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Phrynonax sexcarinatus (WAGLER, 1824)

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Higher TaxaColubridae, Colubrinae, Colubroidea, Caenophidia, Alethinophidia, Serpentes, Squamata (snakes)
Subspecies 
Common NamesE: Northeastern Puffing Snake
Portuguese: Caninana, Cobra-Cipó, Papa-Ovo, Papa-Pinto, Papa-Pinto-de-Papo-Vermelho, Papa-Rato, Rateira, Surucucu-Facão 
SynonymNatrix sexcarinata WAGLER 1824: 35
Ahaetulla polylepis PETERS 1867: 709
Spilotes fasciatus PETERS 1869: 443
Phrynonax fasciatus — BOULENGER 1894: 21
Phrynonax eutropis BOULENGER 1894: 22
Phrynonax lyoni STEJNEGER 1902: 185
Phrynonax atriceps WERNER 1913: 22
Phrynonax poecilonotus polylepis — AMARAL 1929: 313
Pseustes poecilonotus polylepis — BRONGERSMA 1937: 6
Phrynonax lyoni — RENDAHL 1937
Pseustes poecilonotus polylepis — BEEBE 1946: 39
Pseustes sexcarinatus — HOGE 1964: 28
Pseustes poecilonotus polylepis — PETERS & OREJAS-MIRANDA 1970: 258
Pseustes poecilonotus polylepis — GASC & RODRIGUES 1980
Pseutes poecilonotus polylepis — MEHRTENS 1987 (in error)
Pseustes poecilonotus polylepis — GORZULA & SEÑARIS 1999
Pseustes poecilonotus polylepis — WHITHWORTH & BEIRNE 2011
Phrynonax polylepis — JADIN et al. 2013
Pseustes polylepis — FELDMAN et al. 2015
Phrynonax lyoni — NATERA-MUMAW et al. 2015: 82
Phrynonax polylepis — NOGUEIRA et al. 2019
Phrynonax polylepis — LOPES & PASSOS 2023
Phrynonax polylepis — CRNOBRNA et al. 2023
Phrynonax polylepis — MURPHY et al. 2023 
DistributionBrazil (Pará), Venezuela (Trinidad), Ecuador, Peru

Type locality (polylepis): “Suriname”

Neotype locality (sexcarinatus): Porto Trombetas (01°28’01.3”S, 56°22’46.0”W; ca. 40 m above sea level; asl hereafter), municipality of Oriximiná, state of Pará, Brazil

Type locality (Spilotes fasciatus: “angeblich vom Maroni’ [Suriname]  
Reproduction 
TypesNeotype: MNRJ 20302, adult female, collected by Emiliane Pereira Gonçalves and team between November 13 and December 08, 2006. Holotype lost; was ZSM 1744/0, collected by Spix and Martius expedition to Brazil, 1817-1820 [sexcarinatus]
Holotype: ZMB 5899
Holotype: ZMB 6455 [Spilotes fasciatus PETERS 1869]
Holotype: USNM 27826 [Phrynonax lyoni] 
DiagnosisDiagnosis and Definition. Phrynonax sexcarinatus can be distinguished from all congeners by a unique combination of the following characters: dorsal scale rows usually keeled in adult specimens, except for the first series, usually smooth; dorsal scale rows usually 21/23/15; postoculars usually two; temporal formula usually 2+2; supralabials usually eight, fourth to sixth contacting the orbit; infralabials usually 13, the first eight contacting the first pair of chinshields; maxillary teeth 11–19; ventral scales 196–211 in females, 186–215 in males; subcaudal scales 95–142 in females, 110–139 in males; color in preservative (70% ethanol), adults presenting the dorsum of the head brown, supralabials and gular region cream, ventral surface of the body cream anteriorly, darkening to brown posteriorly, ventral surface of the tail brown, dorsum of the body and tail uniform brown; color in preservative (70% ethanol), juveniles with the dorsum of the head beige with scattered brown spots or blotches, supralabials and infralabials cream with brown vertical bars on the posterior edges, a cream gular region, dorsum of the body and tail beige with transverse dark blotches, ventral surface of the body creamish-white anteriorly, darkening posteriorly to beige, belly with scattered dark brown spots or blotches concentrated on the posterior of the body, ventral surface of the tail beige speckled with dark brown spots; maximum SVL 1128 mm in females, 1200 mm in males; relative tail length 32.7–46.7% SVL in females, 34.2–45.7% SVL in males. (LOPES & PASSOS 2023) 
CommentSynonymy partly after VILLA et al. 1988, PETERS & OREJAS-MIRANDA 1970, and LOPES & PASSOS 2023. Listed as synonym of Pseustes poecilonotus by WALLACH et al. 2014: 612. The record of P. poecilonotus polilepis by Lemoine et al (1998) is based on mistake of Pseustes shropshirei (Marco Natera, pers. comm.). LOPES & PASSOS 2023 revalidated sexcarinata so that polylepis becomes a junior synonym.

Habitat: partly arboreal (Harrington et al. 2018).

Distribution: see map for P. polylepis in Nogueira et al. 2019: plate 143. Distribution: Phrynonax sexcarinatus is not in Venezuela fide NATERA-MUMAW et al. 2015. COSTA & BERNILS 2015 excluded this species from their checklist of Brazilian reptiles because 1) the validity of this name is questionable; 2) recent records from Brazil lack vouchers or may be misidentified. 
References
  • Amaral, A. D. 1929. Estudo sobre ophidios neotropicos XX. Revisao do genero Phrynonax Cope, 1862. Mem. Inst. Butantan 4: 301-320 - get paper here
  • Beebe, William 1946. Field notes on the snakes of Kartabo, British Guiana, and Caripito, Venezuela. Zoologica 31: 11-52 - get paper here
  • Boulenger, George A. 1894. Catalogue of the snakes in the British Museum (Natural History). Volume II., Containing the Conclusion of the Colubridæ Aglyphæ. British Mus. (Nat. Hist.), London, xi, 382 pp. - get paper here
  • Brongersma, L. D. 1937. Herpetological Notes XIV - XVI. XV. The type of Dipsas dieperinkii Schlegel, 1837, and the genus Pseustes Fitzinger, 1843. Zoologische Mededelingen 20: 1-10 - get paper here
  • Crnobrna B, Santa-Cruz Farfan R, Gallegos C, López-Rojas JJ, Llanqui IB, Pisco GP, Arbaiza AK 2023. Herpetological records from the Abujao basin, central Peruvian Amazon. Check List 19(4): 433-465 - get paper here
  • Gasc & Rodrigues 1980. Liste preliminaire des Serpents de la Guyane francaise. Bull. Mus. Nat. Hist. Nat. Paris 2 (4): 559-598
  • Gonzalez R. C. et al. 2020. Lista dos Nomes Populares dos Répteis no Brasil – Primeira Versão. Herpetologia Brasileira 9 (2): 121 – 214 - get paper here
  • Gorzula, Stefan & Senaris, J. Celsa 1999. In: Contribution to the herpetofauna of the Venezuelan Guayana. I: a data base. Scientia Guaianae, Caracas, No. 8 [1998], 269+ pp.; ISBN 980-6020-48-0
  • Harrington, Sean M; Jordyn M de Haan, Lindsey Shapiro, Sara Ruane 2018. Habits and characteristics of arboreal snakes worldwide: arboreality constrains body size but does not affect lineage diversification. Biological Journal of the Linnean Society 125 (1): 61–71 - get paper here
  • Hedges SB, Powell R, Henderson RW, Hanson S, and Murphy JC 2019. Definition of the Caribbean Islands biogeographic region, with checklist and recommendations for standardized common names of amphibians and reptiles. Caribbean Herpetology 67: 1–53
  • Hoge, A.R. 1964. Posição sistemática de Natrix sexcarinatus Wagler 1824. Mem. Inst. Butantan 30 (1960-1962): 27-30 - get paper here
  • Jadin, R. C., Burbrink, F. T., Rivas, G. A., Vitt, L. J., Barrio-Amorós, C. L. and Guralnick, R. P. 2013. Finding arboreal snakes in an evolutionary tree: phylogenetic placement and systematic revision of the Neotropical birdsnakes. Journal of Zoological Systematics and Evolutionary Research. 52 (3): 257–264; doi: 10.1111/jzs.12055 - get paper here
  • LOPES, L. V., & PASSOS, P. 2023. Taxonomic status of the enigmatic Natrix sexcarinata Wagler, 1824 (Serpentes: Colubridae: Colubrinae). Zootaxa 5244 (2): 123-144 - get paper here
  • Mehrtens, J.M. 1987. Living snakes of the world in color. Sterling Publ. Co., hic., New York, NY: 480 pp.
  • Natera-Mumaw, Marco; Luis Felipe Esqueda-González & Manuel Castelaín-Fernández 2015. Atlas Serpientes de Venezuela. Santiago de Chile, Dimacofi Negocios Avanzados S.A., 456 pp. - get paper here
  • Nogueira, Cristiano C.; Antonio J.S. Argôlo, Vanesa Arzamendia, Josué A. Azevedo, Fausto E. Barbo, Renato S. Bérnils, Bruna E. Bolochio, Marcio Borges-Martins, Marcela Brasil-Godinho, Henrique Braz, Marcus A. Buononato, Diego F. Cisneros-Heredia, 2019. Atlas of Brazilian snakes: verified point-locality maps to mitigate the Wallacean shortfall in a megadiverse snake fauna. South American J. Herp. 14 (Special Issue 1):1-274 - get paper here
  • Peters, James A.; Donoso-Barros, Roberto & Orejas-Miranda, Braulio 1970. Catalogue of the Neotropical Squamata: Part I Snakes. Bull. US Natl. Mus. 297: 347 pp. - get paper here
  • Peters, Wilhem Carl Hartwig 1867. Über Flederthiere (Pteropus gouldii, Rhinolophus deckenii, Vespertilio lobipes, Vesperugo temminckii) und Amphibien (Hypsilurus godeffroyi, Lygosoma scutatum, Stenostoma narirostre, Onychocephalus unguirostris, Ahaetulla poylepis, Pseudechis scutella Monatsber. königl. Akad. Wiss. Berlin. 1867 (November): 703-712 - get paper here
  • Peters,W.C.H. 1869. Über neue Gattungen und neue oder weniger bekannte Arten von Amphibien (Eremias, Dicrodon, Euprepes, Lygosoma, Typhlops, Eryx, Rhynchonyx, Elapomorphus, Achalinus, Coronella, Dromicus, Xenopholis, Anoplodipsas, Spilotes, Tropidonotus). Monatsber. k. preuss. Akad. Wiss. Berlin, 1869: 432-447 - get paper here
  • Rendahl, H. 1937. Einige Reptilien aus Ecuador und Bolivia. Ark. Zool. 29 (13): 1-19.
  • Stejneger, L. 1902. An annotated list of batrachians and reptiles collected in the vicinity of La Guaira, Venezuela, with descriptions of two new species of snakes. Proc. US Natl. Mus. 24 [1901]: 179-192. - get paper here
  • Torres-Carvajal O, Pazmiño-Otamendi G, Salazar-Valenzuela D. 2019. Reptiles of Ecuador: a resource-rich portal, with a dynamic checklist and photographic guides. Amphibian & Reptile Conservation 13 (1): [General Section]: 209–229 (e178) - get paper here
  • Werner, F. 1913. Neue oder seltene Reptilien und Frösche des Naturhistorischen Museums in Hamburg. Reptilien der Ostafrika-Expedition der Hamburger Geographischen Gesellschaft 1911/12. Leiter: Dr. E. Obst. Reptilien und Amphibien von Formosa. Jb. Hamb. wiss. Anst., 30 [1912], 2. Beiheft: 1-39, 40-45, 45-51 - get paper here
  • Whithworth, A. & Beirne, C. 2011. Reptiles of the Yachana Reserve. Global Vision International, 130 pp. - get paper here
 
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