WWW.IRCF.ORG/REPTILESANDAMPHIBIANSJOURNALTABLE OF CONTENTS IRCF REPTILES & AMPHIBIANSIRCF REPTILES • VOL15, N &O 4AMPHIBIANS • DEC 2008 189 • 21(3):93–95 • SEP 2014

IRCF REPTILES & AMPHIBIANS CONSERVATION AND NATURAL HISTORY

TABLE OF CONTENTS

FEATURE ARTICLES Body-inversion. Chasing Bullsnakes (Pituophis catenifer insayi) in Wisconsin: the Cuban Racer, Cubophis On the Road to Understanding the Ecology and Conservation of the Midwest’s Giant Serpent ...... Joshua M. Kapfer 190 . The Shared History of Treeboas (Corallus grenadensis) and Humans on Grenada: cantherigerusA Hypothetical Excursion ...... cantherigerus (Dipsadidae):Robert W. Henderson 198 RESEARCHDeath-feigning ARTICLES or Warning Signal? . The Texas Horned Lizard in Central and Western Texas ...... Emily Henry, Jason Brewer, Krista Mougey, and Gad Perry 204 . The Knight Anole (Anolis equestris) in Florida 1 2 3 ...... Tomás M. Rodríguez-CabreraBrian J. Camposano, Kenneth, Javier L. Krysko, Torres Kevin M., and Enge, RubenEllen M. Donlan, Marrero and Michael Romero Granatosky 212 1Jardín Botánico de Cienfuegos, Pepito Tey, Cienfuegos, CP 59290, Cuba ([email protected]) CONSERVATION ALERT 2Departamento de Biología Animal y Humana, Facultad de Biología, Universidad de la Habana, La Habana, CP 10400, Cuba ([email protected]) . 3DivisiónWorld’s de Zoología Mammals de in CrisisVertebrados, ...... Instituto de Ecología y Sistemática, La Habana, CP 11900, Cuba ([email protected]) 220 . More Than Mammals ...... 223 . The “Dow Jones Index” of Biodiversity ...... 225

HUSBANDRY . Captive Care of the Central Netted Dragon ...... Shannon Plummer 226 redation is among the main selective pressures that drive body-inversion sensu stricto have ever been reported for any biological evolutionPROFILE (e.g., Futuyma 2013). Snakes are fre- West Indian snakes. Herein we report body-inversion behav- P . Kraig Adler: A Lifetime Promoting Herpetology ...... Michael L. Treglia 234 quent prey of other vertebrates. To avoid predation, they have ior in the Cuban Racer, Cubophis cantherigerus cantherigerus developed a wide varietyCOMMENTARY of defensive behaviors (see Greene (Bibron 1840), from western Cuba, and discuss its possible 1994 for a review). .Anti-predationThe Turtles Have Been Watching behaviors Me ...... reported for relationship to death-feigning Eric Gangloffor warning 238 signal. West Indian colubroidsBOOK include REVIEW immobility, tail-waving, con- cealment, locomotor. escape,Threatened anteriorAmphibians ofbody-raising the World edited by and S.N. dorStuart,- M. Hoffmann, J.S. Chanson, N.A. Cox, R. Berridge, P. Ramani, and B.E. Young ...... Robert Powell 243 soventral cobra-like neck-flattening, gaping, strike-feigning, biting, balling, cloacal CONSERVATION discharge, meal-disgorging, RESEARCH REPORTS: and Summaries rapid of Published Conservation Research Reports ...... 245 body rotation when NATURAL captured HISTORY (see Henderson RESEARCH REPORTS and Powell: Summaries of Published Reports on Natural History ...... 247  NEWBRIEFS ...... 248 2009 for a review). EDITORIAL INFORMATION ...... 251 These anti-predation FOCUS behaviors ON CONSERVATION are relatively: A Project simple You Can and Support ...... 252 generalized for most snakes (e.g., Mattison 1995), but a few species can display more complex patterns, including death- feigning. This is a state of external unresponsiveness to tactile stimulation, exhibited as a last-resortFront Cover.anti-predation Shannon Plummer. tactic Back Cover. Michael Kern Totat et velleseque audant mo Totat et velleseque audant mo for an individual unable to evade captureestibus inveliquo by a predatorvelique rerchil (e.g., estibus inveliquo velique rerchil Greene 1994, Caro 2005, Gregory 2008).erspienimus, Death-feigning quos accullabo. Ilibus can erspienimus, quos accullabo. Ilibus aut dolor apicto invere pe dolum aut dolor apicto invere pe dolum include body-inversion, kinetic (orfugiatis “erratic”) maionsequat display, eumque tonic fugiatis maionsequat eumque immobility, limp-mouth gape, tongue-hanging,moditia erere nonsedis eye ma rotation, sectiatur moditia erere nonsedis ma sectia- ma derrovitae voluptam, as quos tur ma derrovitae voluptam, as autohemorrhaging, cloacal discharge,accullabo. and even bradycardia (e.g., Heterodon platirhinos; see Platt 1969, Gehlbach 1970, McDonald 1974). Body-inversion usually occurs in associa- tion with death-feigning (e.g., Platt 1969, Gehlbach 1970), but Greene (1994) considered them independent behaviors (although he emphasized that they frequently appear in com- bination). Consistent with the separation of both behaviors, a warning function has also been suggested for body-inversion, particularly for certain death-feigning colubroids with strik- ingly marked venters such as Heterodon nasicus, Natrix natrix, and Diadophis punctatus (Heusser and Schlumpf 1962). Death-feigning has been widely documented in the families Colubridae, Dipsadidae, Natricidae, Elapidae, Fig. 1. A Cuban Racer (Cubophis c. cantherigerus) from the Jardín Botánico Leptotyphlopidae, and Tropidophiidae (for a review see Nacional de Cuba exhibiting a cobra-like defensive posture. Photograph by Greene 1994). However, neither death-feigning sensu lato nor Nancy de la C. Llevat.

Copyright © 2014. Tomás M. Rodríguez-Cabrera. All rights reserved. 93 RODRIGUEZ-CABRERA ET AL. IRCF REPTILES & AMPHIBIANS • 21(3):93–95 • SEP 2014

level at the Jardín Botánico Nacional de Cuba (23°00’00”N, 82°20’50”W; Datum: NAD 27 for Cuba), Arroyo Naranjo, La Habana Province, Cuba. The frog jumped to the ground and the snake rapidly followed. Once on the ground, four photographers surrounded the snake. About two minutes after the photographers started their work, the snake adopted the cobra-like defensive posture typical of this species and other alsophiines (Fig. 1; see also Henderson and Powell 2009). Each time the snake attempted to escape it was inter- cepted by one of the photographers, but never touched. After eight minutes of continuous harassment the snake lowered its head (Fig. 2) and suddenly rolled on its back and adopted an inverted semicoiled posture, with the head bent slightly upward (Fig. 3). The snake remained inverted for less than one minute before reverting to the normal position and Fig. 2. The same Cuban Racer lowering its head in transition to an resuming the cobra-like defensive posture for about five more inverted posture. Photograph by Roberto Cruz. minutes, after which it was allowed to escape. The snake was not subjected to any additional disturbance during the time it remained inverted. On 6 January 2012, at 1500 h, we observed a Cuban We consider this behavior as a particular case of death- Racer (ca. 120 cm total length) pursuing a Cuban Tree feigning, less elaborate than the stereotyped behaviors observed Frog (Osteopilus septentrionalis, Hylidae) in a bromeliad in continental species of the genera Heterodon, Masticophis, (Tillandsia valenzuelana) on a tree ca. 3.5 m above ground Diadophis, and Natrix (e.g., Platt 1969, Gehlbach 1970,

Fig. 3. The same snake in an inverted position. Photograph by Raimundo López-Silvero.

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Greene 1994). The individual described herein did not gape, tral coloration and body-inversion could serve as a warning extend its tongue, or discharge its scent glands, and it rapidly signal. However, additional observations are required to cor- reverted to a normal posture. However, we cannot identify roborate body-inversion in this species and to elucidate trig- an apparent advantage other than those related to death- gering factors and function. feigning. Removal of cues (e.g., movement) that elicit prey- killing behavior in a potential vertebrate predator seems rea- Acknowledgements son enough to adopt death-feigning, since it might provide Raimundo López-Silvero, Nancy de la C. Llevat, and Roberto an animal with subsequent opportunities for escape (Alcock Cruz provided valuable information. We calculated time 1975, Gregory 2008). The anti-predation function of death- frames for each behavior from their photographic sequences. feigning may lie in direct interference with the predator’s abil- ity to consume the prey (Honma et al. 2006). However, it Literature Cited may also depend on the predator not immediately consuming Alcock, J. 1975. Animal Behavior: An Evolutionary Approach. Sinauer Assoc., Inc., immobile prey, thereby giving the potential prey a window Sunderland, Massachusetts. Caro, T. 2005. Antipredator Defenses in Birds and Mammals. University of Chicago of opportunity to escape if the predator is even momentarily Press, Chicago, Illinois. inattentive or distracted (Gregory 2008). Futuyma, D.J. (ed.). 2013. Evolution. Third Edition. Sinauer Assoc., Inc., An alternative explanation for this behavior is as a warn- Sunderland, Massachusetts. ing signal, but this individual lacked aposematic coloration on Gehlbach, F.R. 1970. Death-feigning and erratic behavior in leptotyphlopid, colu- broid, and elapid snakes. Herpetologica 26:24–34. its ventral region (which is the rule for its conspecifics, but see Greene, H.W. 1994. Antipredator mechanisms in reptiles, pp. 1–152. In: C. Gans below). An inverted position could be assumed by the Cuban and R.B. Huey (eds.), Biology of the Reptilia. Volume 16, Ecology B. Alan R. Racer after prolonged harassment without direct aggression, and Liss, Inc., New York. as a last-resort tactic after failure of other anti-predation mecha- Gregory, P.T. 2008. Bluffing and waiting: Handling effects and post-release immo- bility in a death-feigning snake (Natrix natrix). Ethology 114:768–774. nisms (e.g., locomotor escape or the cobra-like posture). Death- Henderson, R.W. and R. Powell. 2009. Natural History of West Indian Amphibians feigning in other species (e.g., H. platirhinos and Naja haje) also and Reptiles. University Press of Florida, Gainesville. comes at the end of a repertoire of other defensive behaviors, Heusser, J. and H.U. Schlumpf. 1962. Totstellen bei der Barren-Ringelnatter- including intimidation and odor production (Mattison 1995). Natrix natrix helvetica. Die Aquarien- und Terrarienzeitschrift 15:214–218. Honma, A., S. Oku, and T. Nishida. 2006. Adaptive significance of death feign- Despite the lack of aposematic coloration in the individ- ing posture as a specialized inducible defense against gape-limited predators. ual described above, Schwartz and Henderson (1991) men- Proceedings of the Royal Society of London B Biological Sciences 237:1631–1636. tioned one pattern of C. cantherigerus that includes posterior Jaume, M.L. and O.H. Garrido 1980. Notas sobre mordidas de jubo Alsophis ventral scales “sometimes entirely red, usually dark with more (Serpentes: Colubridae). Miscelánea Zoológica 11:2–3. Mattison, C. 1995. The Encyclopedia of Snakes. Facts On File, Inc., New York. or less bold, red or yellow blotching.” The senior author has McDonald, H.S. 1974. Bradycardia during death-feigning of Heterodon platyrhinos observed individuals of C. c. cantherigerus and C. c. schwartzi Latreille (Serpentes). Journal of Herpetology 8:157–164. with posterior black and red ventral markings. Some indi- Neill, W.T. 1954. Evidence of venom in snakes of the genera Alsophis and Rhadinea. viduals of C. c. pepei from eastern Cuba possess blue throats Copeia 1954:59. Platt, D.R. 1969. Natural history of the Hognose Snakes Heterodon platyrhinos and bicolored bellies (Schwartz and Thomas 1960). Some and Heterodon nasicus. University of Kansas Publications, Museum of Natural specimens from the Península de Guanahacabibes have ven- History 18:253–420. tral scales with yellow ventral markings (J. Torres, pers. obs.). Schwartz, A. and R.W. Henderson. 1991. Amphibian and Reptiles of the West Indies: Descriptions, Distributions, and Natural History. University of Florida Press, Furthermore, C. cantherigerus is known to have Gainesville. Duvernoy’s glands and a venomous bite (Neill 1954, Jaume Schwartz, A. and R. Thomas. 1960. Four new snakes (Tropidophis, Dromicus, and Garrido 1980), which support the idea that striking ven- Alsophis) from the Isla de Pinos and Cuba. Herpetologica 16:73–90.

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