Abstract
Specimens (50) of Leptophis nigromarginatus from northern Brazil, Peru, Ecuador and Colombia were examined for the composition of stomach contents. Most prey items were tree frogs, especially those of the family Hylidae (96%). Most of the anurans identified belong to the Scinax ruber species group (14%) and Scinax sp. (8%). Both sexes preyed on small-sized items (1.8–4.4% of snake snout-vent length), but females capture larger prey than males. Leptophis nigromarginatus seems to manipulate captured prey before ingestion since most of the prey items (68%) were swallowed head-first. There is a positive correlation between female snout-vent length and prey size and between and head length and prey size.
Keywords
Diet; Feeding behavior; Anura; Parrot snake; Leptophis nigromarginatus
Resumo
Espécimes (50) de Leptophis nigromarginatus do Norte do Brasil, Peru, Equador e Colômbia foram examinados quanto à composição do conteúdo estomacal. A maioria das presas eram pererecas, especialmente as da família Hylidae (96%). A maioria dos anuros identificados pertence ao grupo de espécies Scinax ruber (14%) e Scinax sp. (8%). Ambos os sexos predaram itens de pequeno porte (1,8-4,4% do comprimento focinho-cloaca da serpente), mas as fêmeas capturam presas maiores que os machos. Leptophis nigromarginatus parece manipular as presas capturadas antes da ingestão, uma vez que a maioria das presas (68%) foi engolida pela cabeça. Há uma correlação positiva entre o comprimento do focinho-cloaca da fêmea e o tamanho da presa e entre o comprimento da cabeça e o tamanho da presa.
Palavras-chave
Dieta; Comportamento alimentar; Anura; Cobra-papagaio; Leptophis nigromarginatus
Introduction
The Neotropical genus Leptophis Bell, 1825 (parrot snakes) comprises a group of 19 colubrine snakes widely distributed from Mexico through Central- and South America (Uetz et al. 2024UETZ, P., FREED, P. & HOŠEK, J. (Eds.) 2024. The Reptile Database. http://www.reptile-database.org (last access in 16 April 2024).
http://www.reptile-database.org...
). These diurnal serpents inhabit both arboreal and terrestrial environments, and are known to feed predominantly on hylid frogs (e.g., Beebe 1946, Oliver 1948OLIVER, J.A. 1948. The relationships and zoogeography of the genus Thalerophis Oliver. Bull. Am. Mus. Nat. Hist. 92(4):157–280., Albuquerque & Di-Bernardo 2005ALBUQUERQUE, N.R. & DI-BERNARDO, M. 2005. Leptophis ahaetulla marginatus. Diet. Herpetol. Rev. 36(3):325.). However, Oliver (1948)OLIVER, J.A. 1948. The relationships and zoogeography of the genus Thalerophis Oliver. Bull. Am. Mus. Nat. Hist. 92(4):157–280., Hero & Magnusson (1987)HERO, J.-M. & MAGNUSSON, W. 1987. Leptophis ahaetulla. Natural history note. Herpetol. Rev. 18(1):16., Teixeira & Porto (1991)TEIXEIRA, D.M. & PORTO, M. 1991. Leptophis ahaetulla. Natural history note. Herpetol. Rev. 22(4):132., Martins & Oliveira (1999)MARTINS, M. & OLIVEIRA, M.E. 1999. Natural history of snakes in forests of the Manaus region, Central Amazonia, Brazil. Herpetol. Nat. Hist. 6(2):78–150., Albuquerque et al. (2007)ALBUQUERQUE, N.R., GALATTI, U. & DI-BERNARDO, M. 2007. Diet and feeding behaviour of the Neotropical parrotsnake (Leptophis ahaetulla) in Northern Brazil. J. Nat. Hist. 41(17–20):1237–1243. https://doi.org/10.1080/00222930701400954.
https://doi.org/10.1080/0022293070140095...
, Muniz et al. (2013)MUNIZ, S.L.S, MOURA, C.C.M., VEJA, S.F., COUTO, A.A., SILVA, J.S., SANTOS, E.M. & MOURA, G.J.B. 2013. Leptophis ahaetulla marginatus (Swordsnake). Diet. Herpetol. Rev. 44(1):154. and Leite et al. (2022)LEITE, A.K., FAZOLATO, C.P., OLIVEIRA, M.L.T., SILVA, R.S. & AMORIM, P.P.S. 2022. Leptophis ahaetulla marginatus (Southern Green Parrot-Snake). Diet. Herpetol. Rev. 53(1):153. also recorded lizards, snakes, young birds and salamanders in their diet.
Among the currently recognized taxa in the genus, Leptophis nigromarginatus (Günther, 1866) is a diurnal and arboreal snake (Dixon & Soini 1977DIXON, J.R. & SOINI, P. 1977. The Reptiles of the Upper Amazon Basin, Iquitos Region, Peru. Part 1. Lizards and Amphisbaenians. Part 2. Crocodilians, Turtles and Snakes. 1 ed. Milwaukee Public Museum, Contributions in Biology and Geology 12:1–91., Duellman 1978DUELLMAN, W.E. 1978. The biology of an Equatorial herpetofauna in Amazonian Ecuador. Misc. publ. Univ. Kans. Mus. Nat. Hist. 65:1–352.) that can also be found on shrubs and banana plants (Duellman 2005DUELLMAN, W.E. 2005. Cusco Amazónico: The Lives of Amphibians and Reptiles in an Amazonian Rainforest. Cornell University Press, Ithaca.). It is a medium-sized parrot snake, with snout-vent length usually not larger than 835 mm, known from Guyana, the western- and middle Amazon regions of Brazil, the Amazonian lowlands of Colombia, Ecuador, and Peru, and extreme northern Bolivia (Albuquerque & Fernandes 2022ALBUQUERQUE, N.R. & FERNANDES, D.S. 2022. Taxonomic revision of the parrot snake Leptophis ahaetulla (Serpentes, Colubridae). Zootaxa 5153(1):1–69. https://doi.org/10.11646/zootaxa.5153.1.1.
https://doi.org/10.11646/zootaxa.5153.1....
). Although general accounts state that this species feeds on ‘bird eggs, geckos, hylid frogs and young birds’ (Oliver 1948OLIVER, J.A. 1948. The relationships and zoogeography of the genus Thalerophis Oliver. Bull. Am. Mus. Nat. Hist. 92(4):157–280.) some basic aspects of its natural history, including its dietary habits, are still little-known. To date, they are known to consume the following species of frogs: Scinax ruber (Laurenti, 1768) and Osteocephalus alboguttatus (Boulenger, 1882) (Dixon & Soini 1977DIXON, J.R. & SOINI, P. 1977. The Reptiles of the Upper Amazon Basin, Iquitos Region, Peru. Part 1. Lizards and Amphisbaenians. Part 2. Crocodilians, Turtles and Snakes. 1 ed. Milwaukee Public Museum, Contributions in Biology and Geology 12:1–91., Duellman 1978DUELLMAN, W.E. 1978. The biology of an Equatorial herpetofauna in Amazonian Ecuador. Misc. publ. Univ. Kans. Mus. Nat. Hist. 65:1–352., 2005DUELLMAN, W.E. 2005. Cusco Amazónico: The Lives of Amphibians and Reptiles in an Amazonian Rainforest. Cornell University Press, Ithaca.). Nothing else is known about diet in this species.
The goal of this paper is to report the feeding habits of L. nigromarginatus addressing also the following question: Are there differences in the size of prey items ingested by males and females of L. nigromarginatus?
Material and Methods
We analyzed 50 adult specimens of L. nigromarginatus for stomach contents (Appendix I). The snakes are housed in eight herpetological collections and institutional abbreviations follow Sabaj (2023)SABAJ, M.H. 2023. Codes for Natural History Collections in Ichthyology and Herpetology (online supplement). Version 9.5 (10 Nov 2023). Available from: https://asih.org, American Society of Ichthyologists and Herpetologists, Washington, DC. https://doi.org/10.1643/ASIHCODONS2020.
https://asih.org...
. We obtained most of the analyzed stomach contents from snakes collected in northern Brazil, Peru, Ecuador and Colombia. Most of the gut contents were already exposed in the specimens examined. Otherwise, in some cases we made a small incision in the stomach of snake and removed all intact or partially digested prey items for further examination. For the intact prey items, we measured the length (in mm) using Mitutoyo digital callipers (±0.01 mm). We also recorded the direction of ingestion of prey (inferred from orientation in the gut i.e., head-first or tail-first) and the snout-vent length (SVL) of intact prey.
We recorded the following data for each snake: head length (HL), snout-vent length (SVL), tail length (TL) and condition of the tail tip (mutilated or not).
We determined the sex of all snakes by dissecting the base of the tail, and by inspection of gonads. We used linear regression to test for the correlation between prey length versus predator SVL and between prey length versus predator HL, using sex as factor.
We evaluated the assumptions of normality and homoscedasticity using Kolmogorov–Smirnov’s test and the Levene’s test, respectively (Ayres et al. 2007AYRES, M., AYRES JR., M., AYRES, D.L. & SANTOS, A.S. 2007. BioEstat 5.0: Aplicações Estatísticas nas Áreas das Ciências Biológicas e Médicas. Instituto de Desenvolvimento Sustentável Mamirauá-IDSM/MCT/CNPq, Brasília.). Prey items (n = 19) whose length could not be calculated due to lack of measurement data (i.e., fragmented prey) were excluded from the regression analyses. According to Albuquerque & Fernandes (2022)ALBUQUERQUE, N.R. & FERNANDES, D.S. 2022. Taxonomic revision of the parrot snake Leptophis ahaetulla (Serpentes, Colubridae). Zootaxa 5153(1):1–69. https://doi.org/10.11646/zootaxa.5153.1.1.
https://doi.org/10.11646/zootaxa.5153.1....
, there is no significant difference between the TL and the SVL of females and males in L. nigromarginatus.
Results
Table 1 summarizes prey taxa exploited by Leptophis nigromarginatus in the samples. Anurans were the only prey items found, including nine species representing seven genera (Boana, Callimedusa, Dendropsophus, Hemiphractus, Pristimantis, Scinax, and Sphaenorhynchus) of the families Hemiphractidae, Hylidae and Strabomantidae.
Summary of prey taxa in the diet of Leptophis nigromarginatus from northern Brazil, Peru, Ecuador and Colombia.
The most frequent prey items were frogs of the species Scinax ruber (14%) and some predation events occurred in the Andean regions (Figure 1).Thirty-six of the 50 specimens (72%) contained a single prey item. Eight (16%) had two prey items in their stomachs. One (2%) had three and two (4%) had four prey items. Three (6%) of the 50 specimens (TCWC 44090, TCWC 44660 and TCWC 44661) contained frog eggs.
Geographic distribution of Anuran predation by Leptophis nigromarginatus. Map depicts locations (green circles) where L. nigromarginatus predated anurans.
Forty-three prey items (68%) were consumed head-first and nineteen (30%) were consumed tail-first. The direction of ingestion could not be determined for one prey (2%).
Thirteen specimens (46.4%, n = 28) had mutilated tails. Prey lengths ranged from 1.8 to 3.8% of the snake SVL in males (n = 4) and from 1.8 to 4.4% in females (n = 9).
Prey size was not significantly related to SVL (F = 0.9979, P = 0.6593) not to HL (F = 0.9962, P = 0.6588) in males (Figure 2). On the other hand, the linear regression show that ingested prey size increased significantly with both SVL (F = 11.9937, P = 0.0021) and HL (F = 5.5206, P = 0.0246) of females (Figure 3).
Correlation between Leptophis nigromarginatus (A) snout-vent length and prey length and between (B) head length and prey length in males. Values in mm.
Correlation between Leptophis nigromarginatus (A) snout-vent length and prey length and between (B) head length and prey length in females. Values in mm.
Discussion
Our data revealed that Leptophis nigromarginatus specializes on anuran prey (100 % of all prey items), supporting the general consensus of anecdotal reports found in the literature (Oliver 1948OLIVER, J.A. 1948. The relationships and zoogeography of the genus Thalerophis Oliver. Bull. Am. Mus. Nat. Hist. 92(4):157–280., Dixon & Soini 1977DIXON, J.R. & SOINI, P. 1977. The Reptiles of the Upper Amazon Basin, Iquitos Region, Peru. Part 1. Lizards and Amphisbaenians. Part 2. Crocodilians, Turtles and Snakes. 1 ed. Milwaukee Public Museum, Contributions in Biology and Geology 12:1–91., Duellman 1978DUELLMAN, W.E. 1978. The biology of an Equatorial herpetofauna in Amazonian Ecuador. Misc. publ. Univ. Kans. Mus. Nat. Hist. 65:1–352., 2005DUELLMAN, W.E. 2005. Cusco Amazónico: The Lives of Amphibians and Reptiles in an Amazonian Rainforest. Cornell University Press, Ithaca.) and providing additional information on feeding biology. Most prey items were treefrogs of the genus Scinax, followed by Scinax sp., Boana geographica and Sphaenorhynchus lacteus. Boana cinerascens, Callimedusa tomopterna, Dendropsophus parviceps, D. triangulum, Hemiphractus and S. cruentomma are equally represented in the diet. Most of these items demonstrate that hylid frogs constituted the primary prey source for L. nigromarginatus. The new dietary records provided for L. nigromarginatus include seven hylid, one eleutherodactylid, and one hemiphractylid taxa (Table 1).
The species of Scinax are among the most frequently encountered hylid frogs in the Amazon Basin (Duellman & Wiens 1993DUELLMAN, W.E. & WIENS, J.J. 1993. Hylid frogs of the genus Scinax Wagler, 1830, in Amazonian Ecuador and Peru. Occas. Pap. Mus. Nat. Hist. 153:1–57.), and have also been recorded as the major prey for other semi-arboreal snakes such as Chironius exoletus (Linnaeus, 1758) (Dixon et al. 1993DIXON, J.R., WIEST JR, J.A. & CEI, J.M. 1993. Revision of the Neotropical snake genus Chironius Fitzinger (Serpentes, Colubridae). Boll. Mus. Regionale. Sci. Nat. Torino (IPNI), Monografia 14:1–949.), Mesotes strigatus (Günther, 1858) (Bernarde et al. 2000BERNARDE, P.S., MOURA-LEITE, J.C., MACHADO, R.A. & KOKOBUM, M.N.C. 2000. Diet of the colubrid snake, Thamnodynastes strigatus (Günther, 1858) from Paraná State, Brazil, with field notes on anuran predation. R. Bras. Biol. 60(4):695–699.), Leptophis ahaetulla (then the nominotypical subspecies) (Albuquerque et al. 2007ALBUQUERQUE, N.R., GALATTI, U. & DI-BERNARDO, M. 2007. Diet and feeding behaviour of the Neotropical parrotsnake (Leptophis ahaetulla) in Northern Brazil. J. Nat. Hist. 41(17–20):1237–1243. https://doi.org/10.1080/00222930701400954.
https://doi.org/10.1080/0022293070140095...
) and L. marginatus (Lopez et al. 2003LOPEZ, M.S., GIRAUDO, A.R. & ARZAMENDIA, V. 2003. Leptophis ahaetulla marginatus (Southern Green Parrot-Snake). Diet. Herpetol. Rev. 34(1):68–69.).
Although not found in the present study, a specimen of L. nigromarginatus was observed and photographed by William W. Lamar (pers. comm., 1 April 2021) in situ in the act of consuming an anuran of the species Osteocephalus taurinus Steindachner, 1862 at San Andrés, Río Momón, Loreto, Peru (Figure 4), whereas Duellman (1978DUELLMAN, W.E. 1978. The biology of an Equatorial herpetofauna in Amazonian Ecuador. Misc. publ. Univ. Kans. Mus. Nat. Hist. 65:1–352., 2005DUELLMAN, W.E. 2005. Cusco Amazónico: The Lives of Amphibians and Reptiles in an Amazonian Rainforest. Cornell University Press, Ithaca.) recorded O. alboguttatus (Boulenger, 1882) in their diet.
A specimen of Leptophis nigromarginatus in the act of consuming an anuran of the species Osteocephalus taurinus at San Andrés, Río Momón, Loreto, Peru. Photo by William W. Lamar.
The high frequency of mutilated tails in L. nigromarginatus was also observed in other snakes of the L. ahaetulla complex (see Albuquerque & Fernandes 2022ALBUQUERQUE, N.R. & FERNANDES, D.S. 2022. Taxonomic revision of the parrot snake Leptophis ahaetulla (Serpentes, Colubridae). Zootaxa 5153(1):1–69. https://doi.org/10.11646/zootaxa.5153.1.1.
https://doi.org/10.11646/zootaxa.5153.1....
). This condition appears to be most frequently found in diurnal and arboreal snake species (Moura et al. 2023MOURA, M.R., COSTA, H.C., ABEGG, A.D., ALAMINOS, E., ANGARITA-SIERRA, T., AZEVEDO, W.S., CABRAL, H., CARVALHO, P., CECHIN, S., CITELI, N., DOURADO, A.C.M., DUARTE, A.F.V., FRANÇA, F.G.R., FREIRE, E.M.X., GARCIA, P.C.A., MOL, R., MONTERO, R., MORAES-DA-SILVA, A., PASSOS, D.C., … GUEDES, J.J.M. 2023. Unwrapping broken tails: Biological and environmental correlates of predation pressure in limbless reptiles. J. Anim. Ecol. 1–14. https://doi.org/10.1111/1365-2656.13793.
https://doi.org/10.1111/1365-2656.13793...
). According to some authors, it can be related to predator evasion (Hoogmoed & Ávila-Pires 2011HOOGMOED, M.S. & AVILA-PIRES, T.C. 2011. A case of voluntary tail autotomy in the snake Dendrophidion dendrophis (Schlegel, 1837) (Reptilia: Squamata: Colubridae). Bol. Mus. Para. Emílio Goeldi, Ciências Naturais 6:113–117., Lockhart & Amiel 2011LOCKHART, J.S. & AMIEL, J. 2011. Nerodia sipedon (Northern Watersnake). Defensive behavior. Herpetol. Rev. 42(2):296–297.).
The utilization of different-sized prey by males and females in species showing sexual size dimorphism may decrease competition for food between sexes (e.g., Shine 1987SHINE, R. 1987. Ecological comparisons of island and mainland populations of Australian tiger snakes (Notechis: Elapidae). Herpetol. 43(2):233–240., Camilleri & Shine 1990CAMILLERI, C. & SHINE, R. 1990. Sexual dimorphism and dietary divergence: differences in trophic morphology between male and female snakes. Copeia (1990)3:649–658., Forsman 1995FORSMAN, A. 1995. Heating rates and body temperature variation in melanistic and zigzag Vipera berus: does colour make a difference? Ann. Zool. Fenn. 32:365–374., but see Shine & Goiran 2021SHINE, R. & GOIRAN, C. 2021. Sexual dimorphism in size and shape of the head in the sea snake Emydocephalus annulatus (Hydrophiinae, Elapidae). Sci. Rep. 11, 20026 (2021): https://doi.org/10.1038/s41598-021-99113-2.
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). However, this appears not to be the case, since there were no significant differences between sexes in mean values of SVL and TL (Albuquerque & Fernandes 2022ALBUQUERQUE, N.R. & FERNANDES, D.S. 2022. Taxonomic revision of the parrot snake Leptophis ahaetulla (Serpentes, Colubridae). Zootaxa 5153(1):1–69. https://doi.org/10.11646/zootaxa.5153.1.1.
https://doi.org/10.11646/zootaxa.5153.1....
).
Our results suggest that, as in L. ahaetulla (see Albuquerque et al. 2007ALBUQUERQUE, N.R., GALATTI, U. & DI-BERNARDO, M. 2007. Diet and feeding behaviour of the Neotropical parrotsnake (Leptophis ahaetulla) in Northern Brazil. J. Nat. Hist. 41(17–20):1237–1243. https://doi.org/10.1080/00222930701400954.
https://doi.org/10.1080/0022293070140095...
), L. nigromarginatus seems to manipulate captured prey before ingestion because 68% of prey items were consumed head-first. Furthermore, consumption of frog eggs not only suggests visits to multiple nests, but also may reflect the occurrence of L. nigromarginatus in both ground-level and arboreal habitats.
In conclusion, it seems highly desirable to conduct further studies to better understand whether L. nigromarginatus shares food resources in areas of sympatry with L. ahaetulla, a species also specialized in eating hylid frogs (Albuquerque et al. 2007ALBUQUERQUE, N.R., GALATTI, U. & DI-BERNARDO, M. 2007. Diet and feeding behaviour of the Neotropical parrotsnake (Leptophis ahaetulla) in Northern Brazil. J. Nat. Hist. 41(17–20):1237–1243. https://doi.org/10.1080/00222930701400954.
https://doi.org/10.1080/0022293070140095...
) and to verify why females consume larger prey than males.
Acknowledgments
This article began in an earlier form as NA’s Doctoral thesis at the PUCRS, in Brazil. We thank the curators – some now retired – of the following collections for loan of specimens: D. Frost and D. Kizirian (AMNH), Colin McCarthy (BMNH), A. Resetar (FMNH), J.E. Simmons (KU), C.C. Austin (LSUMZ), J.A. McGuire (MVZ), T. Hibbitts (TCWC), R.W. McDiarmid and S. Gotte (USNM). NA thanks his advisors, M. Di Bernardo (In memoriam) and T. de Lema (In memoriam) for their support and encouragement throughout his graduate studies. NA is indebted to D.R. Frost, D. Kizirian as well as the whole team of the Department of Herpetology at the AMNH for the opportunity to develop part of his thesis under their supervision and support. NA also wishes to express his gratitude to R.W. McDiarmid and Julian Faivovich who identified some of the prey items. We thank W.W. Lamar and an anonymous reviewer for their constructive comments, which helped us to improve the manuscript. The FMNH granted to NA a visiting scholarship. CAPES provided NA with a fellowship during his time as a doctoral student. CNPq provided RM with a fellowship during his time as a graduate student. This study was financed in part by the Universidade Federal de Mato Grosso do Sul, Ministério da Educação, Brazil.
Data Availability
The entire dataset supporting the results of this study was published in the article itself.
References
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- ALBUQUERQUE, N.R. & FERNANDES, D.S. 2022. Taxonomic revision of the parrot snake Leptophis ahaetulla (Serpentes, Colubridae). Zootaxa 5153(1):1–69. https://doi.org/10.11646/zootaxa.5153.1.1.
» https://doi.org/10.11646/zootaxa.5153.1.1 - ALBUQUERQUE, N.R., GALATTI, U. & DI-BERNARDO, M. 2007. Diet and feeding behaviour of the Neotropical parrotsnake (Leptophis ahaetulla) in Northern Brazil. J. Nat. Hist. 41(17–20):1237–1243. https://doi.org/10.1080/00222930701400954.
» https://doi.org/10.1080/00222930701400954 - AYRES, M., AYRES JR., M., AYRES, D.L. & SANTOS, A.S. 2007. BioEstat 5.0: Aplicações Estatísticas nas Áreas das Ciências Biológicas e Médicas. Instituto de Desenvolvimento Sustentável Mamirauá-IDSM/MCT/CNPq, Brasília.
- BERNARDE, P.S., MOURA-LEITE, J.C., MACHADO, R.A. & KOKOBUM, M.N.C. 2000. Diet of the colubrid snake, Thamnodynastes strigatus (Günther, 1858) from Paraná State, Brazil, with field notes on anuran predation. R. Bras. Biol. 60(4):695–699.
- CAMILLERI, C. & SHINE, R. 1990. Sexual dimorphism and dietary divergence: differences in trophic morphology between male and female snakes. Copeia (1990)3:649–658.
- DIXON, J.R. & SOINI, P. 1977. The Reptiles of the Upper Amazon Basin, Iquitos Region, Peru. Part 1. Lizards and Amphisbaenians. Part 2. Crocodilians, Turtles and Snakes. 1 ed. Milwaukee Public Museum, Contributions in Biology and Geology 12:1–91.
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- DUELLMAN, W.E. & WIENS, J.J. 1993. Hylid frogs of the genus Scinax Wagler, 1830, in Amazonian Ecuador and Peru. Occas. Pap. Mus. Nat. Hist. 153:1–57.
- FORSMAN, A. 1995. Heating rates and body temperature variation in melanistic and zigzag Vipera berus: does colour make a difference? Ann. Zool. Fenn. 32:365–374.
- HERO, J.-M. & MAGNUSSON, W. 1987. Leptophis ahaetulla Natural history note. Herpetol. Rev. 18(1):16.
- HOOGMOED, M.S. & AVILA-PIRES, T.C. 2011. A case of voluntary tail autotomy in the snake Dendrophidion dendrophis (Schlegel, 1837) (Reptilia: Squamata: Colubridae). Bol. Mus. Para. Emílio Goeldi, Ciências Naturais 6:113–117.
- LEITE, A.K., FAZOLATO, C.P., OLIVEIRA, M.L.T., SILVA, R.S. & AMORIM, P.P.S. 2022. Leptophis ahaetulla marginatus (Southern Green Parrot-Snake). Diet. Herpetol. Rev. 53(1):153.
- LOCKHART, J.S. & AMIEL, J. 2011. Nerodia sipedon (Northern Watersnake). Defensive behavior. Herpetol. Rev. 42(2):296–297.
- LOPEZ, M.S., GIRAUDO, A.R. & ARZAMENDIA, V. 2003. Leptophis ahaetulla marginatus (Southern Green Parrot-Snake). Diet. Herpetol. Rev. 34(1):68–69.
- MARTINS, M. & OLIVEIRA, M.E. 1999. Natural history of snakes in forests of the Manaus region, Central Amazonia, Brazil. Herpetol. Nat. Hist. 6(2):78–150.
- MOURA, M.R., COSTA, H.C., ABEGG, A.D., ALAMINOS, E., ANGARITA-SIERRA, T., AZEVEDO, W.S., CABRAL, H., CARVALHO, P., CECHIN, S., CITELI, N., DOURADO, A.C.M., DUARTE, A.F.V., FRANÇA, F.G.R., FREIRE, E.M.X., GARCIA, P.C.A., MOL, R., MONTERO, R., MORAES-DA-SILVA, A., PASSOS, D.C., … GUEDES, J.J.M. 2023. Unwrapping broken tails: Biological and environmental correlates of predation pressure in limbless reptiles. J. Anim. Ecol. 1–14. https://doi.org/10.1111/1365-2656.13793.
» https://doi.org/10.1111/1365-2656.13793 - MUNIZ, S.L.S, MOURA, C.C.M., VEJA, S.F., COUTO, A.A., SILVA, J.S., SANTOS, E.M. & MOURA, G.J.B. 2013. Leptophis ahaetulla marginatus (Swordsnake). Diet. Herpetol. Rev. 44(1):154.
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Publication Dates
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Publication in this collection
09 Aug 2024 -
Date of issue
2024
History
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Received
26 Apr 2024 -
Accepted
01 July 2024