Liolaemus occipitalis supervivencia

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Playing dead to stay alive: death-feigning in

Liolaemus occipitalis (Squamata: Liolaemidae)

Santos, M.B. et al.

Biota Neotrop. 2010, 10(4): 361-364.

On line version of this paper is available from:

http://www.biotaneotropica.org.br/v10n4/en/abstract?short-communication+bn03110042010

A verso on-line completa deste artigo est disponvel em:

http://www.biotaneotropica.org.br/v10n4/pt/abstract?short-communication+bn03110042010

Received/ Recebido em 27/04/2010 -

Revised/ Verso reformulada recebida em 03/11/2010 - Accepted/ Publicado em 19/11/2010

ISSN 1676-0603 (on-line)


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Biota Neotrop., vol. 10, no. 4

Playing dead to stay alive: death-feigning in

Liolaemus occipitalis (Squamata: Liolaemidae)

Maurcio Beux dos Santos1,3, Mauro Cesar Lamim Martins de Oliveira1,

Laura Verrastro2 & Alexandro Marques Tozetti1

1Laboratrio de Ecologia de Vertebrados Terrestres, Instituto de Cincias Biolgicas,

Universidade Federal do Rio Grande FURG, Av. Itlia, Km 8, CEP 96201-900, Rio Grande, RS, Brazil2Laboratrio de Herpetologia, Departamento de Zoologia, IB, Universidade Federal do Rio Grande do Sul,

Av. Bento Gonalves, 9500, CEP 91501-970, Porto Alegre, RS, Brazil3Corresponding author: Maurcio Beux dos Santos, e-mail: [email protected]

SANTOS, M.B., OLIVEIRA, M.C.L.M., VERRASTRO, L. & TOZETTI, A.M. Playing dead to stay alive: death-feigning in Liolaemus occipitalis (Squamata: Liolaemidae). Biota Neotrop. 10(4): http://www.biotaneotropica.org.br/v10n4/en/abstract?short-communication+bn03110042010.

Abstract: Predation is the major selective orce that drives the development o a series o deense mechanisms bythe species. These mechanisms are ecient in that it limits the ability o predators to detect, recognize and subduetheir prey. In lizards or example, the tail loss and locomotor escape are the most common deensive behavior

reported on the literature. Additionally, in response to external stimuli, some lizards exhibit a peculiar deensivebehavior: death-eigning or thanatosis. Here we describe the rst record o death-eigning in a population oLiolaemus occipitalis in a coastal sand dune habitat in Southernmost Brazil. A total o 86 individuals were tested.During handling 75.6% o lizards eigned death. The duration o this behavior was longer when the observerremained closer to the lizards, suggesting the ability oL. occipitalis to evaluate the predation risk.Keywords: thanatosis, behavior, sand-dune lizard, pampa, restinga.

SANTOS, M.B., OLIVEIRA, M.C.L.M., VERRASTRO, L. & TOZETTI, A.M. Simulando a morte paramanter-se vivo: tanatose emLiolaemus occipitalis (Squamata: Liolaemidae). Biota Neotrop. 10(4): http://www.biotaneotropica.org.br/v10n4/pt/abstract?short-communication+bn03110042010.

Resumo: A presso exercida pela predao a principal ora seletiva que direciona o estabelecimento de umasrie de mecanismos de deesa por parte das espcies. Esses mecanismos so ecientes na medida em que limitama capacidade dos predadores em detectar, reconhecer ou subjugar suas presas. Em lagartos, por exemplo, oscomportamentos deensivos mais requentemente citados na literatura so a autotomia caudal e a uga. Almdisso, em resposta a estmulos externos, alguns lagartos podem exibir um comportamento peculiar: ngem-se demortos (tanatose). Neste estudo descrevemos o primeiro registro de tanatose em uma populao de Liolaemusoccipitalis em rea de dunas costeiras no extremo Sul do Brasil. Um total de 86 indivduos oram testados.Durante o manuseio, 75,6% dos lagartos exibiram o comportamento de tanatose. A durao deste comportamentooi maior quando o observador permanecia a distncias mais curtas dos lagartos, sugerindo uma capacidade deL. occipitalis em avaliar o risco de predao.Palavras-chave: tanatose, comportamento, lagartixa-das-dunas, pampa, restinga.
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Santos, M.B. et al.

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Liolaemus occipitalis Boulenguer, 1885 is a small sand-dune-lizardassociated to coastal habitats and its geographic distribution restrictedto Rio Grande do Sul and Santa Catarina States, southern Brazil(Bujes & Verrastro 2008), and Uruguay (Verrastro et al. 2006). It isprimarily an insectivorous reptile that reproduces between Septemberand March (Verrastro & Krause 1994). Despite being a threatenedspecies (Marques et al. 2002), available data on its natural history and

behavior obtained in the eld are scarce. In addition, the impact thatsand dune habitats have been enduring highlight the importance oecological studies on species associated to this environment (Seeliger2003). Here we report observations o the deensive behavior oL. occipitalis in nature and the rst record o death-eigning in thisspecies.

Methods

The data presented in this study was collected between November2009 and January 2010 in the municipality o Rio Grande, RS, in anarea known as Balnerio Cassino, southernmost Brazil (32 07 54.65- 32 17 35.07 S and 52 06 38.80 - 52 20 53.36 W) locatedat sea level. The area consists o well preserved wet habitats ormed

by meadows with a mosaic o dunes, sand dune vegetation andtemporary lagoons. The climate is classied as subtemperate (Malu2000), with an average annual maximum temperature o 23.3 Cand average annual minimum temperature o 12.7 C. The seasonsare well dened, with rainall homogeneously distributed along theyear and averaging 1252 mm between 1931and1990 (INMET 2010).During eld surveys o the regional herpetoauna, we captured86 individuals oLiolaemus occipitalis using pitall traps with dritences (Brasileiro et al. 2005). The lizards remained a maximum oeight hours in buckets containing water and shelter to protect themagainst the sunlight, minimizing the stress on captured animals.All lizards were handled careully during measurements (body sizeand mass), sex determination, and assessment o the reproductivecondition o emales by palpation (Fitch 1987). While being held,some individuals eigned death (Figure 1). The animals that exhibitedthis behavior were immediately placed on the ground and observedrom two distances, 1 and 7 m, whose order was set at random.In one o the distances, individuals were observed until recovery.The animal was then captured by hand and i death-eigning wasobserved again, the observation was repeated rom a second distancerom the lizard. The duration o the behavior was recorded in eachobservation. Only animals that exhibited this behavior rom the twodistances rom the observers (treatments) were considered. Becausethis species exhibit intraspecic agonistic interactions (M.B. Santos,personal observation), we only tested individuals that were alone inthe traps. This measure was taken to reduce the stress level o lizardsprior to manipulation. Each observation was considered a sample.

Comparisons between the durations o the behavior rom dierentdistances were conducted with a paired Wilcoxon test (Zar 1999).Ater recording the behavior, lizards were toe-clipped (Waichman1992) beore being released ve meters rom traps. Recapturedanimals were not included in the behavior assessment.

Results and Discussion

Eighty-six lizards (47 males, 31 emales, and 8 juveniles) werecaptured. The average size o animals was 47.5 9.3 mm SVL(range - emales: 37.0 66.0 mm; males: 40.0 63.0 mm; juveniles:22.0 30.0 mm) and the average body mass was 3.60 1.6 g (range -emales: 1.3 5.3 g; males: 1.8 7.8 g; juvenile: 0.3 0.8 g). Duringhandling, 65 lizards (75.6%) eigned death, 35 were males, 23 wereemales and seven were juveniles (Figure 2). This represents 74.5%o males, 74.2% o emales, and 87.5% o juveniles.

Introduction

In general, animals show a series o traits and deensivemechanisms to avoid predation and stay alive. Organisms reduce theprobability o being predated by using many adaptations or behavioralstrategies, which may present specic behaviors (displays) that candiscourage a predators attack (Greene 1988). Predation pressure

is a undamental selective orce or the development o a series otraits that increase prey survival, by limiting the predator ability todetect, recognize, approach, subjugate or consume the prey (Endler1986, Greene 1988, Lima & Dill 1990). In general, antipredatormechanisms are an important actor in the evolutionary processes omany animals (e.g., Vermeij 1982, Lima & Dill 1990). Vertebrates,in particular the Squamata, exhibit extremely diverse antipredatortactics (e.g., Pough et al. 2004, Rocha 1993), making this groupan interesting model to examine evolutionary mechanisms to avoidpredation (e.g., Greene 1988). In snakes, many deensive displayshave been characterized including various ways o intimidating oravoiding aggressors (Greene 1988, Tozetti et al. 2009). Unortunately,much less inormation is available on the deensive behavior o lizardscompared to snakes. The most reported deensive behavior o lizards

in the literature is tail loss (autotomy) and locomotor escape (Greene1988, Roberts et al. 1998, Rocha-Barbosa et al. 2008). Additionally,some lizards exhibit a peculiar deensive behavior: death-eigning.Death-eigning (thanatosis) is a state o immobility assumed bymany animals in response to external stimuli. This behavior has beenconsidered a deense mechanism against predators and is shared bymammals (e.g., Francq 1969), birds (e.g., Sargeant & Eberhardt1975), shes (e.g., Howe 1991), amphibians (e.g., Gargaglioni et al.2001, Bertoluci et al. 2007), reptiles (e.g., Greene 1988), andinsects (e.g., Acheampong & Mitchell 1997). Death eigning is alsoknown as catalepsy, or tonic immobility. In most cases, animals thatexhibit this behavior play dead by maintaining a rigid posture orby simulating ully relaxed muscles (e.g. ainting; Greene 1988).

Apparently thanatosis is a behavior that rarely occurs spontaneously(Greene 1988). Immobility might discourage sequential attacks,allowing escape, as demonstrated in experiments using invertebrates(Miyatake et al. 2004). While eciency in this deensive behaviorhave been poorly tested, its occurrence in several vertebrate groupssuggests a avorable evolutionary pressure toward its maintenance(Miyatake et al. 2004). In rogs, or example, the thanatosis waspointed out as an important secondary deense mechanism inPhillomedusa (Sazima 1974). In lizards, death-eigning has beenreported or several dierent amilies, such as Anelytropsidae(Torres-Cervantes et al. 2004), Crotaphytidae (Gluesing 1983), andScincidae (Langkilde et al. 2003). Among tropidurid lizards, thisbehavior has been observed inEurolophosaurus nanuzae Rodrigues,1981 (Galdino & Pereira 2002), E. divaricatus Rodrigues, 1984

(Gomes et al. 2004, Kohlsdor et al. 2004) previously Tropidurusnanuzae and T. divaricatus (Frost et al. 2001), T. torquatus Wied,1820 and T. hispidus Spix, 1825 (Bertoluci et al. 2006). Bertoluci et al.(2006) has also reported anecdotal records or several other speciesin the tropidurid amily. Among Liolaemidae, this behavior has beenreported only inLiolaemus lutzae Mertens, 1938 (Rocha 1993). Thisamily consists o 229 species (including subspecies) belongingto the genera Ctenoblepharys, Phymaturus, and Liolaemus. Thelatter comprises most o the species o the amily, with 200 species(Pincheira-Donoso et al. 2008a). Considering the great diversity inecological specializations and morphology,Liolaemus is an importantgroup to study adaptive radiations (Pincheira-Donoso et al. 2008b).The South AmericanLiolaemus lizardsoccurs in Argentina, Bolivia,Brazil, Chile, Paraguay, Peru, and Uruguay, which represents thewidest range o environments occupied by a single lizard genus. The


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The death-eigning behavior observed in our study was similarto that described in Tropidurus species by Bertoluci et al. (2006).During handling,L. occipitalis lizards exhibited very relaxed muscles,remained immobile in the observers hand, and maintained thisposture when placed on the ground (Figure 1). During death-eigning,

simultaneous slow and lateral tail movements and intermittentmovements o opening and closing the eyes were observed. Somestudies show that habitat characteristics coner dierent probabilitieso predation o lizards (Shepard 2007). Although L. occipitalis isassociated with open environments like sand dunes, the presenceo shrubs, even sparse ones, provides shelter and protection againstpredators (Bujes & Verrastro 1998). In addition, the cryptic colorationand the ability o digging burrows are probably its main strategiesagainst predation. Also, L. occipitalis is an ambush orager (M.B.Santos, personal observation), which might decrease the time oexposure to visually-oriented predators. Apparently, when a predatoris detected, the main active deense strategy oL. occipitalis isto escape and the capacity o burying (Bujes & Verrastro 1998).However, the capacity o escape, especially the speed, depends onbody temperature as well as reproductive conditions (Shine 1980,

Plummer 1997, Miles et al. 2000). As a result o these constraints, adiversication in deense strategies might increase the probability osurvival o individuals. Because thanatosis only occurs i stimulated(e.g. handling), this could indicate that this strategy represents anextreme measure used ater a rst attack by the predator. Thus, escapeand cryptic behavior combined represents ways to avoid perceptionor capture, while death-eigning might be a strategy used when thelizard has been already captured by the predator. It should be pointedout that the duration o thanatosis was signicantly longer (T = 0.0;P = 0.028; n = 12) when the observed remained near (averageduration = 10.2 1.99 minutes; range = 7.2 - 12.7 minutes) thanwhen the observed was arther away rom lizards (average duration =2.0 1.14 minutes; range = 0.5 - 3.83 minutes), suggesting that the

animals are capable o evaluating the threat level.It should be pointed out that during previous studies conducted

with other populations oL. occipitalis, death-eigning was rarelyobserved (M.B. Santos, personal observation). The high requencyo records o this behavior in the population o the present studymight be associated with a selective process toward this deensivemechanism. Studies on experimental handling demonstrated thattheduration o death eigning observed in insects was a result odierent selection pressures caused by predation (Prohammer & Wade1981). However, in our study, this hypothesis still needs to be urtherinvestigated with experimental studies in which the actual predationpressure is quantied, as well as the ecacy o death-eigning toavoid the action o predators.

Acknowledgments

We are grateul to Col. Augusto Martins or providingtransportation to the study area. The study was conducted withlaboratory assistance o several biologists, especially Tatiane PenteadoGonalves and Alexandre Hartmann. We also thank CNPq (process476661/2008-0) and FAPERGS (process 0902139) or grants receivedand ICMBio or collecting permits.

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a b

Figure 1. a) general aspect o an individual oLiolaemus occipitalis Boulenguer, 1885; and b) an individual eigning death.

Figura 1. a) aspecto geral de um indivduo deLiolaemus occipitalis Boulenguer, 1885; e b) indivduo exibindo o comportamento de tanatose.

Figure 2. Absolute number oLiolaemus occipitalis lizards captured bypitall traps with drit ence and number o lizards that eigned death duringhandling.

Figura 2. Nmero absoluto de indivduos deLiolaemus occipitaliscapturadosem armadilha de interceptao e queda e que apresentaram o comportamentode tanatosedurante a manipulao.


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Received 27/04/2010

Revised 03/11/2010

Accepted 19/11/2010


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Liolaemus occipitalis supervivencia