Herpetology Notes, volume 13: 649-660 (2020) (published online on 05 August 2020)

Unsuccessful predation attempts by snakes on anuran amphibians: How successful are snakes?

William P. Costa1,* and César C. Trevelin2

Abstract. Snakes and anurans are commonly involved in predator-prey interactions. The large number of records of predation events involving snakes and anurans suggests that snakes are frequently successful in their attempts to prey on these amphibians. However, two aspects of the data may lead to overestimates of the efficiency of the predator in these cases. One factor is that the principal source of the data on snake diets is the analysis of the stomach contents of snake specimens deposited in museums. The second is the paucity of data on the actual interaction between the predator and its prey, and the frequency with which the anurans are either able to escape the predator or are abandoned by snakes. Given this general lack of data on unsuccessful predator-prey interactions between snakes and anurans, the present study describes four unsuccessful predation attempts and discusses the potential consequences for both the predator and prey. These events involved three snakes of the family Dipsadidae and one of the family Viperidae, which are all venomous, and three anuran species of the family Leptadactylidae. The events were recorded at four localities in the Brazilian states of São Paulo, Minas Gerais, and Goiás between 2016 and 2018. In one of the four cases (Event 1), the anuran was able to escape, while in the other three events, the prey were abandoned by the snake. In one case (Event 3), the anuran was dead, while in the other two cases (Events 2 and 4), the animals were left with reduced mobility, and were thus more vulnerable to other predators. In two of these events (1 and 3), the predator attacked the anuran by grasping its posterior members, while in event 2, the strike was to the throat of the anuran, and in event 4, the anuran was grasped by its right flank. In the only case in which the prey was able to escape unharmed, the combination of the size of the animal and its resistance to the attack of the predator appear to have been decisive. In the three cases in which the anuran was abandoned by the predator, the initial point of contact may have determined the effectiveness of the attack, given the anatomical resistance exerted by the prey, even when it remains immobile. The fact that all the snakes included in the present study are venomous did not ensure the success of the predation attempts, which would have resulted in the waste of this resource (venom). Despite the difficulty of analysing unsuccessful predation attempts (and their consequences for both predator and prey) systematically, studies of this kind are necessary to advance the understanding of this type of interaction and, ultimately, to evaluate more conclusively the ecological dynamics of the relationship between snakes and anurans.

Keywords. snakes, anurans, ecology interaction, predation attempts, predator-prey

Introduction 1993; Martins and Oliveira, 1998; Wells, 2007; Toledo et al., 2007; Bernarde and Abe, 2010; Vargas-Salinas Snakes and anurans are important components of and Aponte-Gutierrez, 2013). The large number of terrestrial ecological communities, and are often records of this type of interaction indicate that anurans involved in predator-prey interactions, normally as are an important component of the diets of a number of predator and prey, respectively (Strüssmann and Sazima, different snake species (e.g. Martins et al., 1993; Toledo et al., 2007; Moya and Maffei, 2012; Bovo and Sueiro, 2012; Forti and Bertoluci, 2012; Falkenberg et al., 2013; Santos-Silva et al., 2014; Passos et al., 2017; Muscat and Moroti, 2018; Thaler et al., 2018). In rare cases, 1 Departamento de Bioestatística, Biologia Vegetal, however, large anurans are also known to have subdued Parasitologia e Zoologia, Instituto de Biociências, and ingested snakes, although these events appear to Universidade Estadual “Júlio de Mesquita Filho” (UNESP), be highly opportunistic (Branch, 1976; Silva, Costa campus Botucatu, SP, Brasil. and Feio, 2007; Camargo Filho et al., 2008; Fonseca 2 Departamento de Economia, Sociologia e Tecnologia, et al., 2012). It is worth mentioning that, one of the Faculdade de Ciências Agronômicas, Universidade Estadual “Júlio de Mesquita Filho” (UNESP), campus Botucatu, SP, advantages of exploiting anurans as a food is their lack Brasil. of anatomical structures that are difficult to digest, such * Corresponding author. E-mail: [email protected] as fur, claws or scales, which is further reinforced by 650 William P. Costa & César C. Trevelin the fact that many species form large aggregations of Paulo, and União de Minas (-19.2825°S, -50.4224°O), individuals in the vicinity of bodies of water during the in the state of Minas Gerais. The sites are located within breeding season (Duellman and Trueb, 1986; Wells, an area of transition between the Cerrado savannah of 2007). central Brazil and the Atlantic Forest of eastern Brazil, The typically discreet nature of predation events in and were originally covered by a mosaic of Cerrado the wild contributes to the scant and sporadic records of sensu stricto savanna vegetation and semi-deciduous this type of behaviour, which hamper a more systematic seasonal forest (Ab’Saber, 2003). All four localities are analysis of the phenomenon. In fact, the vast majority also within an area dominated by a tropical savannah of reported cases are derived from the analysis of the climate (Aw in the Köppen classification), with a dry stomach contents of specimens deposited in scientific season that coincides with the austral winter and rains collections, which represent the end result of successful concentrated during the austral summer (Rolim et al., predation events (Pombal, 2007). The efficiency of 2007; Cardoso et al., 2015; Reboita et al., 2015). this type of predatory behaviour in snakes nevertheless The events at Acreúna and Ouroeste were recorded at remain unclear, given the scant records of unsuccessful the margins of permanent lakes, while the record from attacks (Rocha-Lima et al., 2018). União de Minas was obtained at a temporary pond, and Unsuccessful predation events may result from errors the event at Palmeiras de Goiás took place in a fragment in the evaluation of the relative size of the prey, which of forest (Figures 1a, 2a, 3a, 4a). The encounters with is more typical of young snakes (Sazima and Martins, the animals occurred during nocturnal surveys of the 1990), in addition to the intrinsic predator-defence respective study sites. Once an attempted predation mechanisms available to the anurans (see Ferreira et al., was observed, the subsequent sequence of events was 2019). Unsuccessful attempts may result in energetic recorded from a distance of approximately two meters, attrition in both the predator and the prey which, in with a minimum of artificial illumination (red lights), in extreme cases, may even result in the deaths of both an attempt to minimise possible interference from the animals (Bernal and Palma, 2011; Cavalcanti et al., 2012; presence of the observers (Vargas-Salinas and Aponte- Caramaschi and Niemeyer, 2012; Fong et al., 2013). In Gutierrez, 2013; Mario-da-Rosa et al., 2020). some cases, the prey is too large to be swallowed by the predator, which may even be suffocated in the attempt Results and Discussion (e.g. Caramaschi and Niemeyer, 2012; Nogueira et al., The observed predation attempts involved four 2013; Filho et al., 2017), or it may escape, but eventually species of snake, including three representatives of the die from exhaustion or the toxic effects of the venom family Dipsadidae, Helicops modestus Günther, 1861, (Sazima and Martins, 1990; Rocha-Lima et al., 2018). Thamnodynastes strigatus (Günther, 1858), Xenopholis The systematic evaluation of these unsuccessful undulatus (Jensen, 1900), and one member of the family predation attempts, and their relevance to both the Viperidae, Bothrops moojeni Hoge, 1966. The potential predator and the prey, is a complex task, although it prey was all frogs of the family Leptodactilydae, that may provide fundamental insights into the dynamics of is, Leptodactylus chaquensis Cei, 1950, Leptodactylus the ecological interactions between snakes and anurans. labyrinthicus (Spix, 1824), and Physalaemus centralis In order to broaden the existing perspective on this Bokermann, 1962. All the snake species are known to important type of ecological interaction, the present include anurans in their diets (see Nogueira et al., 2003; study describes four unsuccessful predation attempts França et al., 2008; Scartozzoni, 2009), although none by snakes on anurans, with the aim of evaluating the of the anurans recorded as prey in the present study have eventual consequences for both predator and prey, from been recorded previously in the diets of any of the four the moment that the predator loses interest in the prey or snake species. it manages to escape. Helicops modestus vs. Leptodactylus chaquensis in Material and Methods Acreúna, Goiás (Event 01): this event occurred at 18:39 The events described in the present study were recorded h on 05-29-2016 near the internal margins of a lake (Fig. at four localities in central and southeastern Brazil– 1a) and it was perceived due to the distress vocalisations Acreúna (-17.3786°S, -17.3786°O) and Palmeiras de emitted by the anuran. Initially, an individual of the Goiás (-16.8758°S, -49.9702°O) in the state of Goiás, species Leptodactylus chaquensis was observed with Ouroeste (-20.0077°S, -50.4282°O), in the state of São its body slightly stretched out, partially submerged in Unsuccessful predation attempts by snakes on anuran amphibians 651 the water, with its right leg in this substrate and the left aquatic environments (Scartozzoni, 2005). The snakes leg hidden by the mud and vegetation (Fig. 1b). After of this tribe feed exclusively or predominantly on approximately one minute, a snake (Helicops modestus) fish (Scartozzoni, 2009), although anurans are often emerged from the substrate and moved its body in among the items recorded in studies of the diets of an attempt to turn the frog belly-up (Fig. 1c). At this Helicops species (Feltrim and Cechin, 2000; Martins moment, it was possible to confirm that the snake had and Duarte, 2003; Aguiar and Di-Bernardo, 2004; attacked the frog by seizing its left posterior member, Ávila et al., 2006; Teixeira et al., 2017; Thaler et al., near the heel. The efforts of the snake to ingest the frog 2018). Scartozzoni (2009) also recorded semi-aquatic lasted for approximately three minutes. After resisting lizards in the diet of the brown-banded water snake, the snake, the frog was able to return to its initial Helicops angulatus (Linnaeus, 1758). Few data are position and escape. At this moment, the snake retreated available on the composition of the diet of H. modestus, and dived into the water (Fig.1d). however, although anurans of the families Hylidae and The genera Helicops, Hydrops, and Pseudoerys are Leptodactylidae are common items (Lema et al., 1983; members of the tribe Hydropsini, in the subfamily Sawaya et al., 2008; Costa et al., 2009; Scartozzoni, Xenodontinae of the family Dipsadidae (Vidal et al., 2009). One important element of the ecology of 2000; Zaher, 1999; Zaher, 2009) and form a clade this species is the presence of Duvernoy’s glands, characterised by adaptations for the exploitation of which produce toxins that help the snake subdue and

Figure 1. Attempted predation of Leptodactylus chaquensis by Helicops modestus: (a) general view of the lake at the study site; (b) the frog, emitting distress vocalisations, attempts to escape from the predator; (c) the snake attempts to turn the frog belly-up; (d) the snake loses its prey and submerges itself in the substrate. 652 William P. Costa & César C. Trevelin immobilise its prey, as discussed by Oliveira et al. any reaction to the attack (Fig. 2c). The snake then (2017). According to these authors, the time taken to continued trying to swallow its prey, opening its mouth subdue the prey is correlated with its relative size and, and moving its lower jaw. This behaviour continued for once the venom is injected, it may take as long as 30 approximately 10 minutes, when the snake moved its minutes to act. body laterally at an angle of around 180°, from left to In addition to the fact that L. chaquensis is a robust right, and stopped trying to swallow its prey, which it frog, the site of the attack was an environment that both released (Fig. 2d). Once released, the frog attempted hampered the movement of the snake and supported the to escape from the site of the attack by recoiling its frog, a combination of factors that may have contributed posterior members and moving erratically through to the failure of the attack. Thaler et al. (2018) recorded the substrate. It presented signs of internal lesions, as a similar situation that resulted in a successful attack of indicated by the presence of blood in its mouth (Fig. a congener, Helicops infrataeniatus (Jan, 1865), on L. 2d). The snake moved immediately away from the site chaquensis. The principal similarities were observed and disappeared into the vegetation. in the way in which the snake attacked the frog. It is The genus Thamnodynastes Wagler, 1830 is composed important to note, however, that in this case, the authors of 20 species of small- to medium-sized snakes with removed predator and prey from the site of the attack, opistoglyphous dentition (Bailey, 1967; Franco, 1999; in a body of water, which may have facilitated the Franco and Ferreira, 2002; Bailey et al., 2005; Uetz et subsequent immobilisation of the frog by the snake, al., 2019). These snakes are dietary generalists that feed using constriction. In the present study, the fact that the primarily on vertebrates, although their preference for frog was able to escape meant that it was impossible anurans and the degree of specialisation for the predation to determine whether the attack was ultimately fatal for of these amphibians has resulted in the genus being the prey. Even so, Rocha and López-Baucells (2013) classified as anurophagous (see Bernarde et al., 2000; reported on the survival of a predation attempt by H. Ruffato et al., 2003; Bellini et al., 2013; Dorigo et al., angulatus on Boana boans (Linnaeus, 1758). In this 2014; Bortolanza-Filho et al., 2019). Thamnodynastes case, the frog survived, but presented swelling and strigatus (Günther, 1858) is typical of the genus, depigmentation of the area of its body that was attacked and is known to prey on an ample variety of anurans by the snake, and it seems likely that the L. chaquensis (Bernarde et al., 2000; Ruffato et al., 2003; Winkler et attacked in the present study may have suffered a similar al., 2011; Preuss and Tozetti, 2018; Bortolanza-Filho et fate. The sum of this evidence indicates that medium- al., 2019). The ecological characteristics of this snake, to large-sized anurans, while part of the diets of some such as its nocturnal activity, semi-arboreal habits, and Helicops species, may be particularly difficult to subdue preference for habitats associated with bodies of water, and immobilise by these snakes, and that their venom bring it into contact with the breeding aggregations of may not always subdue the prey effectively. In general, many anuran species, favouring interactions between then, it seems likely that these snakes will typically prey predator and prey (Bernarde et al., 2000; Marques et al., on the juveniles or naturally debilitated individuals of 2001; Franco and Ferreira, 2002; Ruffato et al., 2003). medium- to large-sized anuran species. In addition to anurans, the diet of T. strigatus includes some fish, reptiles, and mammals, which are usually Thamnodynastes strigatus vs. Physalaemus centralis taken opportunistically (Bernarde et al., 2000; Ruffato in Ouroeste, São Paulo (Event 02): this event occurred et al., 2003; Bortolanza-Filho et al., 2019). between 20:45 h and 20:57 h (duration of approximately The diversity of anuran species in the diet of T. 12 minutes) on 10-29-2017, at the external margin of strigatus (see Bernarde et al., 2000; Ruffato et al., a lake (Fig. 2a) during fieldwork at the study site. The 2003; Bortolanza-Filho et al., 2019) indicates that observations were initiated following the first visual any encounter with P. centralis would normally contact, when it was possible to confirm that the snake favour the predator, although, as the frog maintained had attacked the frog in the region of the throat, after its head stretched backward throughout the whole of which, it had attempted to swallow its prey (Fig. 2b). the encounter, it may have impeded the snake from The position of the attack left the frog in a lateral position swallowing it, which eventually led to the predator in relation to the substrate, with its head stretched abandoning its attack. T. strigatus is an opistoglyphous backward (Fig. 2c). During the attempted predation, the snake, and produces toxins capable of poisoning its frog maintained all four of its members outstretched, prey, which help to subdue it, avoid potentially harmful with its body slightly inflated, without demonstrating reactions, and facilitate ingestion (Bernarde et al., 2000). Unsuccessful predation attempts by snakes on anuran amphibians 653

Given the presence of these toxins, the fact that the frog and hence may explain the non-occurrence of the was abandoned by the snake and appeared to have the predation event. energy to try to escape, did not mean that it would have survived, given that it presented signs of poisoning, Xenopholis undulatus vs. Physalaemus centralis in including blood in its mouth. Observations of the snake União de Minas, Minas Gerais (Event 03): this event also revealed the presence of a discontinuous swelling was recorded between 00:49 h and 01:13 h (duration of in the middle of its body, which appeared to indicate approximately 25 minutes) on 11-06-2017 at the external that it had already ingested a prey earlier that same margin of a temporary pond (Fig. 3a) during fieldwork night. Bernarde et al. (2000) and Ruffato et al. (2003) at the study site. At first visual contact, the body of the both recorded individuals of T. strigatus with more than prey, Physalaemus centralis, was inflated, belly-up, one prey item in their digestive tracts at a similar stage with no signs of resistance, and no vocalisations being of digestion, which indicates that, under favourable emitted (Fig. 3b). The predator (Xenopholis undulatus), conditions, this species may be capable of successfully in turn, was pressing its head against the left flank of attacking a number of preys in a single night. The snake the frog in the vicinity of the inguinal region (Fig. 3b), in this study presented clear signals that it had just fed. in an attempt to swallow its prey. After approximately This may have led the snake to become less propense to seven minutes, the snake moved its mandible in an attack a prey that could impose resistance to predation, attempt to advance over the immobile body of the frog,

Figure 2. Attempted predation of Physalaemus centralis by Thamnodynastes strigatus: (a) general view of the lake at the study site; (b) the snake attempts to swallow its prey; (c) the snake trying to swallow the frog, which maintained its body stretched backward; (d) the snake abandons its prey, which then attempts to escape. 654 William P. Costa & César C. Trevelin although it was unsuccessful, and retreated, appearing presence of relatively large-bodied anurans in the diet to regurgitate the left posterior member of the anuran. of X. undulatus, as observed by Kokubum and Maciel At this moment, it was possible to observe that the frog (2010), Costa et al. (2013) and Guedes (2018), indicates had an open wound in its skin, into which the snake had that the large size of P. centralis would probably not partially introduced its head (Fig. 3c). After regurgitating have been a factor determining the withdrawal of the the posterior left member of the frog as far as the knee snake in the event documented here. It does seem likely, joint, the snake attempted once again to swallow its however, that the attack on the posterior members of the prey, reaching the inguinal region. At this moment, the prey may have reduced its efficiency. body of the frog had deflated, but even so, the snake was An attack on the posterior members of the prey may be unable to swallow any further parts of its body. After hampered by the position of these members in relation regurgitating again, the snake retreated, exposing the to the rest of the body, given the insertion between the whole of the frog’s posterior member, and moved back a inguinal and cloacal regions, which may reinforce the few centimetres (Fig. 3d). The snake then moved around mechanical resistance of the frog, enabling it to avoid the body of the frog from the left side until reaching its being swallowed. In contrast with the event recorded by right posterior member (Fig. 3e). On touching the right Teles et al. (2018), the P. centralis subsequently died, posterior member of the frog, the snake retreated once possibly as a consequence of either poisoning and/or again, and then moved out of sight (Fig. 3f). At the end the lesions suffered during the predation attempt. In this of the event, at 01:13 h, it was possible to examine the specific case, the outcome for both individuals were frog, whose left thigh was swollen and reddened, while negative, given the death of the prey and the wasted part of the right thigh had been affected by the cut in the energy expended by the predator. skin of the inguinal region (Fig. 3f). The genus Xenopholis Peters, 1869 has three species Bothrops moojeni vs. Leptodactylus labyrinthicus (Uetz, 2018), X. undulatus (Jensen, 1900), X. scalaris in Palmeira de Goiás, Goiás (Event 04): this (Wucherer, 1861), and X. werdingorum Jansen, Álvarez event occurred between 20:03 h and 20:33 h, lasting and Köhler, 2009. No systematic data are available on approximately 30 minutes, on 10-28-2018, during the the diet of X. werdingorum, although anurans are the inspection of the area surrounding a forest fragment exclusive or predominant prey of the other two species, at the study site (Fig. 4a). On this occasion, the snake, with species of a number of different families being Bothrops moojeni, was observed striking the frog, targeted (see Cunha and Nascimento, 1993; Strüssmann Leptodactylus labyrinthicus, which, when attempting and Sazima, 1993; Martins and Oliveira, 1998; França to escape, landed in front of the snake with all four and Araújo, 2006; Bernarde and Abe, 2010; Ringler et members stretched backward (Fig. 4b). The snake al., 2010; Silva et al., 2010; Santos-Costa et al., 2015). then remained motionless in the same position for the However, only two predation events involving X. subsequent 30 minutes, only occasionally inspecting it undulatus have been observed in the wild, and only one prey with movements of its tongue (Fig. 4b-d). of these was successful (Kokubum and Maciel, 2010; The Brazilian Lancehead, B. moojeni Hoge, 1966 Teles et al., 2018). In captivity, in addition to anurans, has been recorded widely in inventories of the snakes X. undulatus has been observed preying on the gecko, or herpetofauna of Cerrado savannah localities (e.g. Hemidactylus mabouia (Moreau de Jonnès, 1818) Santos et al., 2014; Silva et al., 2015; Freitas et al., (Costa et al., 2013). Costa et al. (2013) also recorded 2016; Madella-Auricchio et al., 2017; Ramalho et al., prey selectivity in this snake which, when offered 2018; Neves et al., 2019), where its distribution broadly three potential anuran prey–Physalaemus cuvieri overlaps with that of L. labyrinthicus. Even so, this Fitzinger, 1826, Boana albopunctata (Spix, 1824), and is the first time that an interaction between these two Dendropsophus rubicundulus (Reinhardt and Lütken, species has been observed under natural conditions, with 1862)–demonstrated interest only in P. cuvieri, which it an unsuccessful outcome for the snake. Ectothermic preyed on head-first. animals are an important component of the diets of the In addition to the unsuccessful attempt recorded in the pitvipers of the genus Bothrops (see Monteiro et al., present study, involving P. centralis, Teles et al. (2018) 2006; Toledo et al., 2007; França et al., 2008; Palmuti et also recorded an unsuccessful attack of X. undulatus al., 2009; Leão et al., 2014; Bisneto and Kaefer, 2019), on P. cuvieri. In both cases, the prey was attacked in and in the case of B. moojeni, in particular, anurans are the posterior members, in contrast with the successful amply preyed on by juveniles (Andrade et al., 1996; head-first attacks observed by Costa et al. (2013). The Nogueira et al., 2003). Although anurans are known to Unsuccessful predation attempts by snakes on anuran amphibians 655

Figure 3. Attempted predation of Physalaemus centralis by Xenopholis undulatus: (a) general view of the temporary pond at the study site; (b) the predator attempts to swallow its prey; (c) second attack by the snake on the frog; (d) complete regurgitation of the posterior right member of the frog; (e) the snake moves around the frog; (f) the snake abandons its prey.

be a prominent component of the diet of B. moojeni, few 1799) and Boana albopunctata (Spix, 1824), while data are available on the anuran species preyed on by this Moura et al. (2012) also confirmed that its diet includes snake. Even so, França et al. (2008) confirmed that B. Boana crepitans (Wied-Neuwied, 1824). moojeni preys on both Leptodactylus fuscus (Schneider, 656 William P. Costa & César C. Trevelin

L. labyrinthicus has rarely been reported as the prey of Conclusions other animals. Toledo (2003) reported the predation of The events reported here involved snakes that use young individuals by the water bug Belostoma elongatum venom to subdue their prey, although this capacity was Montandon, 1908, while Sazima and Martins (1990) not sufficient to ensure the predation, given that one prey and Prado et al. (2005) register the same interaction was able to escape and the other three were abandoned. predator–prey on adult of L. labyrinthicus. One of the Unsuccessful predation attempts involving snakes and factors that may minimise the wider predation of L. amphibians may be related to the relative sizes of the labyrinthicus is the secretion of unpalatable substances predator and prey, skin secretions, and physical resistance in its skin. These secretions may represent an efficient on the part of the anuran (e.g. Sazima and Martins, 1990; anti-predator defence (Brizzi and Corti, 2007), and may Toledo et al., 2007; Solé et al., 2010; Bernal and Palma, contain substances such as amines, bioactive peptides, 2011; Fong et al., 2013; Vargas-Salinas and Aponte- and alkaloids, all of which may inhibit the behaviour Gutierrez, 2013; Theles et al., 2018; Rocha-Lima et of the predator (Ferreira et al., 2019). Given this, the al., 2018; Yeager et al., 2019). In the specific case of loss of interest in its prey observed in B. moojeni in the L. chaquensis, the combination of the body size of the present study may have been the result of its perception frog and its resistance to predation probably favoured of these substances in the skin of L. labyrinthicus. its escape. In the case of the three abandoned prey, the factors that may have influenced the behaviour of the predator are unclear, and the observation of additional

Figure 4. Attempted predation of Leptodactylus labyrinthicus by Bothrops moojeni: (a) general view of the fragment of forest at the study site; (b) the prey attempts to escape, but falls in front of the snake; (c) and (d) the snake inspects its prey. Unsuccessful predation attempts by snakes on anuran amphibians 657 events under similar conditions would be necessary to unsuccessful predation events, as well as more detailed provide more conclusive insights into the phenomenon. studies of the predatory behaviour and diet of snakes, to However, mainly the results of the present study better comprehend the importance of the predator-prey nevertheless indicate that body size (Event 03) and interaction. the presence of toxins in the skin of the anuran (Event 04) may be important factors, as observed in previous Acknowledgments. We are grateful to the Paulo Sérgio Bernarde studies (see Brizzi and Corti, 2007; Dorado-Rodrigue and Ana Beatriz Carollo Rocha-Lima for critical revision, et al., 2012; Manoel and Almeida, 2017; Oliveira et al., corrections and suggestions on the manuscript; Sônia Cristina da Silva Belentani and José Fernando Pacheco for assistance and 2017; Teles et al., 2018; Ferreira et al., 2019). support during fieldwork. An additional determinant of the effectiveness of a snake attack is the region of the prey’s body affected by References the strike. In an analysis of the diet of T. strigatus, Ruffato et al. (2003) observed that anteroposterior ingestion was Ab’Saber, A.N. (2003): Os domínios de natureza no Brasil: typical of anuran prey, and that only relatively small potencialidades paisagísticas. Ateliê Editorial, São Paulo. Aguiar, L.F.S., Di-Bernardo, M. (2004): Diet and Feeding prey were ingested in the opposite orientation, that Behaviour of Helicops infrataeniatus (Serpentes: Colubridae: is, in a posteroanterior fashion. Records of successful Xenodontinae) in Southern Brazil. Studies on Neotropical predation tend to involve attacks to the head or anterior Fauna and Environment 39(1): 7–14. region of the prey (e.g. Toledo et al., 2007; Moya and Andrade, D.V., Abe, A.S., Dos Santos, M.C. (1996): Is the venom Maffei, 2012; Falkenberg et al., 2013; Santos-Silva et related to diet and tail color during Bothrops moojeni ontogeny? al., 2014; Passos et al., 2017; Muscat and Moroti, 2018; Journal of Herpetology 30: 285–288. Ávila, R.W., Ferreira, V.L., Arruda, J. (2006): Natural history of the Thaler et al., 2018), whereas unsuccessful attempts tend South American water snake Helicops leopardinus (Colubridae: to involve attacks directed at the posterior members of Hydropsini) in the Pantanal, Central Brazil. Journal of the prey (e.g. Bernarde et al., 2000; Rocha and López- Herpetology 40: 274–279. Baucells, 2013; Manoel and Almeida, 2017; Teles et al., Bailey, J.R. (1967): The synthetic approach to Colubrid 2018; Rocha-Lima et al., 2018; Yeager et al., 2019). The classification. Herpetologica 23: 155–161. evidence from the present study further reinforces the Bailey, J.R., Thomas, R.A., Silva, N.J. (2005): A revision of the conclusion that unsuccessful predation is related to the South American snake genus Thamnodynastes Wagler, 1830 (Serpentes, Colubridae, Tachymenini). I. Two new species of region of the body attacked by the snake, although this Thamnodynastes from Central Brazil and adjacent areas, with can only be confirmed more conclusively with a much a redefinition of and neotype designation for Thamnodynastes. larger number of observations of unsuccessful attempts. Phyllomedusa 4: 83–101. One of the greatest challenges for the study of predator- Bellini, G.P., Arzamendia, V., Giraudo, A.R. (2013): Ecology of prey interactions under natural conditions, in particular Thamnodynastes hypoconia in Subtropical-Temperate South in the case of anuran communities, is the difficulty of America. Herpetologica 69: 67–79. Bernal, M.H., Palma, G.M. (2011): Micrurus mipartitus (redtail observing and quantifying predation events (Pombal, coralsnake). Diet. Herpetological Review 42: 617. 2007). In general, these events are observed only very Bernarde, P.S., Abe, A.S. (2010): Hábitos alimentares de serpentes sporadically, and most published reports are no more em Espigão do Oeste, Rondônia, Brasil. Biota Neotropica 10(1): than brief descriptions of the event, focusing primarily 167–173. on successful attacks (e.g. Vargas-Salinas and Aponte- Bernarde, P.S., Moura-Leite, J.C., Machado, R.A., Kokubum, Gutierrez, 2013; Falkenberg et al., 2013; Dorigo et al., M.N.C. (2000): Diet of the colubrid snake, Thamnodynastes strigatus (Günther, 1858) from Paraná state, Brazil, with field 2014; Carrillo 2017). The number of observed events, notes on anuran predation. Revista Brasileira de Biologia 60: even when combined with data on stomach contents (e.g. 695–699. Zanella and Cechin, 2009; Bernarde and Abe, 2010), Bisneto, P.F., Kaefer, I.L. (2019): Reproductive and feeding nevertheless provide only a very limited perspective on biology of the common lancehead Bothrops atrox (Serpentes, the ecological relationship between snakes and anuran Viperidae) from central and southwestern Brazilian Amazonia. amphibians (Marques and Sazima, 2004; Pombal, 2007). Acta Amazonica 49(2): 105–113. Given this, it would be fundamentally important to Bortolanza-Filho, D., Lourenço-de-Moraes, R., Otani1, M., Lemos, G.F., Zawadzki1, C.H. (2019): New records of the dipsadid compile the records of unsuccessful predation attempts, snake Thamnodynastes strigatus (Günther, 1858) preying on in order to provide a more comprehensive perspective the characid fish Astyanax bockmanni Vari & Castro, 2007 in on this type of interaction. In this context, it is important the Atlantic Forest of Southern Brazil. Herpetology Notes 12: to recognise the need for the more ample divulgation of 613–615. 658 William P. Costa & César C. Trevelin

Bovo, R.P., Sueiro, L.R. (2012): Records of predation on Itapotihyla K.H., Brodie Jr., E.D. (2019): Antipredator mechanisms of langsdorffii (Anura: Hylidae) by Chironius bicarinatus post-metamorphic anurans: a global database and classification (Serpentes: Colubridae) with notes on foraging substrate. system. Behavioural Ecology and Sociobiology 73: 769. Herpetology Notes 5: 291–292. Filho, L.M.C., Nascimento, D.G., Argôlo, A.J.S. (2017): Death of Branch, W.R. (1976): Two exceptional food records for the african Oxyrhopus petolarius (Linnaeus, 1758) (Serpentes: Dipsadidae) bullfrog, Pyxicephalus adspersus (Amphibia, Anura, Ranidae). after an unsuccessful predation event on Tropidurus torquatus Journal of Herpetology 10: 266–268. (Wied-Neuwied, 1820) (Sauria: Tropiduridae). Herpetology Brizzi, R., Corti, C. (2007): Cutaneous antipredatory secretions and Notes 10: 71–73. pheromones in anurans and urodeles. Marine and Freshwater Fong, G.A., Bignotte, G.I., Maure, G.K. (2013): Unsuccessful Behaviour and Physiology 40: 225–231. predation on the toad Peltophryne peltocephala (Bufonidae) Camargo Filho, C.B., Costa, H.C., Silva, E.T., Ribeiro Filho, O.P., by the Cuban snake Tropidophis melanurus (Tropidophiidae). Feio, R.N. (2008): Lithobates catesbeianus (American Bullfrog) Herpetology Notes 6: 73–75. Prey. Herpetological Review 39: 338–338. Fonseca, E., Lanna, F., Carvalho, R., Gehara, M. (2012): Predation Caramaschi, U., Niemeyer, H. (2012): Unsuccessful predation on Sibynomorphus neuwiedi (Serpentes: Dipsadidae) by of Elapomorphus quinquelineatus (Serpentes: Colubridae) on Leptodactylus labyrinthicus (Anura: Leptodactylidae) in Amphisbaena microcephala (Amphisbaenia: Amphisbaenidae). southeastern Brazil. Herpetology Notes 5: 167–168. Herpetology Notes 5: 429–430. Forti, L.R., Bertoluci, J. (2012): Distress call of Hypsiboas Cardoso, M.R.D., Marcuzzo, F.F.N., Barros, J.R. (2015): faber (Anura: Hylidae) during a Liophis miliaris (Serpentes: Classificação Climática de Köppen-Geiger para o Estado de Colubridae) attack. Herpetology Notes 5: 187–188. Goiás e o Distrito Federal. Acta Geográfica 8(16): 40–55. França, F.G.R., Araújo, A.F.B. (2006): The conservation status of Carrillo, J.F.C. (2017): Predation of Thamnodynastes chaquensis snakes in central Brazil. South American Journal of Herpetology (Serpentes, Colubridae) upon Elachistocleis matogrosso (Anura, 1(1): 25–36. Microhylidae) in the Brazilian Pantanal. Herpetology Notes 10: França, F.G.R., Mesquita, D.O., Nogueira, C.C., Araújo, A.F.B. 355-357 (2008): Phylogeny and Ecology Determine Morphological Cavalcanti, L.B.Q., Santos-Protázio, A., Albuquerque, R.L., Pedro, Structure in a Snake Assemblage in the Central Brazilian C.K.B., Mesquita, D.O. 2012. Death of a coral snake Micrurus Cerrado. Copeia 2008 (1): 23–38. ibiboboca (Merrem, 1820) (Elapidae) due to failed predation Franco, F.L. (1999): Relações filogenéticas entre os gêneros da tribo on bigger prey: a cat-eyed night snake Leptodeira annulata Tachymenini Bailey, 1967 (Serpentes, Colubridae). Doctoral (Linnaeus, 1758) (Dipsadidae). Herpetology Notes 5: 129–131. dissertation. São Paulo, University of São Paulo. Costa, H.C., Nascimento, M.C., Oliveira, M.C.G. (2013): Franco, F.L., Ferreira, T.G. (2002): Descrição de uma nova espécie Xenopholis undulatus (Serpentes: Xenodontinae) Reprodução e de Thamnodynastes Wagler, 1830 (Serpentes, Colubridae) alimentação em cativeiro. Herpetologia Brasileira 2: 36–38. do nordeste brasileiro, com comentários sobre o gênero. Costa, H.C., Pezzuti, T.L, Leite, F.S.F., Cientifuegos, C. (2009): Phyllomedusa 1(2): 57–74. Helicops modestus (water snake): Prey. Herpetological Bulletim Freitas, M.A., Colli, G.R., Estiuspe-Neto, O.M., Trinchão, L., 109: 35–36. Araújo, D., Lima, T.O., França, D.P.F., Gaiga, R., Dias, P. da Cunha, O.R., Nascimento, F.P. (1993): Ofídios da Amazônia. (2016): Snakes of Cerrado localities in western Bahia, Brazil. As cobras da região leste do Pará. Boletim do Museu Paraense Check List 12(3): 1896. Emílio Goeldi - Zoologia 9(1): 0–191. Guedes, T.B. 2018. Xenopholis undulatus (Jensen’s Groundsnake). Dorigo, T.A., Vrcibradic, D., Borges-Junior, V.N.T., Rocha, C.B.D. Diet. Herpetological Review 49: 142–142. (2014): New records of anuran predation by snakes of the genus Kokubum, M.N.C., Maciel, N.M. (2010): Natural history notes. Thamnodynastes Wagler, 1830 (Colubridae: Dipsadinae) in the: Scinax fuscovarius. Predation. Herpetological Review 41: Atlantic rainforest of southeastern Brazil. Herpetology Notes 7: 480–481. 261–264. Leão, S.M., Pelegrin, N., Nogueira, C.C., Brandão, R.A. (2014): Dorado-Rodrigues, T.F., Campos, V.A., dos Santos M.M., Natural history of Bothrops itapetiningae Boulenger, 1907 Pansonato A., Strüssmann C. (2012): Circumstances and (Serpentes: Viperidae: Crotalinae), an endemic species of the bioacoustics of the distress call of Leptodactylus chaquensis Brazilian Cerrado. Journal of Herpetology 48: 324–331. (Anura: Leptodactylidae) during predation by Thamnodynastes Lema, T., Araújo, M.L., Azevedo, A. (1983): Contribuição ao chaquensis (Serpentes: Dipsadidae) in the Brazilian Pantanal. conhecimento da alimentação e do modo alimentar de serpentes Salamandra 48: 237–240. do Brasil. Comunicação do Museu de Ciência e Tecnologia da Duellman, W.E., Trueb, L. (1986): Biology of amphibians. New Pontifícia Universidade Católica do Rio Grande do Sul, Série York, McGraw-Hill, 670 pp. Zoologia 26: 41–12. Falkenberg, L.M., Protázio, A.S., Albuquerque, R.L., Mesquita, Madella-Auricchio, C.R., Auricchio, P., Soares, E.S. (2017): Reptile D.O. (2013): Predation of Phyllomedusa nordestina (Anura: species composition in the Middle Gurguéia and comparison Hylidae) by Leptodeira annulata (Serpente: Dipsadidae) in a with inventories in the eastern Parnaíba River Basin, State of temporary pond. Herpetology Notes 6: 97–98. Piauí, Brazil. Papéis Avulsos de Zoologia 57: 375–386. Feltrim, A.C., Cechin, S.T.Z. (2000): Helicops infrataeniatus Manoel, P.S., Almeida, S.C. (2017): Predation attempt on the tree (NCN). Diet. Herpetological Review 31: 46. frog Hypsiboas faber (Wied- Neuwied, 1821) by the snake Ferreira, R.B., Lourenço-de-Moraes, R., Zocca, C., Duca, C., Beard, Thamnodynastes hypoconia (Cope, 1860). Herpetology Notes Unsuccessful predation attempts by snakes on anuran amphibians 659

10: 433–434. by the colubrid snake Chironius flavolineatus (Jan, 1863) in a Mario-da-Rosa, C., Abegg, A.D., Malta-Borges, L., Righi, A.F., montane rocky grassland. Revista Brasileira de Zoociências Bernarde, P.S., Cechin, S.Z., Santos, T.G. (2020): A fisherman’s 18(1): 47–52. tale: Activity, habitat use and the first evidence of lingual lure Pombal Jr., J.P. (2007): Notas sobre predação em uma taxocenose behavior in a South American snake. Salamandra 56 (1): 39–47. de anfíbios anuros no Sudeste do Brasil. Revista Brasileira de Marques, O.A.V., Eterovic, A., Sazima, I. (2001): Serpentes da Zoologi 24(3): 841–843. Mata Atlântica - guia ilustrado para a Serra do Mar. Ribeirão Prado, C.P.A., Toledo, L.F., Zina, J., Haddad, C.F.B. (2005): Preto. Holos. 184 pp. Trophic eggs in the foam nests of Leptodactylus labyrinthicus Marques, O.A.V., Sazima I. (2004): História natural dos répteis da (Anura, Leptodactylidae): an experimental approach. The Estação Ecológica Juréia-Itatins, p. 257-277. In: O.A.V. Marques Herpetological Journal 15(4): 279–284. & W. Dulepa (Eds). Estação Ecológica Juréia-Itatins. Ambientes Preuss, J.F., Tozetti1, A.M. (2018): A record of predation and físico, flora e fauna. Ribeirão Preto, Holos, 384p. ingestion of Phyllomedusa tetraploidea (Anura, Hylidae) by Martins, I.A., Duarte, M.R. (2003): Physalaemus nattereri. Thamnodynastes strigatus (Serpentes, Dipsadidae), in the Predation. Herpetological Review 34: 233. municipality of São Miguel do Oeste, state of Santa Catarina, Martins, M., Oliveira, M.E. (1998): Natural history of snakes Brazil. Herpetology Notes 11: 945–947. in forests of the Manaus region, Central Amazonia, Brazil. Ramalho, W.P., Silva, J.R., Soares, P.T., Ferraz, D., Arruda, Herpetological Natural History 6(2): 78–150. F.V., Prado, V.H.M. (2018): The anurans and squamates of a Martins, M., Sazima, I., Engler, S.G. (1993): Predators of the nest periurban Cerrado remnant in the State of Goiás, Central Brazil. building gladiator frog, Hyla faber, in southeastern Brazil. Herpetology Notes 11: 573–583. Amphibia-Reptilia 14: 307–309. Reboita, M.S., Rodrigues, M., Silva, L.F., Alves, M.A. (2015): Monteiro, C, Montgomery, C.E., Spina, F., Sawaia, R.J., Martins, Aspectos climáticos do estado de Minas Gerais. Revista M. (2006): Feeding, Reproduction, and Morphology of Brasileira de Climatologia 17: 206–226. Bothrops mattogrossensis (Serpentes, Viperidae, Crotalinae) in Ringler, M., Ursprung, E., Hӧdl, W. (2010): Predation on Allobates the Brazilian Pantanal. Journal of Herpetology 40: 408–413. femoralis (Boulenger 1884; Anura: Arombatidae) by the colubrid Moura, M.R., Godinho, L.B., Feio, F.N. (2012): Bothrops moojeni snake Xenopholis scalaris (Wucherer 1861). Herpetology Notes (Squamata, Viperidae) predation on Hypsiboas crepitans 3: 201–304. (Anura: Hylidae) in southeastern Brazil. Herpetology Notes 5: Rocha, R., López-Baucells, A. (2013): Predation attempt of 247–248. Hypsiboas boans (Anura: Hylidae) by Helicops angulatus Moya, G.M., Maffei, F. (2012): Predation on Dendropsophus (Squamata: Dipsadidae) with notes on defensive behaviour. elianeae (Napoli & Caramaschi, 2000) (Anura: Hylidae) Alytes 30(1): 78–81. by Thamnodynastes hypoconia (Cope, 1860) (Squamata: Rocha-Lima, A.B.C., Santos, I., Suigh, L., Duarte, C., Costa, Colubridae). Herpetology Notes 5: 261–262. W.P. (2018): Erythrolamprus miliaris orinus (Reptilia, Muscat, E., Moroti, M.T. (2018): Predation of Rhinella ornata Squamata, Dipsadidae): tentativas de predação de Boana faber (Anura: Bufonidae) by the water snake Erythrolamprus miliaris e Leptodactylus latrans. Boletim do Museu Paraense Emílio (Squamata: Dipsadidae) in São Paulo, Brazil. Herpetol Notes Goeldi - Ciências Naturais 13(3): 455–460. 11: 449–450. Rolim, G. de S., Camargo, M.B.P.de, Lania, D.G., Moraes, J.F.L.de. Neves, M.O., Yves, A., Pereira, E.A., Alves, L., Vasques, J.B., (2007): Classificação climática de Köppen e de Thornthwaite e Coelho, J.F.T., Silva, P.S. (2019): Herpetofauna in a highly sua aplicabilidade na determinação de zonas agroclimáticas para endangered area: the Triângulo Mineiro region, in Minas Gerais o Estado de São Paulo. Bragantia 66: 711–720. State, Brazil. Herpetozoa 32: 113–123. Ruffato, R., Di-Bernardo, M., Maschio, G.F. (2003): Dieta de Nogueira, C., Sawaya, R.J., Martins, M. (2003): Ecology of Thamnodynastes strigatus (Serpentes: Colubridae) no sul do Bothrops moojeni (Serpentes: Viperidae: Crotalinae) in the Brasil. Phyllomedusa 2(1): 27–34. Brazilian Cerrado. Journal of Herpetology 37(4): 653–659. Santos, D.L., Andrade, S.P., Victor-Jr., E.P., Vaz-Silva, W. (2014): Nogueira, C.H.O., Figueiredo-de-Andrade, C.A., Freitas, N.N. Amphibians and reptiles from southeastern Goiás, Central (2013): Death of a juvenile snake Oxyrhopus petolarius Brazil. Check List 10(1): 131–148. (Linnaeus, 1758) after eating an adult house gecko Hemidactylus Santos-Costa, M.C., Maschio, G.F., Prudente, A.L.C. (2015): mabouia (Moreau de Jonnès, 1818). Herpetology Notes 6: 39– Natural history of snakes from Floresta Nacional de Caxiuanã, 43. eastern Amazonia, Brazil. Herpetology Notes 8: 69–98. Oliveira, S., Silva, D., Fachi, M., Morais, A.R. (2017): Predation on Santos-Silva, C.R., Andrade, I.S., Araújo, M.L.N., Barros, L.C.S., Rhinella mirandaribeiroi (Gallardo, 1965) (Anura; Bufonidae) Gomes, L., Ferrari, S.F. (2014): Predation of six anuran species by a snake from Central Brazil. Herpetology Notes 10: 151– by the banded cat-eyed snake, Leptodeira annulata (Serpentes: 155. Dipsadidae), in the Caatinga scrub of northeastern Bahia, Brazil. Palmuti, C.F.S., Cassimiro, J., Bertoluci, J. (2009): Food habits of Herpetology Notes 7: 123–126. snakes from the RPPN Feliciano Miguel Abdala, an Atlantic Sawaya, R.J., Marques, O.A.V., Martins, M. (2008): Composição e Forest fragment of southeastern Brazil. Biota Neotropica 9(1): história natural das serpentes de Cerrado de Itirapina, São Paulo, 263–269. sudeste do Brasil. Biota Neotropica 8(2): 127–149. Passos, D.C., Glauss, L.H.A., Galdino, C.A.B. (2017): Predation of Sazima, I., Martins, M. (1990). Presas grandes e serpentes jovens; the hylid frog Bokermannohyla alvarengai (Bokermann, 1956) quando os olhos são maiores que a boca. Memórias do Instituto 660 William P. Costa & César C. Trevelin

Butantan 52(3): 73–39. Toledo, L.F., Ribeiro, R.S., Haddad, C.F.B. (2007): Anurans as Scartozzoni, R.R. (2005): Morfologia de serpentes aquáticas prey: an exploratory analysis and size relationships between neotropicais: um estudo comparativo. Masters thesis, Institute predators and their prey. Journal of Zoology 271: 170–177. of Biosciences, University of São Paulo, São Paulo. 102 pp. Uetz, P., Freed, P., Hošek J. (eds.). (2019): The Reptile Database, Scartozzoni, R.R. (2009): Estratégias reprodutivas e ecologia http://www.reptile-database.org, accessed on 08/08/2019. alimentar de serpentes aquáticas da tribo Hydropsini Vargas-Salinas, F., Aponte-Gutierrez, A. (2013): A race for (Dipsadidae, Xenodontinae). Doctoral dissertation, Interunit survivorship: failed predation on the toad Rhinella humboldti Program in Biotechnology, Butantan Institute and the University (Gallardo, 1965) by the cat-eyed snake Leptodeira septentrionalis of São Paulo, São Paulo. 161 pp. (Kennicott, 1859). Herpetology Notes 6: 189–191. Silva, E.T., Costa, H.C., Feio, R.N. (2007). Rana catesbeiana Vidal, N., Kindl, S.G., Wong, A., Hedges, S.B. (2000): Phylogenetic (American Bullfrog). Prey. Herpetological Review 38: 443– relationships of xenodontine snakes inferred from 12s and 443. 16s ribosomal RNA sequences. Molecular Phylogenetics and Silva, M.C., Oliveira, R.H., Morais, D.H., Kawashita-Ribeiro, Evolution 14: 389–402. R.A., Brito, E.S., Ávila, R.W. (2015): Amphibians and reptiles Warkentin, K.M. (2011): Plasticity of hatching in amphibians: of a Cerrado area in Primavera do Leste municipality, Mato evolution, trade-offs, cues and mechanisms. Integrative and Grosso State, Central Brazil. Salamandra 51(2): 187–194. Comparative Biology 51: 111–127. Silva, M.V., Souza, M.B., Bernarde, P.S. (2010). Riqueza e dieta Wells, K.D. (2007): Ecology and Behaviour of Amphibians. de serpentes do Estado do Acre, Brasil. Revista Brasileira de Chicago and London, The University of Chicago Press. 1400 Zoociências 12(2): 55–66. pp. Solé, M., Marciano-Jr., E., Dias, I.R., Kwet, A. (2010): Predation Winkler, F.J.M.,Waltenberg, L.M., Almeida-Santos, P., attempts on Trachycephalus cf. mesophaeus (Hylidae) by Nascimento, D.S., Vrcibradic, D., Sluys, M.V. (2011): New Leptophis ahaetulla (Colubridae) and Ceratophrys aurita records of anuran prey for Thamnodynastes strigatus (Günther, (Ceratophryidae). Salamandra 46(2): 101–103. 1858) (Serpentes: Colubridae) in a high-elevation area of Strüssmann, C., Sazima, I. (1993): The snake assemblage of southeast Brazil. Herpetology Notes 4: 123–124. the pantanal at Poconé, western Brazil: faunal composition Yeager, J., Zarling, A., Rodríguez, C. (2019): Successful multimodal and ecological summary. Studies on Neotropical Fauna and amphibian defence in the neotropical frog Trachycephalus, Environmental 28: 157–168. including handling and recovery costs to would-be predators. Teixeira, C.C., Montag, L.F.A., Santos-Costa, M.C. (2017): Diet Herpetology Notes 12: 279–280. Composition and Foraging Habitat Use by Three Species of Zaher, H. (1999): Hemipenial morphology of the South American Water Snakes, Helicops Wagler, 1830, (Serpentes: Dipsadidae) xenodontine snakes, with a proposal for a monophyletic in Eastern Brazilian Amazonia. Journal of Herpetology 51(2): Xenodontinae and a reappraisal of colubroid hemipenes. Bulletin 215–222. of the American Museum of Natural History 240:1–168. Teles, A., Sena, A., Ribeiro, M.V. (2018): Predation attempt of Zaher, H., Grazziotin, F.G., Cadle, J.E., Murphy, R.W., Moura- Xenopholis undulatus (Serpentes, Dipsadidae) on Physalaemus Leite, J.C.D., Bonatto, S.L. (2009): Molecular phylogeny of cuvieri (Amphibia, Leptodactylidae). Herpetology Notes 11: advanced snakes (Serpentes, Caenophidia) with an emphasis 829–830. on South American Xenodontines: a revised classification Thaler, R., Folly, H., Galvão, C., Silva, L.A. (2018): First predation and descriptions of new taxa. Papéis Avulsos de Zoologia 49: report of Leptodactylus chaquensis (Anura, Leptodactylidae) by 115–153. Helicops infrataeniatus (Squamata, Dipsadidae) and distribution Zanella, N., Cechin, S.Z. (2009): Influência dos fatores abióticos e extension for this water snake in the Brazilian Cerrado ecoregion. da disponibilidade de presas sobre comunidade de serpentes do Herpetology Notes 11: 539–541. Planalto Médio do Rio Grande do Sul. Iheringia, Serie Zoologia Toledo, L.F. (2003): Predation on seven south american anuran 99(1): 111–114. species by water bugs (Belostomatidae). Phyllomedusa 2(2): 105–108.

Accepted by Mirco Solé