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Leptodeira annulata (LINNAEUS, 1758)

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Higher TaxaColubridae (Dipsadinae), Colubroidea, Caenophidia, Alethinophidia, Serpentes, Squamata (snakes)
Subspecies 
Common NamesE: Banded Cat-eyed Snake
G: Bananennatter
Portuguese: Cacaual, Cobra-Cipó, Cobra-Olho-de-Gato, Dormideira, Jararaca, Jararaca-de-Parede, Jararaca-de-Patioba, Jararaca-de-Tabuleiro, Jararaca-do-Rabo-Fino, Jararaquinha, Jiboia-Dormideira, Olho-de-Gato, Rabo-Fino 
SynonymColuber annulatus LINNAEUS 1758: 224
Sibon annulatus — FITZINGER 1826: 60 (non Leptognathus annulata GÜNTHER 1872)
Leptodeira annulata — FITZINGER 1843: 27
Dipsas annulata — DUMÉRIL & BIBRON 1854: 1141
Dipsas septentrionalis KENNICOTT in BAIRD 1859
Eteirodipsas annulata var. rhomboidalis JAN 1863: 105 (nomen nudum)
Leptodira annulata var. COPE 1866
Dipsas approximans GÜNTHER 1872: 32
Sibon annulatum — COPE 1875: 157
Sibon annulatum — COPE 1878: 33
Sibon annulatum — GARMAN 1884: 17
Eteirodipsas Wieneri SAUVAGE 1884: 146
Leptodeira annulata — GÜNTHER 1895
Leptodeira polysticta GÜNTHER 1895
Leptodeira ocellata GÜNTHER 1895
Leptodeira personata — GÜNTHER 1895
Leptodeira rhombifera GÜNTHER 1895
Leptodira annulata – BOULENGER 1896: 95 (part)
Leptodira albofusca – BOULENGER 1896: 95 (part)
Leptodira nycthemera WERNER 1901: 598
Leptodeira annulata annulata — AMARAL 1929: 78 (part)
Leptodeira annulata polysticta — STUART 1935: 52
Leptodeira annulata septentrionalis — SMITH 1944
Leptodeira septentrionalis — DITMARS 1946
Leptodeira annulata annulata — BEEBE 1946: 31
Leptodeira annulata septentrionalis — TAYLOR 1949: 211
Leptodeira maculata — WERLER & SMITH 1952
Leptodeira annulata — TAYLOR 1954
Leptodeira rhombifera — TAYLOR 1954
Leptodeira ocellata — BRATTSTROM & HOWELL 1954
Leptodeira annulata annulata — DUELLMAN 1958: 47
Leptodeira annulata — PETERS & OREJAS-MIRANDA 1970: 153
Leptodeira annulata annulata — DUELLMAN 1978: 247
Leptodeira annulata annulata — GASC & RODRIGUES 1980
Leptodeira annulata polysticta — WEBB 1984
Leptodeira annulata — LINER 1994
Leptodeira annulata — PONTES et al. 2008
Leptodeira annulata — WALLACH et al. 2014
Leptodeira annulata — COSTA et al. 2022 
DistributionPanama, Colombia (incl. Valle del Cauca), Venezuela, French Guiana, Guyana, Brazil (Paraná [HR 27: 216], Amapá, Amazonas, Pará, Rondonia, Roraima, Espírito Santo, Bahia, S Ceará, Rio de Janeiro, Acre, Rio Grande do Norte, Pernambuco, Piauí, Sergipe), Bolivia, Paraguay, N Peru (Pasco etc.)
Argentina (Corrientes, Entre Rios, Santa Fe, Chaco, Formosa, Salta, Tycumán), Ecuador

Type locality: “America”; restricted to “lower Rio Amazon, Pará, Brazil” by DUELLMAN 1958.  
Reproductionoviparous. Females of this species may store sperm for up to 5 years and then give birth to young (PETZOLD 1982). 
TypesSyntypes: NRM 69 and NRM without voucher number; also given as UUZM (= ZMUU) (Zoologicka Institutionen Uppsala) 9 and 35. 
DiagnosisDiagnosis: Leptodeira annulata can be distinguished from their South American congeners by the following combination of characters: (1) dorsal scales 17–23/17–23/13–17; (2) presubocular absent; (3) supralabial scales 6–9/6–9; (4) infralabial scales 9–12/9–12; (5) ventral counts in males 178–202 and fe- males 180–203; (6) subcaudal counts in male 84–108 and females 78–104; (7) dorsum of the head light brown to grayish brown generally without spots; (8) occipital region light brown to grayish brown without stripes; (9) first dorsal blotch with half-moon shape generally fused with other dorsal blotches in the first third of body forming a zigzag pattern; (10) number of dark brown dorsal blotches in males 25–51 and females 23–53; (11) number of dorsal blotches in tail in male 13–38 and females 11–23; (12) SVL in males 302–655 mm and females 305–782 mm; and (13) tail long in males 26.72% of TTL and females 24.33% of TTL.
Leptodeira annulata differs from L. ashmeadii, L. bakeri, L. larcorum, L. ornata, and L. pulchriceps by the absence or only a medial thin line in the occipital region (vs. occipital region with well-defined dark stains); from L. tarairiu sp. nov. and L. pulchriceps by the absent of presubocular (vs. presubocular present); and from L. approximans by having the basal region of the hemipenis covered by small spines that extend in a line along the sulcus spermaticus (vs. basal region of the hemipenis covered by small spines that do not extend to the capitulum). (Costa et al. 2022, supplement S2). 
CommentSynonymy after PETERS & OREJAS-MIRANDA 1970, Villa et al. 1988, and Costa et al .2022. Barrio-Amorós 2019 thinks that pulchriceps is probably a synonym of L. annulata but Costa et al. revalidated pulchriceps. Daza et al. (2009) erroneously synonymized L. maculata with L. cussiliris. Because maculata has priority, cussiliris becomes a synonym of maculata. Kaiser et al. 2013 rejected the (sub-) generic names Crossmanus Hoser 2012, Ginafabaserpenae Hoser 2012, Lukefabaserpens Hoser 2012 invalid and rejected their use instead of Leptodeira.

The review by Barrio-Amorós 2019 is difficult to follow as he does not provide a list of specimens studied or a map with localities, although he does provide a lot of photos that nicely show the variation within the genus.

Subspecies: Leptodeira annulata rhombifera GÜNTHER 1872 and Leptodeira annulata pulchriceps DUELLMAN 1958 have been elevated to full species.

Distribution: Reports from Mexico, Guatemala, Belize ?, El Salvador, Nicaragua, Costa Rica represent L. rhombifera. See map in Nogueira et al. 2019 and revised map in Costa et al. 2022 (Fig. 3 in supplement S2). Not in Nayarit, where it is replaced by L. polysticta, fide LOC-BARRAGÁN et al. 2024.

Type species: Coluber annulatus LINNAEUS 1758 is the type species of the genus Leptodeira FITZINGER 1843. The name of the genus is derived from Greek (leptos = slender, deire = neck).

Tribus Imantodini. MYERS 2011 erected the new tribus Imantodini for the genera Leptodeira and Imantodes. In addition to molecular support, the loss of bifurcation of the sulcus spermaticus (Imantodes and some Leptodeira) or reduction to a small terminal fork or expanded flat area (other Leptodeira) sets off the mainly scansorial Imantodini from all other Dipsadinae except the terrestrial Mesoamerican-North American night-snakes, which appear to form an unnamed clade.

Hybridization: ROZE 1958 reports Leptodeira annulata annulata x ashmeadii hybrids.

Habitat: partly arboreal (Harrington et al. 2018). 
EtymologyThe generic name is derived from the Greek words leptos, meaning "thin" and deire, meaning "neck."

The specios has been named after the diminutive form (“annul-”) of Latin “anus” = ring.

ashmeadi has been named after the collector of the types, Samuel Ashmead. 
References
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