplocercus spinosus, Mabuya agilis, M. macrorhyn-
cha, Polychrus acutirostris, Tropidurus etheridgei, T. torquatus,
Tupinambis merianae, Tupinambis teguixin, and Dicrodon gut-
tulatum (Ávila and Silva 2010. J. Venomous Animals and Tox-
ins including Tropical Diseases 16:543–572). The intermediate
hosts of Physaloptera are arthropods such as crickets, locusts,
cockroaches, and beetles (Gray and Anderson 1982. Can. J. Zool.
60:2134–2142). To our knowledge, this is the first record of Physa-
loptera sp. infecting A. vermicularis.
ÉRICA GOMES DA SILVA (e-mail: ericagomesdasilva127@gmail.
OLIVEIRA, Laboratório de Zoologia/Parasitologia, Universidade Regional
do Cariri –URCA, Campus Pimenta, CEP 63100-000, Crato, Ceará, Brazil;
de Pós-Graduação em Ciências Biológicas (Zoologia), Laboratório/Coleção
de Herpetologia, Universidade Federal da Paraíba – UFPB , Cidade Univer-
sitária, Campus I, CEP 58059-900, João Pessoa, Paraíba, Brazil; ANTONIO
tamento de Química Biológica, Campus Pimenta, Universidade Regional
do Cariri – URCA, Rua Cel. Antônio Luiz, 1161, CEP 63105-100, Crato, CE,
is a recently described species of the Ameivula ocellifera group,
found in the Caatinga lowlands of Bahia state, Brazil (Arias et
al. 2011. Zootaxa 3022:1–21). There is little known of the natural
history of this lizard, and nothing known of its diet. Consider-
ing other species of the A. ocellifera group, the diet is composed
mainly of arthropods, with one case of cannibalism reported
(Sales et al. 2010. Herpetol. Rev. 41:217–218). Here, we report the
first recorded case of cannibalism for A. nigrigula from the Caat-
inga of northeast Bahia.
On 5 March 2015, we captured a male A. nigrigula (SVL = 63.9
mm, mass = 9 g before regurgitation) in a semi-arid area of the
Lipari Mineração LTDA near Nordestina in the state of Bahia,
Brazil (10.9026°S, 39.4227°W; SAD69). In this specimen’s stomach
we found an undigested juvenile A. nigrigula (SVL = 33.8 mm,
mass = 1 g) (Fig. 1).
Our survey was part of the fauna rescue program of Lipari
Mineração LTDA (license INEMA nº 8598). The specimen was
deposited in the herpetological collection of the Centro de Eco-
logia e Conservação Animal, Universidade Católica do Salvador,
Salvador, Bahia. (CHECOA 3657).
erta Consultoria, Projetos & Assessoria Ambiental LTDA, Av Tancredo Neves,
939 - s-1305, Caminho Árvores, Salvador, Bahia , Brazil; Centro de Ecologia e
Conservação Animal, Universidade Católica do Salvador, Av. Prof. Pinto de
Aguiar, 2589, 41740-090, Pituaçu, Salvador, Bahia, Brazil.
Saurophagy by other lizards is considered common in semi-
arid environments such as in Australia (Pianka 1973. Annu. Rev.
Ecol. Syst. 4:53–74), and may also be common in the semi-arid
Caatinga of northeastern Brazil (Vitt 1995. Occas. Pap. Oklahoma
Mus. Nat. Hist. 1:1–29). Ameivula is a teiid genus that includes
species widely distributed from northeastern Brazil to northern
Fig. 1. Cannibalism in Ameivula nigrigula: A) stomach being remo-
ved; B) predator and prey.
Herpetological Review 48(3), 2017
Argentina (Arias et al 2011. Zootaxa 2787:37–54; Arias et al 2011.
Zootaxa 3022:1–21). Ameivula ocellifera is a terrestrial species
that actively forages during the day (Mosque and Colli 2003. J.
Herpetol 37:498–509). Its diet is generalized, mainly composed
of adult insects and larvae (Sales and Freire 2015. J. Herpetol.
49: 579–585); however, adults can be cannibalistic on juveniles
(Sales et al 2010. Herpetol. Rev. 41:217–218). Tropidurus hispidus
(Peter’s Lava Lizard, family Tropiduridae) is a habitat and dietary
generalist distributed throughout South America. It is common
in Caatinga regions where it is easily observed in rocky outcrops
(Kolodiuk et al. 2010. South Am. J. Herpetol. 5:35–44). Herein, we
report an observation of an A. ocellifera preying upon Tropidurus
Our observation occurred during a survey of the herpeto-
fauna in the Caatinga, on a rocky outcrop at Sítio Cacimbas,
municipality of Itapetim, state of Pernambuco, Brazil (7.4053°S,
37.1868°W, WGS 84; elev. 647 m). At 1437 h on 28 July 2015, we
observed an adult A. ocellifera (SVL ~ 70 mm) subduing a juvenile
T. hispidus (SVL ~ 31 mm) on the ground near a cluster brome-
liad (Encholirium spectabile). At that time the A. ocellifera held
the T. hispidus firmly within its mouth, with only the head and
forelimbs exposed (Fig. 1). The predator then moved approxi-
mately 0.5 m towards the adjacent herbaceous vegetation, where
it continued to try and swallow its prey. The A. ocellifera moved a
second time into vegetation (approximately 0.3 m) and, when it
stopped, completely swallowed its prey. The total duration of the
predation event was approximately one minute.
Teiid lizards are active foragers, and they capture mainly
sedentary prey using a well-developed visual and chemorecep-
tion system (Huey and Pianka 1981. Ecology 62: 991–999). Both
A. ocellifera and T. hispidus share dimensions of the ecological
niche including activity period and various food items, and they
use similar habitats (Andrade et al 2013. Biota Neotrop. 13:199–
209). This can result in increased competition, although there are
segregation mechanisms that can enable the co-occurrence of
species (Huey and Pianka 1977. Ecology 58:119–128). In our case,
the predation event might be explained by 1) the fact that a large
prey item like a lizard is a high-energy food item that can reduce
the need to feed for several days (Vitt 2000. Herpetol. Monogr.
14:388–400); and/or 2) the elimination of potential competitors.
We are grateful to Coordenação de Aperfeiçoamento de Pes-
soal de Nível Superior (CAPES) for financial support, and to Sítio
Cacimbas owner for assistance during fieldwork.
SOUSA (e-mail: í, ÂNDERSON B. P. ARAÚJO (e-
mail:, and MARCELO N. C. KOKUBUM,
Laboratório de Herpetologia and Programa de Pós-graduação em Ciên-
cias Florestais, Centro de Saúde e Tecnologia Rural, Universidade Federal
de Campina Grande, Caixa Postal 61, Santa Cecília, CEP 58708-110, Patos,
Paraíba, Brazil (e-mail:
and ANT INVASION. Some of the highland gymnophthalmid
lizard species share the behavior of communal nesting (Swain
et al. 1980. J. Herpetol. 14:321–326; Ramos-Pallares et al. 2013.
Herpetol. Rev. 44:226–229). Anadia bogotensis is a Colombian
endemic that has been recently designated as Vulnerable in
the national threatened reptile species list. This species is
likely to be threatened by cattle grazing expansion, agriculture,
fire, urbanization, and unsustainable ecotourism (Morales-
Betancourt et al. 2015. Libro Rojo de Reptiles de Colombia.
Instituto de Investigación de Recursos Biológicos Alexander
von Humboldt, Universidad de Antioquia, Colombia. 258
pp.). Records of communal nesting in A. bogotensis include
the descriptions of nests composed of 26–578 eggs in different
development stages, and the presence of old eggshells next to
new eggs demonstrated that nests are continuously used. The
number of eggs laid under the same rock during a breeding
season varied from 2–60; these eggs incubate for 6–7 months.
Communal nests from this species are typically found under
flat rocks and under low necromass on highlands (paramos)
near Bogotá reaching elevations of up to 3900 m. Nesting in
this species has been observed from July to November 2006 and
from March to May 2009 (Medina-Rangel. 2013. Herpetol. Rev.
44:312–313). Here we further describe communal nesting in A.
bogotensis, and the subsequent invasion of ants into those nests.
During 2012 and 2015 we monitored seven A. bogotensis
nests located in the eastern mountains of Bogotá near La
Calera (4.6827139°N, 74.0123167°W, WGS84; 3109 m elev.).
Communal nests were observed from May to November (Fig.
1). During monitoring in 2014, we noticed that Carpenter Ants
(Camponotus cf. nitens) had built colonies on six of the nests
initially utilized by the lizards (Fig. 2). Ants persisted throughout
2014 and 2015, suggesting that once ant colonies are established
they will prevail for more than two years under the same rock.
Since these six nests were not used by lizards again (Fig. 2), it
Fig. 1. Adult Ameivula ocellifera preying on a juvenile Tropidurus his-
pidus on a rocky outcrop.
Herpetological Review 48(3), 2017
is likely that the ants are interfering with nesting behavior and
hatchling development by driving adults away, or consuming
eggs or hatchlings. Communal nesting can result from limited
nest sites (Doody et al. 2009. Quart. Rev. Biol. 84:229–251); thus, in
paramo habitats, humidity and temperature gradients may limit
the availability of places suitable for egg laying and successful
incubation. Moreover, communal nesting has been suggested to
increas fitness in hatchlings of other lizard species (Radder et al.
2007. J. Anim. Ecol. 76:881– 887).
Climate change has been identified as one of the drivers that
have caused ecological shifts in reproductive phenology and
high extinction rates of native species, caused by abrupt chang-
es in distribution (Parmesan et al. 2003. Nature 421:37–42). Ant
populations have shown different responses to global warming
such as increasing their altitudinal range and abundance (Kwon
et al. 2015. J. Asia-Pacific Biodivers. 8:49–65). However, studies
on high altitude ants in the neotropical region are scarce, and the
dynamics of their range and population trends influenced by cli-
mate change remain unknown. We note that prior to 2010, nests
without ants were common. The invasion by these ants suggests
that Camponotus cf. nitens are expanding to higher elevations
(showing an increase in the number of colonies), a trend that
may threaten Andean lizards that lay communally. Nest displace-
ment by ant colonization is thus an environmental stressor for A.
bogotensis that could cause significant behavioral changes in this
species, affecting survival and recruitment. Finally, we suggest
further research on this interaction since it might be an emergent
threat brought by climate change, affecting this vulnerable and
endemic high altitude lizard.
JUAN SALVADOR MENDOZA R., Universidad de los Andes, Bogotá,
Colombia (e-mail:; CAMILA A. RODRI-
GUEZ-BARBOSA, University of Florida (e-mail: camila.rodriguez@u.edu)
ANNIELLA GRINNELLI (Bakersfield Legless Lizard). PREDA-
TION. Anniella grinnelli is a recently described species of angui-
morph lizard known only from Kern and San Luis Obispo counties
in south-central California, USA (Papenfuss and Parham 2013.
Breviora 536:1–17). Due to the secretive nature and recent recog-
nition of this species, information on the predators of A. grinnelli
is absent from the literature. On 26 March 2017 we conducted
surveys for Anniella at the Pixley National Wildlife Refuge in Tu-
lare County, California, USA. During these surveys, we encoun-
tered a dead adult A. grinnelli (Fig. 1) which had been impaled by
Lanius ludovicianus (Loggerhead Shrike) on a barbed wire fence
at the edge of the property (35.90635°N, 119.33169°W; WGS 84).
The specimen was collected and deposited in the Museum of
Vertebrate Zoology, University of California, Berkeley, California,
USA (MVZ 272791) along with a Uta stansburiana (MVZ 272790)
found similarly impaled on the same stretch of fence. Lanius
ludovicianus is a known predator of reptiles including A. pulchra,
a more widely distributed relative of A. grinnelli (Clark 2011. Son.
Herpetol. 24:20–22). However, to our knowledge this observation
represents the first confirmed predator of A. grinnelli in the lit-
erature. Additionally, this specimen represents the first record of
A. grinnelli in Tulare County, CA (Papenfuss and Parham, op. cit.).
This research was conducted under a California Scientific
Collection Permit issued to T. J. Papenfuss. The work was suppor-
ted by the Museum of Vertebrate Zoology, University of Califor-
nia, Berkeley and a grant from the California Department of Fish
and Wildlife.
MARK W. HERR, Department of Herpetology, University of Kansas
Natural History Museum, 1345 Jayhawk Blvd., Lawrence, Kansas 66045,
of Vertebrate Zoology and Department of Integrative Biology, 3101 VLSB,
University of California Berkeley, California 94720-3160, USA.
ANOLIS CAROLINENSIS (Green Anole). DIET. Anolis carolinen-
sis is native to the coastal plain of the southeastern United States
and is considered a dietary generalist, feeding on a variety of in-
sects, spiders, and other arthropods (Conant and Collins 1998.
Reptiles and Amphibians of Eastern and Central North America.
Fig. 1. A communal nest site of Anadia bogotensis, showing eggs and
hatched eggshells.
Fig. 2. A former Anadia bogotensis communal nest site, now colo-
nized by carpenter ants.
Herpetological Review 48(3), 2017
3rd Ed. Houghton Mifflin, New York. 616 pp.). As generalists, their
diet is apparently only restricted by size. Herein, we report on a
rare observation of A. carolinensis feeding on a vertebrate. Our
observation also represents what we believe to be the first record
of an A. carolinensis regurgitating another lizard.
On 13 October 2016, between 1200–1215 h, an adult female
A. carolinensis (SVL = 49 mm, total length = 139 mm, weight =
2.1 g) was detected inside an upright PVC pipe set up as part of a
sampling array for a long-term study of amphibian use of upland
Longleaf Pine ephemeral sinkhole ponds in the Ocala National
Forest (29.32391°N, 81.74823°W). Skies were sunny and the tem-
perature was approximately 27°C. While weighing this individu-
al, it regurgitated about half of an adult Scincella lateralis (Little
Brown Skink), and, after a few minutes, the complete body. With
a body girth (~6–8 mm) more than a quarter that of its predator
and a total length (~80 mm) over half that of its predator, the S.
lateralis was an exceptionally large prey item for A. carolinensis.
The S. lateralis had not yet lost any coloration, although some
exterior scales were partially digested. We have captured dozens
of A. carolinensis during each of the last 24 years in the field, but
this was the first time that we witnessed one regurgitating prey
upon capture, suggesting that prey size and (or) species was a
contributing factor to this behavior. This observation occurred
under Florida Fish and Wildlife Conservation Commission per-
mit # LSSC–12-00021A.
SKY T. BUTTON, Department of Wildlife Ecology and Conservation,
University of Florida, Gainesville, Florida 32611, USA (e-mail: skybutton@
u.edu); CATHRYN H. GREENBERG, USDA Forest Service, Upland Hard-
wood Ecology and Management RWU-4157, 1577 Brevard Rd., Asheville,
North Carolina 28806, USA (e-mail:; JAMES D. AUS-
TIN, Department of Wildlife Ecology and Conservation, University of Flori-
da, Gainesville, Florida 32611, USA (e-mail: austinj@u.edu).
Anolis equestris is native to Cuba and established in southern
Florida (Meshaka 2011. Herpetol. Conserv. Biol. 6:1–101). Regen-
eration of multiple tails is thought to be due to tail injury without
tail breakage (Bellairs and Bryant 1985. In Gans and Billett [eds.],
Biology of the Reptilia, Volume 15, Development B, pp. 301–410.
Wiley-Interscience, New York). Tail trifurcation has been report-
ed in the lizard genera Ctenosaura (Iguandiade; Ariano-Sanchez
and Gil-Escobedo 2016. Herpetol. Rev. 47:463–464), Cyclura
(Iguanidae; Hayes et al. 2012. Biodivers. Conserv. 21:1893–1899),
Lacerta (Lacertidae; Graper 1909 In Bellairs and Bryant, op. cit.),
Hemidactylus (Gekkonidae; Evans and Bellairs 1983 In Bellairs
and Bryant, op. cit.), Gekko (Gekkonidae; Das 1933 In Bellairs and
Bryant, op. cit.), and Mabuya (Scincidae; Brindley 1898 in Bellairs
and Bryant, op. cit.).
At 2045 h, 22 May 2016 we found an adult male A. equestris
(Fig. 1; 157 mm SVL, 175 mm tail length, 80 g) sleeping on a thin
branch ~3 m aboveground in second growth forest adjacent to an
avocado orchard (25.425483°N, 80.501383°W, WGS 84; 22 m elev.)
southwest of Florida City, Miami-Dade County, Florida, USA. The
trifurcation (Fig. 1) was 113 mm posterior to the cloaca and con-
sisted of a split on the ventral side of the tail resulting in two new-
ly regenerated tail tips, a left prong (27.2 mm) and right prong
(36.4 mm) that projected from beneath the central axis of the tail
which extended 61.7 mm beyond the split. The right prong was
the same color as the proximal part of the tail and may have rep-
resented the original portion of the tail. To our knowledge this is
the first report of tail trifurcation in this species.
BENJAMIN T. CAMPER, 723 Sidney Ave., Florence, South Carolina
29505, USA (e-mail:; JEFFREY D. CAMPER, De-
partment of Biology, Francis Marion University, Florence, South Carolina
29506, USA.
ANOLIS EQUESTRIS (Cuban Knight Anole). DIET. Anolis eques-
tris is native to Cuba and present as a nonindigenous species
Fig. 1. Anniella grinnelli (MVZ 272791) impaled on a barbed wire
fence by Lanius ludovicianus.
Fig. 1. Adult male Anolis equestris with a trifurcated tail. Inset show-
ing that the central axis of the tail is yellow.
Herpetological Review 48(3), 2017
in Florida, California, Hawaii, central Mexico, and coastal Bra-
zil. Previous studies have indicated the diet of A. equestris to
be principally insectivorous and to a lesser degree frugivorous
(Brach 1976. Copeia 1976:187–189; Dalrymple 1980. J. Herpe-
tol. 14.4:412–415). Individuals within Florida will also consume
smaller congeners, A. sagrei and A. porcatus (Meshaka et al. 2004.
Exotic Amphibians and Reptiles of Florida. Krieger Publishing
Company, Malabar, Florida. 61 pp.). While cannibalism is well
documented in some other species of anoles, the possible oc-
currence, prevalence, and age classes involved in cannibalism
among A. equestris are unknown. At 2214 h on 10 March 2015,
while surveying for exotic species in Homestead, Florida, USA
(25.447488°N, 80.533897°W; WGS 84) we captured by hand a
juvenile A. equestris (SVL = 7.35 cm). A prey item was readily vis-
ible protruding from the animal’s throat as it gaped defensively
(Fig. 1A). The item was gently removed and identified as a small-
er juvenile A. equestris (SVL = 5.2 cm). While the posterior end
of the prey item was intact, the anterior had been digested. This
exposed a skull and clear dentition with blunt, conical teeth (Fig.
1B,C). The cannibalizing anole was in good body condition and
not presenting any obvious negative effects from the large size
of this prey item. This observation demonstrates cannibalism
among the A. equestris juvenile age class in the introduced range
of this species and demonstrates the selection by an A. equestris
juvenile of an extraordinarily large prey item relative to current
dietary accounts.
HOLLIS DAHN, University of Toronto, 27 King’s College Cir, Toronto,
Ontario M5S, Canada (e-mail:; JEFF SHARPE,
University of Central Florida, 4000 Central Florida Blvd, Orlando, Florida
32816, USA.
fuscoauratus is an iguanid (Polychrotinae) lizard that is widely
distributed in northern South America east of the Andes and in
the Atlantic Forest in eastern Brazil. This species is commonly
found in dense forests and exhibits predominantly arboreal
Fig. 1. Juvenile Anolis equestris captured in Homestead, Florida. A)
Animal gaping defensively after capture. B) Animal and prey item
size comparison. C) Anterior end of prey item showing dentition
consistent with A. equestris.
Fig. 1. Whipscorpion (Heterophrynus longicornis) preying on Anolis
Herpetological Review 48(3), 2017
habits. It is usually found on trunks, vines, and leaves, generally
within a few meters of the ground (Vitt et al. 2008. Guide to the
Frogs of Reserva Adolpho Ducke, Central Amazonia. Áttema De-
sign Editorial, Manaus. 176 pp.). Here we report on observation
of predation on A. fuscoauratus by a whipscorpion.
At 0215 h on 1 April 1 2012, a specimen of Heterophrynus
longicornis (Amblypygi: Phrynidae) was observed preying on an
adult A. fuscoauratus (Fig. 1) in an ombrophilous forest in the
Amazonian Federal University (UFAM) Experimental Farm, lo-
cated in Manaus city, Amazonas, Brazil (2.65885°S, 60.06603°W;
WGS 84). The predation event occurred on a tree trunk approxi-
mately 1.6 m above ground. The lizard was a male (SVL = 44 mm,
TL = 70 mm). The whipscorpion was holding the lizard by the
head and thoracic region with its pedipalps (Fig. 1). Ingestion
had begun on the lateral region of the lizard’s head, and a large
part of the head and neck there were already partially digested,
probably due to the action of the whipscorpion’s digestive en-
Our record corroborates the hypothesis that whipscorpions
remain oriented downward, with their pedipalps raised and out-
stretched, waiting for prey (Dias and Machado 2007. J. Arach-
nol. 34:540–544). When potential prey enters the field of view, it
is caught and immobilized by the raptorial pedipalps and then
consumed at the site in which it was captured. Our observation is
the first record of Heterophrynus preying upon a vertebrate, and
increases the list of invertebrates that can prey on A. fuscoauratus.
DIEGO M. M. MENDES, Laboratório de Entomologia Sistemática Ur-
bana e Forense, Instituto Nacional de Pesquisas da Amazônia - Campus II,
Av. André Araújo, 2936, 69080-971 Manaus, AM, Brazil (e-mail: diego.mello.; ANDRÉ L. BARROS, Departamento de Ecologia, In-
stituto Nacional de Pesquisas da Amazônia – Campus V8, Av. Egênio Sales,
2239, 69060-020, Manaus, Amazonas, Brazil (e-mail: andrelima1701@gmail.
com); DIEGO A. PIRES, Departamento de Ecologia, Instituto Nacional de
Pesquisas da Amazônia – Campus V8, Av. Egênio Sales, 2239, 69060-020,
Manaus, AM, Brazil (e-mail:; PATRIK F.
VIANA, Laboratório de Genética Animal, Instituto Nacional de Pesquisas
da Amazônia - Campus II, Av. André Araújo, 2936, 69080-971 Manaus, Ama-
zonas, Brazil (e-mail:
Sleep-site choice can directly influence survival in many spe-
cies due to their vulnerability to predation (Clark and Gillingham
1990. Anim. Behav. 39:1138–1148). Like many diurnal lizards,
Anolis spp. are commonly observed sleeping at night on fern
fronds and small tree branches, sites that would appear to ex-
pose them to the risk of predation (Singhal et al. 2007. Behaviour
144:1033–1052). To begin to understand why particular sleep-
sites are chosen, we assessed sleep-site choice and fidelity in
Anolis gemmosus in the Andes Mountains of Ecuador.
We conducted our observations at Reserva Las Gralarias
(RLG, Province of Pichincha) (2068 m elev.) and made repeated
nightly observations on 20 anoles at 13 sleep-sites along two
open-canopy trails (each trail approximately 250 m long). Aver-
age temperature during the sampling period was 19.4°C (± 0.05°C
SE) with intermittent light rainfall. We sampled Road Trail, a ru-
ral roadside with limited automobile disturbance, from 2–7 May
2016, and Granny’s Trail, a walking trail with limited human dis-
turbance (foot traffic only), from 3–7 May 2016. We walked the
trails each night during 2000–2300 h to observe anole sleeping
behavior. To limit human interference, we did not mark indi-
vidual lizards, but rather used photographs of each anole for
individual animal identification. The unique skin coloration of
individuals allowed us to recognize previously sighted anoles.
Our results confirmed sleeping site fidelity in A. gemmosus
with 17/20 anoles observed re-sighted at least once within 1 m
of their initial sleeping site (mean ± SE 1.68 ± 0.30 re-sightings). A
Wilcoxon rank sign test indicated that our two trails did not sig-
nificantly differ in their number of re-sighted anoles (W = 79.50,
p = 0.16).
During our survey, we found anoles utilizing several different
types of sleeping perches, typically in an oblique position respec-
tive to the ground. Sleeping perches consisted of herbaceous
plants, fern fronds, and thin branches of woody plants at heights
ranging 1.6–4 m. Unexpectedly, seven of our sites contained
multiple (2 –3) anoles within a 5-m radius, which suggests there
may be competition for sleeping sites, or reduction in individual
predation risk via grouping behavior. Our results indicating that
A. gemmosus exhibit sleep-site fidelity at both disturbed and un-
disturbed sites provides novel natural history information for the
species as we are unaware of any other studies examining this
behavior in the species.
Future investigations into A. gemmosus behavior should ex-
amine whether this species exhibits intraspecific competition
for sleeping sites, and what habitat characteristics (such as perch
diameter, elevation above land, anole density, etc.) may predict
such behavior. If competition for sleeping sites does occur, this
may influence an individual’s fidelity to a particular site.
We are grateful to RLG volunteer, Ray So Lok, and the stu-
dents of the Grand Valley State University study abroad program
for field assistance, and to Jane Lyons and Reserva Las Gralarias,
for hosting our research.
ALEXA S. WAGNER, Department of Biology, Kent State University,
Kent, Ohio 44240, USA (e-mail:; ALEXANDRIA L. LA-
VALLEY, Department of Biology, Grand Valley University, Allendale, Michi-
gan 49401, USA (e-mail:; KATHERINE L. KRYNAK,
Department of Biological and Allied Health Sciences, Ohio Northern Uni-
versity, Ada, Ohio 45810, USA (e-mail:; TIM J. KRYNAK,
Natural Resources Department, Cleveland Metroparks, Cleveland, Ohio
44109, USA (e-mail:; ERIC B. SNYDER, De-
partment of Biology, Grand Valley University, Allendale, Michigan 49401,
USA (e-mail:
Anolis gundlachi is an endemic Puerto Rican anole, classified as
a forest-interior “trunk-ground” ecomorph (sensu Williams 1972.
In Dobzhansky et al. [eds.], Evolutionary Biology, Volume 6, pp.
47–89. Meredith Corporation, New York, New York; Reagan 1996.
In Reagan and Waide [eds.], The Food Web of a Tropical Rain For-
est, pp. 321–345. University of Chicago Press, Chicago, Illinois). It
is distributed throughout montane areas in Puerto Rico and has
an adult mean snout–vent length range of 42–72 mm, (Reagan
1996, op. cit.). It has a generalist diet that includes a wide vari-
ety of invertebrates (mostly associated with the forest litter) and
vertebrate prey that includes treefrogs (Eleutherodactylus sp., E.
wightmanae) and other lizards (A. stratulus, A. krugi, Sphaero-
dactylus klauberi) (reviewed in Henderson and Powell 2009. Nat-
ural History of West Indian Reptiles and Amphibians. University
Press of Florida. Gainesville, Florida. 528 pp.; Ríos-López et al.
2014. LEB 3:137–148). Cannibalism has been reported in 19 West
Indian anoles, including A. evermanni and A. cristatellus from
Puerto Rico (reviewed in Powell and Watkins 2014. IRCF Reptiles
and Amphibians 21:136–137). Herein we document the first ac-
count of cannibalism in A. gundlachi, an event observed as part
of a long-term survey of malarial parasites in anoles.
Herpetological Review 48(3), 2017
On 13 January 2017, in El Verde Field Station, Luquillo Ex-
perimental Forest in northeastern Puerto Rico (18.3213°N,
65.8194°W, WGS 84; 357.9 m elev.), we observed and collected a
male A. gundlachi (mass = 7.2 g; SVL = 65 mm) that consumed
a juvenile A. gundlachi (mass = 1.0 g; SVL 35 mm), head first.
Three quarters of the tail of the prey was visible protruding from
the mouth of the predator; it regurgitated the prey when we ap-
proached. Both individuals were identified as A. gundlachi by a
combination of characters: blue-eye coloration and the distinc-
tive yellow-colored chin. This is, to our knowledge, the first re-
port of cannibalism by A. gundlachi.
Cannibalism in reptiles is believed to be an opportunistic re-
sponse to high conspecific densities or starvation (Polis and My-
ers 1985. J. Herpetol. 19:99–107). At El Verde, population density
of A. gundlachi reaches 2000 individuals ha-1, with relative abun-
dance fluctuation from a mean 86.5% (± 2.6% SD, N = 4 census
transects) sighting at ground level during the wet season (May to
end of year) to a mean 64.0% (± 7.0% SD, N = 4 census transects)
sighting during the dry season (January to April) (Reagan 1996,
op. cit.). Abundance of invertebrate prey available to A. gund-
lachi also varies seasonally, increasing during the wet season
and decreasing during the dry season (reviewed in Reagan and
Waide 1996. The Food Web of a Tropical Rain Forest. University
of Chicago Press, Chicago, Illinois. 616 pp.). Although less food
resources due to the winter dry season and the generalist diet of
A. gundlachi may be hypotheses that explain our observation,
we cannot discard the possibility that this may have been an un-
common behavior in the species.
mail:, NICOLE MARTÍNEZ-LLAURADOR (e-mail:, and MIGUEL A. ACEVEDO, University of
Puerto Rico - Río Piedras, P.O. Box 23360, 00936, San Juan, Puerto Rico (e-
DIET. At 0054 h on 14 December 2015, while conducting
fieldwork in Cozumel Island, Quintana Roo, Mexico, we collected
an adult male A. georgeensis at ca. 0054 h near Puerto de Abrigo,
a marina on the northwest coast of the island (20.52878°N,
86.93867°W, WGS 84; 7 m elev.). The gecko was located on a
chacah tree, Bursera simaruba, in an unkept botanical garden
with both introduced and local flora (Téllez Valdez et al. 1989.
Las Plantas de Cozumel [Guía Botánico-Turística de la Isla de
Cozumel, Quintana Roo]. Instituto de Biología, Universidad
Nacional Autónoma de México, México, D.F. 75 pp.). The
specimen’s stomach contents were later removed and identified
by Rachael Alfaro and Kelly B. Miller to the order Scorpiones (Fig.
1). The gecko was deposited in the Museo de Zoologia “Alfonso L.
Herrera,” Facultad de Ciencias, Universidad Nacional Autónoma
de México, México (MZFC-HE 30638).
Geckos of the genus Aristelliger are known to have a broad
diet that includes a variety of arthropods, hatchling geckos and
eggs, Anolis lizards, berries, and flowers (Cloud 2013. Cryptic
Diversity, Evolution, and Biogeography of Caribbean Croaking
Geckos (Genus: Aristelliger). Master of Science thesis, The Pen-
nsylvania State University, University Park, Pennsylvania. 44 pp.).
The diet of Aristelliger includes Arachnida, with previous studies
indicating the orders Araneae and Pseudoscorpiones, but reports
of the order Scorpiones are absent from the literature (Gifford et
al. 2000. Caribb. J. Sci. 36:3–4). Ours is the first record of preda-
tion on Scorpiones by A. georgeensis.
Fieldwork was conducted under the authority of collecting
permit FAUT 0243 issued to Uri O. García-Vázquez by the Secre-
taría de Medio Ambiente y Recursos Naturales.
BRITTNEY A. WHITE (e-mail: and LEVI N.
GRAY, Department of Biology and Museum of Southwestern Biology, Uni-
versity of New Mexico, Albuquerque, New Mexico 87131, USA; CARLOS
J. PAVÓN-VÁZQUEZ*, Laboratorio de Herpetología, Museo de Zoología,
Departamento de Biología Evolutiva, Facultad de Ciencias, Universidad Na-
cional Autónoma de México, Apartado Postal 70-153, México 04510, D.F.,
México; URI O. GARCÍA-VÁZQUEZ, Carrera de Biología, Facultad de Estu-
dios Superiores Zaragoza, Universidad Nacional Autónoma de México, Ba-
talla 5 de mayo s/n, Ejército de Oriente, México 09230, D.F., México; ALEXIS
S. HARRISON, Department of Organismic and Evolutionary Biology, Har-
vard University, Cambridge, Massachusetts 01238, USA. *Current address:
Division of Ecology and Evolution, Research School of Biology, Australian
National University, Acton, ACT 2601, Australia.
ASPIDOSCELIS INORNATA (Trans-Pecos Striped Whiptail).
PREDATION. Aspidoscelis inornata is a diurnal, active foraging
teiid lizard native to the southwestern United States and north
central Mexico (Jones and Lovich 2009. Lizards of the American
Southwest: A Photographic Field Guide. Rio Nuevo Publishing,
Tucson, Arizona. 567 pp.). Until recently, information on the
predators of A. inornata was absent from the literature. However,
in recent years we have documented predation on this species by
Fig. 1. Stomach contents of Aristelliger georgeensis (MZFC-HE 30638).
Herpetological Review 48(3), 2017
the Coachwhip (Coluber flagellum; Graham and Kelehear 2015.
Herpetol. Rev. 46:267), Western Massasauga (Sistrurus tergemi-
nus; Graham and Kelehear 2015. Herpetol. Rev. 46:107), and Log-
gerhead Shrike (Larius ludovicianus; Graham and Kelehear 2016.
Herpetol. Rev. 47:132). Here we contribute an additional obser-
vation of an avian predator of A. inornata.
On 28 May 2017, in Brewster County, Texas, USA (30.69633°N,
103.21686°W; WGS 84), we observed a Greater Roadrunner (Geo-
coccyx californianus) perched in a shrub feeding upon an indidi-
vual A. inornata (Fig. 1). Close inspection of the interaction with
10x40 binoculars at 35 m confirmed the identity of the lizard. To
our knowledge, this is the first documented instance of G. cali-
fornianus preying upon A. inornata (Clark 2011. Son. Herpetol.
MARK W. HERR, Department of Herpetology, University of Kansas
Natural History Museum, 1345 Jayhawk Blvd., Lawrence, Kansas 66045,
USA (e-mail:; LAINE A. GIOVANETTO, Biology De-
partment, New Jersey City University, Jersey City, New Jersey 07305, USA;
SEAN P. GRAHAM, Department of Biology, Geology, and Physical Sci-
ences, Sul Ross State University, Alpine, Texas 79830, USA (e-mail: sean.
notable lizard capture techniques are utilized by ecologists when
attempting to collect these animals alive in the field (Fitzgerald
2012. In McDiarmid et al. [eds.], Reptilian Biodiversity: Standard
Methods for Inventory and Monitoring, pp. 77–88. University of
California Press, Berkeley, California; Schemnitz et al. In Silvy
[ed.] The Wildlife Techniques Manual – Research. Vol. 1, 7th ed. pp.
64–117. Johns Hopkins University Press, Baltimore, Maryland).
Other than direct hand grabbing, possibly the most common and
most fundamental methods used to collect live lizards are with
funnel and pitfall traps and with the use of a noose. A less well-
known technique that can be especially effective for securing
live, unharmed whiptail lizards of the genus Aspidoscelis (family
Teiidae) involves removing them from their temporary dens
(i.e., activity burrows) by finger probing their burrow system in
conjunction with tool excavation (Trauth 1977. Herpetol. Rev.
8:33). In the following, I provide details regarding a modification
of the finger probing/tool excavation method specifically
designed to easily capture these fast and elusive lizards alive.
During the summer of 1982, I designed a simple capture de-
vice that proved useful in collecting Aspidoscelis sexlineata sex-
lineata in sandy, flat-ground, disturbed (open area and sparsely
vegetated) habitats. This method requires some basic knowledge
about the biology of this species. For example, one must be able
to recognize freshly excavated activity burrows with their char-
acteristic open mouths (Fig. 1), which are normally conspicuous
on the ground surface. Activity burrows constructed by whiptail
Fig. 1. Aspidoscelis inornata preyed upon by Geococcyx californian-
Fig. 1. Capture device for securing live Aspidoscelis sexlineata sexlin-
eata. A) Device set into the ground on 11 July 1982 in a vacant lot
located in the Temple Terrace subdivision of Tampa, Hillsborough
County, Florida, USA (28.055582°N, 82.312866°W, WGS 84; 25 m
elev.) and approximately 4 km SE of the University of South Florida
campus. B) Trap placement over an activity burrow (end of arrow). C)
The captured lizard (end of arrow) measured 65 mm in snout–vent
Herpetological Review 48(3), 2017
lizards have been described elsewhere for A. gularis (Trauth
1987. Southwest Nat. 32:279–281), A. laredoensis (Walker et al.
1986. Southwest. Nat. 31:408–410), and A. sexlineata (Trauth et
al. 2013. Herpetol. Rev. 44:668–669).
The capture device is relatively inexpensive and easily carried
by a researcher in the field. It consists of a piece of sturdy, but
flexible, aluminum flashing (approximately 2.0 m × 0.3 m) and a
coupling “C” clamp (Fig. 1A, B). The trapping technique is as fol-
lows: 1) a freshly constructed activity burrow of the lizard is iden-
tified by its sandy tailings strewn about in a fan-shaped pattern
in a direction away from the burrow mouth (Fig. 1B). This dis-
turbance normally indicates the presence of the lizard within an
individual burrow or within passageways radiating out from the
primary burrow. 2) The aluminum flashing is curled into a circle
with its ends firmly attached together with the clamp. The “cir-
cle” trap is then placed over the burrow site and firmly embed-
ded into the sandy soil (Fig. 1B). 3) A short-handled garden tool,
preferably a four-prong rake is gently plunged into the soil near
the entrance to the burrow. Successive digging a swath through
the soil in the vicinity of the burrow mouth normally induces the
hidden lizard into an escape behavior, at which time the animal
explodes out the distal end of its burrow shelter and runs into
the wall of the enclosure (Fig. 1C). I have successfully used this
device on numerous occasions in flat, sandy habitats in Florida
and in similar habitats throughout this species’ range.
STANLEY E. TRAUTH, Department of Biological Sciences, Arkan-
sas State University, P.O. Box 599, State University, Arkansas, USA; e-mail:
ASPIDOSCELIS UNIPARENS (Desert Grassland Whiptail).
REPRODUCTION. Triploid parthenogenetic Aspidoscelis
uniparens is a hybrid-derived species (Reeder et al. 2002. Amer.
Mus. Novitat. 3365:1–61) distributed through parts of Arizona,
New Mexico, and Texas in the USA and states of Chihuahua
and Sonora in Mexico (Wright 1968. J. Herpetol. 1:1–20; Lemos-
Espinal and Smith 2007. Anfibios y Reptiles del Estado de
Chihuahua, México/Amphibians and Reptiles of the State of
Chihuahua, Mexico. UNAM-CONABIO. México, D.F. 613 pp.).
We assessed size and reproductive characteristics in the species
near the southern limits of its range in Mexico based on 54
specimens from a site where it can be readily observed and easily
collected. Data for SVL and clutch size based on measurements
and dissections of lizards in the sample are presented herein by
ranges of variation and/or means (to one decimal place) ± 1 SE.
Statistical analyses were performed at the University of Arkansas
on a PC loaded with institutionally licensed JMP software
(Version 12, SAS Institute, Inc., Cary, North Carolina, Copyright
© 2015).
The sample was collected by JAL-E on 21 July 2002, during
the herpetofaunal study of Chihuahua published by Lemos-Es-
pinal and Smith (op. cit.), from a site at Caseta Galeana, Llanos
de Flores Magón (30.008306°N, 107,253222°W, WGS 84; elev. 1500
m) which is ~189 km due south of a point on the New Mexico
state boundary. This is an extensive plains area surrounded by
high mountains. Most of the plains consisted of gravelly soil
with deposits of sand apparent mostly along rivulets. The do-
minant vegetation consisted of Creosote Bush (Larrea triden-
tata), with interspersed Torrey Yucca (Yucca torreyi) and Honey
Mesquite (Prosopis glandulosa). Aspidoscelis uniparens was the
most common lizard species observed in this area, becoming
active at ~0830 h. Gonochoristic Texas Spotted Whiptail (A. gu-
laris) was also present, but in lesser numbers. Each specimen
of A. uniparens was identified by JMW based on color pattern,
qualitative characters of scutellation, and results of statistical
treatments of meristic characters. These lizards were assigned
the following numbers of Laboratorio de Ecología, Unidad de
Biotecnología y Prototipos (= LEUBIPRO) 9718–9724, 9731–9740,
9744–9748, 9750–9781 (N = 54).
We sorted the specimens into subsamples consisting of gra-
vid females and females devoid of clutch development, respecti-
vely. Data for females with either ovarian (N = 28, 84.8%) or ovi-
ductal (N = 5, 15.2%) egg development included: body size range
60–76 mm SVL; mean 65.8 ± 0.64 mm; N = 33 (61.1% of entire
sample). Numbers of clutches of two (N = 22), followed by clut-
ches of three (N = 10), presented by SVL to nearest mm included:
60 (N = 1, 0); 62 (4, 2); 63 (2, 1); 64 (5, 0); 65 (1, 2); 66 (1, 1); 67 (3,
1); 68 (0, 1); 69 (2, 0); 70 (2, 1); 71 (1, 0); 73 (0, 1). Only the largest
lizard in the entire sample, namely a female of 76 mm SVL had a
clutch of four eggs. The sample of 54 A. uniparens included only
five individuals of over 69 mm SVL (70, 70, 70, 73, 76). Data for
females devoid of clutches included: range 55–72 mm SVL; mean
64.3 ± 0.89 mm; N = 21 (38.9%). These specimens presented by
SVL to nearest mm were: 55 (1); 56 (1); 62 (3); 63 (3); 64 (2); 65 (1);
66 (3); 67 (2); 68 (3); 70 (1); 72 (1). Only the two smallest lizards
devoid of clutched were below the minimum size at maturity
based on the smallest gravid female. Based on 33 gravid fema-
les from the study site in Chihuahua the usual clutch sizes were
either two (SVL range 60–71, mean 65.3 ± 0.69 mm; N = 22) or
three (SVL range 62–73, mean 66.1 ± 1.03 mm; N = 10) eggs. Clu-
tch data for the entire sample were: range 2–4; mean 2.4 ± 0.10;
symmetrical clutches N = 16 (48.5%); asymmetrical clutches N
= 17 (51.5%). Linear regression calculated in JMP indicated the
following relationship between SVL and clutch size for the 33
gravid Chihuahua females of A. uniparens (adjusted r2 = 0.1019; P
0.0393). We offer the hypothesis that many of the lizards devoid
of clutches had already oviposited a minimum of one clutch in
the summer they were collected and that many of the lizards with
clutches had also deposited a previous clutch that year.
Specimens referenced herein were collected in Chihuahua by
JAL-E and imported to the University of Colorado for study by
HMS under authority of permits provided by the government of
Mexico. The specimens were then shipped to the University of
Arkansas for further analyses by JMW. They were subsequently
deposited in the collections of the University of Colorado Mu-
seum of Natural History in Boulder, Colorado, and Laboratorio
de Ecología, Unidad de Biotecnología y Prototipos, Facultad de
Estudios Superiores Iztacala, UNAM, Estado de México, Mexico.
JULIO A. LEMOS-ESPINAL, Laboratorio de Ecología-UBIPRO, FES Izta-
cala UNAM, Av. Los Barrios No. 1, Los Reyes Iztacala, Tlalnepantla, México –
54090 (e-mail:; JAMES M. WALKER, Department of Bio-
logical Sciences, University of Arkansas, Fayetteville, Arkansas 72701, USA
(e-mail:; HOBART M. SMITH, University of Colorado
Museum of Natural History, Boulder, Colorado 80309, USA.
cus vittatus is native to Central and South America, and was
introduced to South Florida in the late 1970s as a result of the
exotic pet trade (Wilson and Porras 1983. The Ecological Impact
of Man on the South Florida Herpetofauna. Special Publication
No. 9, University of Kansas Museum of Natural History and World
Wildlife Fund-US, Lawrence, Kansas. 89 pp.). The species has
since been documented in nine Florida counties (Krysko et al.
2011. Atlas of Amphibians and Reptiles in Florida. Final Report,
Project Agreement 08013, Florida Fish and Wildlife Conservation
Herpetological Review 48(3), 2017
Commission, Tallahassee, Florida. 524 pp.). Little is known re-
garding the natural history of introduced B. vittatus in Florida.
The only known predators of B. vittatus in Florida are snakes
(Meshaka et al. 2004. The Exotic Amphibians and Reptiles of
Florida. Krieger Publishing Company, Malabar, Florida. 155 pp.),
wading birds, and large fish (Flaherty and Friers 2014. Southeast.
Nat. 13:N57–N58). Here we report an observation of predation on
B. vittatus by a Northern Mockingbird.
In July 2015 a mixed-age group of five B. vittatus were ob-
served basking at the edge of a canal fringed with dense thi-
ckets of Schinus terebinthifolia (Brazilian Pepper) at Homestead
Air Reserve Base, Miami-Dade County, Florida (25.489166°N,
80.371666°W; WGS 84). An adult Mimus polyglottos (Northern
Mockingbird) was observed landing among the group, grasping
a hatching B. vittatus in its beak, and swallowing it whole. The re-
maining adults and sub-adults fled into the adjacent vegetation.
This is the first documented case of M. polyglottos preying
upon B. vittatus. M. polyglottos is a generalist omnivore, known
to feed on native Anolis lizards (Farnsworth et al. 201l. In P. G.
Rodewald [ed.], The Birds of North America. Cornell Lab of Or-
nithology, Ithaca, New York; retrieved from
Species-Account/bna/species/normoc), as well as non-native
Leiocephalus carinatus (Northern Curly-tailed Lizard) (Smith et
al. 2006. Herpetol. Rev. 37:224). Because of this, it is likely that M.
polyglottos is a frequent predator of B. vittatus hatchlings where
the two species co-occur in South Florida.
JAMES P. FLAHERTY, Central Florida Zoo’s Orianne Center for Indigo
Conservation, Eustis, Florida 32726, USA; JOSHUA FRIERS, United States
Department of Agriculture Wildlife Services, Homestead, Florida 33030,
USA (e-mail:
CALOTES MYSTACEUS (Moustached Crested Lizard). DEFEN-
SIVE BEHAVIORS. Calotes mystaceus is a diurnal, arboreal, and
insectivorous agamid common in forested areas throughout
southeast Asia. Adult males can change color (Zug et al. 2010.
Salamandra 46:104–107), notably turning blue during their sum-
mer breeding season (Chan-ard et al. 2015. A Field Guide to the
Reptiles of Thailand. Oxford University Press, New York, New
York. 352 pp.). Herein we report color change and an apparent
ontogenetic shift in the defensive tactics of adult and juvenile C.
Our five observations occurred between September and
November 2016 in the Sakaerat Biosphere Reserve (SBR), located
in Nakhon Ratchasima Province, in northeastern Thailand
(14.44–14.55°N, 101.88–101.95°E; WGS 84). We located an adult
C. mystaceus (SVL ca. 10.0 cm) about 2 m up a deciduous tree
at 2045 h on 12 November 2016. Ambient temperature was
22.4°C, ground temperature was 22.7°C, and ambient relative
humidity was 100%. We approached within 1–2 m of the lizard
and photographed the individual prior to disturbing it. Upon
closer approach (< 1 m) the individual twitched and became a
blue color, but did not flee. We then took another photograph to
document the animal’s response color (Fig. 1). Subsequently, we
handled the lizard and the blue color became more apparent.
Researchers reported two additional observations of the
same defensive color change by adult C. mystaceus in the SBR
(Cameron Hodges and Sean Laughlin, pers. comm.).
We observed two juvenile C. mystaceus at 1352 h on 25 No-
vember 2016, on a sunny day with few clouds. The lizards were in
the station headquarters parking area on a tree (Shorea obtusa)
surrounded by a 1-m ring of dirt and woody shrubs (Phyllanthus
emblica; Wrightia tomentosa). Ambient temperature was 33.2°C,
ground temperature on the soil was 33.8°C, and ambient relative
humidity was 95%. When approached within 1–2 m, the smaller
juvenile (SVL ca. 3.0 cm) fled towards the brush without chan-
ging color. The larger juvenile (SVL ca. 6.0 cm) displayed dark red
spots on its dorsum (Fig. 2). This individual also fled further up
the tree in short bursts of about 0.5 m, visibly pausing twice. Du-
ring both pauses the animal jerked its head straight backwards
2–3 times.
Individuals in all life stages appeared to first rely on cryptic
defense, but upon approach differed in defensive tactics. The
smaller juvenile, most vulnerable to predation, quickly fled. The
larger juvenile may have used bright red spots and head jerks in
an attempt to deter a predator, whereas the adults initially chan-
ged color and stood their ground. Our observations suggest that
C. mystaceus undergo an ontogenetic shift in defensive strate-
gies, with older individuals utilizing color change. However, it is
also possible that the color change associated with our approach
was an artifact of increased hormone levels (e.g., testosterone)
due to the approach.
WAENGSOTHORN, Sakaerat Environmental Research Station, Nakhon
Ratchasima, Thailand; COLIN T. STRINE, Suranaree University of Technol-
ogy, Nakhon Ratchasima, Thailand (e-mail:
Fig. 1. Calotes mystaceus adult before (A) and after (B) approach.
Fig. 2. Calotes mystaceus larger juvenile before (A) and after (B)
approach, and the smaller juvenile (C) after approach.
Herpetological Review 48(3), 2017
COLOBOSAURA MODESTA (Bahia Colobosaura). DIET. The
Gymnophthalmidae currently includes 235 species of small
spectacled lizards (; accessed 23
January 2017). Lizards from this family are widely distributed
in Central and mostly South America, from southern Mexico
to Argentina, including some Caribbean and South American
islands, and exhibit an array of morphological specializations,
with different degrees of body elongation and limb reduction
(Ávila-Pires 1995. Zool. Verhand. 299:1–706; Pellegrino et al.
2001. Biol. J. Linn. Soc. 74:315–338).
There is published information on the diet of
gymnophthalmids from different biomes, including the Amazon
forest (Vitt et al. 1998. Can. J. Zool. 76:1671–1680; Vitt et al. 1998.
Can. J. Zool. 76:1681–1688; Caldwell and Vitt 1999. Oikos 84:383–
397), Cerrado (Vitt 1991 J. Herpetol. 25:79–90; Mesquita et al.
2006. Copeia 2006:460–471), Caatinga (Oliveira and Pessanha
2013. Biota Neotrop. 13:193–198), dry forest (Werneck et al. 2009.
Aust. Ecol. 34:97–115), open areas (Vitt and Carvalho 1995. Co-
peia 1995:305–329), Andes (Doan 2008. J. Herpetol. 42:16–21),
and coffee plantations (Anaya-Rojas et al. 2010. Pap. Avulsos
Zool., São Paulo 50:159–166). Data on the diet of Atlantic forest
gymnophthalmids are restricted to few species (Rocha 1991.
Herpetol. Rev. 22:40–42; Teixeira and Fonseca 2003. Bol. Mus.
Biol. Mello Leitão 15:17–28; Eisemberg et al. 2004. Herpetol. Rev.
35:336–337; Maia et al. 2011. Zoologia 28:587–592).
Colobosaura modesta (Reinhardt and Lütken, 1862) is
distributed in Brazil, Paraguay, and Argentina (Ávila-Pires 1995.
Zool. Verhand. 299:1–706; Cacciali 2010. Rep. cient. FACEN
1:10–19). In Brazil, it is found in the Cerrado (Brazilian savanna),
Amazon and Atlantic forests and forest enclaves areas across the
states of Bahia, Ceará, Distrito Federal, Goiás, Maranhão, Mato
Grosso, Mato Grosso do Sul, Minas Gerais, Pará, São Paulo, and
Tocantins (Freire et al. 2012. Check List 8:970–972 and references
therein). Biological data on C. modesta are rare. Based on 13
individuals from a Cerrado area, Mesquita et al. (2006. Co-
peia 2006:460–471) found orthopterans as the main food item,
followed by Araneae and other insects (Formicidae, Homoptera,
Mantodea and Solifuga), while Werneck et al. (2009. Aust. Ecol.
34:97–115), based on eight individuals from another Cerrado
area, found only Blattaria in the stomachs. Here we report
additional (new) dietary items for C. modesta.
Between October 2008 and June 2009, we captured 31
individual C. modesta in pitfall traps with drift fences in two areas
of Atlantic rainforest located at Gália (N = 1) and Lupércio (N =
6) municipalities and in three areas of eucalyptus plantations in
Avaí (N = 4), Piratininga (N = 15) and Arealva (N = 5) municipalities
(Brassaloti 2010. M.S. Thesis, Universidade Estadual Paulista,
São José do Rio Preto, São Paulo, Brazil. 135 pp.); all localities
were in the state of São Paulo, southeastern Brazil. Animals
were collected under the ICMBio collection license number
18204-1, euthanized in a CO2 saturated atmosphere, fixed in 10%
formaldehyde and preserved in 70% ethanol. Voucher specimens
were deposited in the herpetological collection of the Escola Su-
perior de Agricultura Luiz de Queiroz, Universidade de São Pau-
lo, Brazil (VESALQ 1033–1063).
Gastrointestinal tracts were extracted and analysed under
a stereomicroscope. Three individuals (9.7 %) had empty
stomachs. Food items were identified to the lowest possible
taxonomic level based on arthropod identification keys (spiders:
Kaston et al. 1978. How to Know the Spiders. McGraw-Hill Sci-
ence, Boston, Massachusetts. 288 pp.; isopods: Schmalfuss
2003. World Catalog of Terrestrial Isopods [Isopoda: Oniscidea].
Stuttgarter Beitrage zur Naturkunde, Serie A, Nr. 654, Stuttgart,
Germany. 341 pp.; insects: Rafael et al. 2008. Insetos do Brasil: Di-
versidade e Taxonomia. Holos Editora, Ribeirão Preto, São Paulo,
Brazil. 810 pp.).
The diet of Colobosaura modesta was composed of 30 food
items distributed across eight prey categories (Table 1). The
most abundant and frequent item in the stomachs was Araneae
(spiders), followed by Blattaria (cockroaches). The less abundant
items were Scorpiones, Phthiraptera (lice), insect larvae and
Based on the variety of taxa found in the stomachs, we
conclude that Colobosaura modesta has a generalist diet with a
major focus on spiders. However, because we did not sample the
available prey, we cannot be certain of this. Our data are similar
to those of Mesquita et al. (op. cit.), although these authors also
found items not present in our sample (e.g., Hymenoptera,
Hemiptera, Orthoptera, and Solifuga). Usually, spiders are an
important part of the diet of gymnophthalmid lizards, such
as Ecpleopus gaudichaudii (Eisemberg et al. 2004. Herpetol.
Rev. 35:336–337; Maia et al. 2011. Zoologia 28:587–592) and
Prionodactylus eigenmanni (Vitt et al. 1998. Can. J. Zool. 76:1681–
1688). Araneae has been considered a secondary prey for other
Gymnophthalmidae, such as Anotosaura vanzolinia (Oliveira
and Pessanha 2013. Biota Neotrop. 13:193–198), Ptychoglossus
bicolor (Anaya-Rojas et al. 2010. Pap. Avulsos Zool., São Paulo
50:159–166), Proctoporus bolivianus, P. pachyurus, P. sucullucu,
and P. unsaacae (Doan, op. cit.). Therefore, the importance of
Araneae in Gymnophthalmidae is probably species-specific.
Phthiraptera is not a common prey item in lizard diets. Some
Phthiraptera have phoretic behavior (Harbison et al. 2009. Int.
J. Parasitol. 39:569–575) using flies, fleas, dragonflies, butterflies,
and beetles as transport vectors. The Phthiraptera found in our
sample belonged to the family Gyropidae, which are typical
rodent parasites (Pedro M. Linardi, pers. comm.). The lizard that
ate Phthiraptera, also ate Blattaria, Isoptera, and larva. However,
there is no record of Phthiraptera using Isoptera or Blattaria for
phoresy. The pitfall traps also captured rodents, which could
potentially transport Phthiraptera.
Our study recorded two new food items (e.g., Isopoda and
Phthiraptera) that were not previously found in C. modesta
table 1. Food items identified in the diet of the lizard Colobosaura
modesta from southeastern Brazil.
Food items Abundance Frequency of
of item (%) stomachs with item
Araneae 15 (50) 4 (29)
Scorpiones 1 (3.3) 1 (7)
Blattaria 6 (20) 4 (29)
Isoptera 3 (10) 1 (7)
Mantodea 2 (6) 1 (7)
Phthiraptera (Gyropidae) 1 (3.3) 1 (7)
Insect larva 1 (3.3) 1 (7)
Isopoda 1 (3.3) 1 (7)
Total 30 (100) 14 (100)
Herpetological Review 48(3), 2017
stomachs. In fact, it was the first time Phthiraptera was found as
a food item for any gymnophthalmid lizard.
We thank Pedro Marcos Linardi (Universidade Federal de
Minas Gerais, Brazil) for the identification of the louse and CNPq
(Conselho Nacional de Pesquisa e Desenvolvimento Tecnoló-
gico) for grants to GAT, CCE and JB (processes 147271/2014-2,
233418/2014-8 and 309017/2016-5, respectively).
GUSTAVO A. TORELLI, Escola Superior de Agricultura Luiz de Queiroz,
Universidade de São Paulo, Piracicaba, SP, Brazil; CARLA C. EISEMBERG,
Research Institute for the Environment and Livelihoods, Charles Darwin
University, Darwin, NT, Australia; RICARDO A. BRASSALOTTI, JAIME
BERTOLUCI, Escola Superior de Agricultura Luiz de Queiroz, Universidade
de São Paulo, Piracicaba, SP, Brazil (e-mail:
CYRTODACTYLUS YOSHII ( Yoshi’s Bow-fingered Gecko). RE-
PRODUCTION. Cyrtodactylus yoshii is endemic to northern Bor-
neo (Das 2004. Lizards of Borneo. Natural History Publications,
Kota Kinabalu, Malaysia. 83 pp.). To my knowledge the reproduc-
tion of this species has not been studied. In this note I report the
initial reproductive information for C. yoshii.
A sample of 39 C. yoshii consisting of 13 adult males (mean
SVL = 87.5 mm ± 6.1 SD, range = 75–98 mm), 23 adult females
(mean SVL = 95.5 mm ± 5.3 SD, range = 82–101 mm) and 3 suba-
dult females (mean SVL = 66.7 mm ± 2.3 SD, range = 64–68 mm)
from Sabah, Malaysia collected from 1986 to 1993 and deposi-
ted in the herpetology collection of the Field Museum of Natu-
ral History (FMNH), Chicago, Illinois, USA was examined: (by
district) Lahad Datu: FMNH 230087–230089, 230105, 235118,
235121, 235125, 235136, 235141, 235144, 235146, 235147, 240622,
246200, 246209, 246212, 246214, 246216, 246217, 246221, 246222,
246225–246228, Ranau: FMNH 249765, 249766, 249768, 251008,
Tawau: FMNH 248069–248071, 248072, 248074–248077, 248978,
A cut was made in the lower abdominal cavity and the left
testis or ovary was removed, embedded in paraffin, cut into 5-µm
sections and stained by Harris hematoxylin, followed by eosin
counterstain. Enlarged follicles (> 4 mm) or oviductal eggs were
counted. Histology slides were deposited in FMNH.
The only stage noted in the testicular cycle was spermiogene-
sis in which seminiferous tubules are lined by sperm or clusters
of metamorphosing spermatids. Males undergoing spermioge-
nesis were collected in January (N = 1), July (N = 2), August (N
= 2), September (N = 1), October (N = 5), November (N = 2). The
smallest reproductively active male (spermiogenesis) measured
75 mm SVL (FMNH 248071) and was collected in July.
Four stages were observed in the ovarian cycle (Table 1): 1)
quiescent, no yolk deposition; 2) early yolk deposition, basophilic
vitellogenic granules in the ooplasm; 3) enlarged follicles > 4
mm; 4) oviductal eggs. Mean clutch size (N = 19) = 1.99 ± 0.32 SD,
range = 1–2. One female (FMNH 235136) collected in December
contained one oviductal egg and also exhibited concurrent early
yolk deposition in a separate follicle, indicating C. yoshii may
produce multiple clutches in the same reproductive season. The
smallest reproductively active female (2 eggs > 4 mm) measured
82 mm SVL (FMNH 249765) and was collected in December.
Three smaller C. yoshii (SVLs = 64, 68, 68 mm) contained
quiescent ovaries and were considered to be subadults.
While it is clear C. yoshii, like other south Asian Cyrtodactylus
lizards, exhibits an extended reproductive cycle (Goldberg and
Grismer 2014. Herpetol. Rev. 45:327; Goldberg and Grismer 2015.
Herpetol. Rev. 46:89–90; Goldberg and Grismer 2016. Herpetol.
Rev. 47:135–136), examination of additional monthly samples are
needed before all aspects of the reproductive cycle can be known.
I thank Alan Resetar (FMNH) for permission to examine C.
STEPHEN R. GOLDBERG, Whittier College, Department of Biology,
Whittier, California 90608, USA; e-mail:
ELGARIA COERULEA (Northern Alligator Lizard) AQUATIC
HABITAT USE. Aquatic habitat use by terrestrial lizards has
generally been reported as either escape or foraging behaviors
by swimming in streams or ponds. North American lizards
exhibiting this behavior include Aspidoscelis sexlineatus (Dillon
and Baldauf 1945. Co peia 1945:174; Trauth et al. 1996. Herpetol.
Rev. 27:20), Crotaphytus collaris (Burt and Hoyle 1934. Trans.
Kansas Acad. Sci. 37:193– 216), Elgaria multicarinata (Cowles
1946. Copeia 1946:105), Gambelia wislizenii (Medica 2010.
Herpetol. Rev. 41:354–355), Sceloporus clarkii (Zylstra and Weise
2010. Herpetol. Rev. 41:86), Sceloporus torquatus (Marquez et
al. 2014. Herpetol. Rev. 45:134), Scincella lateralis (Akin and
Towsend 1998. Herpetol. Rev. 19:43), and Uma exsul (Estrada-
Rodriguez and Leyva-Pacheco 2007. Herpetol. Rev. 38:84–85).
Alligator lizards (genus Elgaria) can often be found near or
along stream banks, riparian edges, or moist canyons in a variety
of upland habitats in the North American West (Stebbins 2003. A
Field Guide to Western Reptiles and Amphibians, 3rd ed. Houghton
Mifflin Co., Boston, Massachusetts. 533 pp.). Elgaria multicarna-
ta (Southern Alligator Lizard) (Stebbins and McGinnis 2012. Field
Guide to Amphibians and Reptiles of California, 2nd ed. University
of California Press, Berkeley, California. 538 pp.) and E. coerulea
(Storm and Leonard 1995. Reptiles of Washington and Oregon,
Seattle Audubon Society, Seattle, Washington. 176 pp.) are both
known to be relatively good swimmers by undulating their body
and using their long tail to move through the water.
Elgaria coerulea is a predominantly terrestrial and secretive
anguid lizard, ranging from the Pacific Northwest south through
the Coast Ranges to Monterey County, California and through the
Cascades Range and the Sierra Nevada to Kern County, Califor-
nia (Stebbins 2003, op. cit.). It typically inhabits cool mesic wood-
lands, forests, and grasslands, and especially frequents vegetated
areas near or along stream banks, creeks, and cobble bars (Lais
1976. Cat. Amer. Amphib. Rept. 178:1–4; Sabo and Powers 2002.
Ecology 83:1860–1869; Stebbins and McGinnis 2012, op. cit.). To
our knowledge, use of aquatic habitat beyond these general de-
scriptions has not been documented. Herein we report repeated
aquatic habitat use by E. coerulea in montane wet-meadows and
fens in the southern Cascades Range of California.
table 1. Monthly stages in the ovarian cycle of 23 adult female
Cyrtodactylus yoshii from Sabah, Malaysia. *One December female