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Erythrolamprus aenigma ENTIAUSPE-NETO, ABEGG, KOCH, NUÑEZ, AZEVEDO, MORAES, TIUTENKO, BIALVES & LOEBMANN, 2021

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Higher TaxaColubridae (Dipsadinae), Colubroidea, Caenophidia, Alethinophidia, Serpentes, Squamata (snakes)
Subspecies 
Common NamesE: Savannah Racer Snake
P: Corredeira de Savana
S: Guarda-camino de Sabana 
SynonymErythrolamprus aenigma ENTIAUSPE-NETO, ABEGG, KOCH, NUÑEZ, AZEVEDO, MORAES, TIUTENKO, BIALVES & LOEBMANN 2021
Erythrolamprus poecilogyrus schotti – ENTIAUSPE-NETO et al. 2016: 61 (part.)
Erythrolamprus poecilogyrus – AZARAK & FARIAS 2017: 45 (non WIED-NEUWIED) 
DistributionBrazil (Roraima), Guyana (Kuruwini), Venezuela (Bolívar)

Type locality: Boa Vista (2°49’12” N, 60°40’19” W, 90 m a.s.l.), Roraima state, Brazil.  
Reproduction 
TypesHolotype: IBSP 90722, Adult female (Figs 3–4), col. M. Duarte, F. L. Franco, R. Feltran and W. Azevedo on 03 March 2019.
Paratypes: INPA-H 34377, Adult female, col. P. Azarak in 2012, from Rorainópolis (0°50’30” N, 60°40’48” W, 200 m a.s.l.), Roraima, Brazil; adult female (CHFURG 5068), col. D. Loebmann in 2016, from Boa Vista (2°49’12” N, 60°40’19” W, 90 m a.s.l.), Roraima, Brazil; adult female (IBSP 90743), col. M. Duarte, F. L. Franco and W. Azevedo in 2019, dissected head, from Boa Vista (2°49’12” N, 60°40’19” W, 90 m a.s.l.), Roraima, Brazil; adult female (IBSP 90727, Fig. 5E), col. M. Duarte, F. L. Franco and W. Azevedo in 2019, dissected head, from Boa Vista (2°49’12” N, 60°40’19” W, 90 m a.s.l.), Roraima, Brazil; juvenile female (IBSP 90660), col. M. Duarte, F. L. Franco and W. Azevedo in 2019, from Boa Vista (2°49’12” N, 60°40’19” W, 90 m a.s.l.), Roraima, Brazil; juvenile female (AMNH-R 60803), from Kuyuwini (2o12’30” N, 58o28’30” W), Guyana.
Referred specimens: Unsexed adult (MTR 20581), from Fazenda Salvamento, Boa Vista (2°49’12” N, 60°40’19” W, 90 m a.s.l.), Roraima, Brazil; unsexed adult (MTR 20551) from Estação Ecológica Maracá, Amajari (3°39’07” N, 61°22’15” W, 100 m a.s.l.), Roraima, Brazil; unsexed adult (MTR 20546) from Estação Ecológica Maracá, Ama- jari (3°39’07” N, 61°22’15” W, 100 m a.s.l.), Roraima, Bra- zil; unsexed adult (MZUSP 10389, field tag LJV 2570), col. L.J. Vitt, from Novo Brasil, Boa Vista (2°49’12” N, 60°40’19” W, 90 m a.s.l.), Roraima, Brazil; unsexed adult (CVULA, voucher unknown, photographic voucher), col. C.B. Amorós, reportedly lost, from “Barinitas” (un- known locality), Bolívar, Venezuela; unsexed adult (CVU- LA, voucher unknown, photographic voucher), col. C.B. Amorós, reportedly lost, from Santa Elena de Uairen (4o36’8” N, 61o06’36” W, 900 m a.s.l.), Bolívar, Venezuela. 
DiagnosisDiagnosis: The new species is assigned to Erythrolamprus based on molecular evidence, considering that until now, no unambiguous morphological synapomorphy is known for this genus [see Zaher et al. (2009) and Grazziotin et al. (2012)]. The new species can be differentiated from all other Erythrolamprus species by the following combination of characters: (1) 19/19/15 dorsal scales; (2) preocular present, contacting loreal, supraocular, 3rd and 4th supralabials; (3) loreal present; (4) temporals 1+2 (1+1+2 in a single specimen); (5) supralabials eight, 4th–5th in contact with orbit; (6) infralabials 10/10 (11/10 in a single specimen), 1st–5th in contact with anterior chinshields; (7) ventrals 149–154 in females; (8) subcaudals 45–50 in females; (9) dorsal background coloration light grey, scales reticulated, with black margins and diffuse light blue lateral edges; (10) ventral and subcaudal coloration white with conspicuous diffuse and irregular small black blotches or dots; (11) black and white nuchal collars present, arrow-shaped, black nuchal collar over parietals, temporals and occipitals, up to one and a half scales wide; white nuchal collar over interoccipitals and dorsals, up to two scales wide, with outer black margins; (12) SVL 167–476 mm, TL 40–114 mm (Entiauspe-Neto et al. 2021).

Variation: Only female specimens are available. SVL 167– 476 mm (365 ± 105.1, n = 5), TL 40–114 mm (85.6 ± 28.5, n=5),headlength10–18mm(14±3,n=5),headwidth 6–9 mm (8.25 ± 1.5, n = 5). Ventrals 149–154 (152 ± 9, n = 5), subcaudals 45–50 (47 ± 2.5, n = 5). One specimen has 1+1+2 temporals on left side of head (IBSP 90722, holotype); another specimen possesses 10/11 infralabials (IBSP 90727). Largest specimen SVL 478 mm, and TL 105 mm (Entiauspe-Neto et al. 2021).

Coloration in life: Head uniformly light grey dorsally and laterally, scales with black margins; head uniformly white ventrally; supralabials light grey or light brown, with black margins; black and white nuchal collars, arrowshaped; black nuchal collar over parietals, temporals and occipitals, up to one and a half scales wide (inconspicuous in IBSP 90743); white nuchal collar over interoccipitals and dorsals, up to two scales wide, with black margins; dorsal background coloration light grey or light brown (Fig. 5), scales reticulated, with black margins (vestigial markings in IBSP 90743) and diffuse light blue lateral edges (white in IBSP 90743); ventral and subcaudal coloration white with conspicuous diffuse and irregular small black blotches (Entiauspe-Neto et al. 2021).

Comparisons (data for other species in parenthesis): Erythrolamprus aenigma is most likely to be confused with taxa from the E. poecilogyrus species complex, since these species share 19/19/17 dorsals.
In order to assure that E. aenigma is not allocable to any of the available names (including ‘forgotten’ synonyms of other taxa), the taxonomic history of E. poecilogyrus needs to be briefly summarized here. An overview of the geographic distribution of available valid names associated to E. poecilogyrus is also given (Fig. 2), As is an overview of osteological comparisons (Table 1).
Dixon & Markezich (1992) recognized four valid subspecies for E. poecilogyrus, and 28 available names associated with this species. The recognized subspecies were: (i) E. p. poecilogyrus, described based on specimens from “Rio Jucú”, in what is currently the Brazilian state of Espirito Santo, likely in the vicinity of Vitória municipality, in the Atlantic Rainforest of southeastern Brazil; (ii) E. p. schotti (Schlegel, 1837), described based on a specimen from São Paulo state, in southeastern Brazil; (iii) E. p. sub­ lineatus (Cope, 1860), described based on a specimen from Buenos Aires, Argentina; (iv) E. p. caesius (Cope, 1862), described based on a specimen from Santa Fé, Argentina.
Of these names, none were described based on specimens from ‘open areas’ of northern Brazil, which could possibly represent E. aenigma. Three names have unknown or imprecise localities, i.e. Coluber alternans Lichstenstein, 1823, described from “Brazil”, Liophis verecundus Jan, 1863, described from an unknown locality, and Liophis cobella flaviventris Jan & Sordelli, 1866, described from “Brazil”. Liophis verecundus is currently considered a junior synonym of Erythrolamprus almadensis (Dixon, 1989). Liophis cobella flaviventris is currently considered a junior synonym of Erythrolamprus p. poecilogyrus (Wied-Neuwied, 1825) (cf. Dixon & Markezich 1992). The names Coluber m­nigrum Raddi, 1820, Coluber alternans Lichtenstein, 1823 and Natrix forsteri Wagler, 1824 were considered conspecific with, i.e., junior synonyms of, E. p. poe­ cilogyrus by Dixon & Markezich (1992) [...] (Entiauspe-Neto et al. 2021).

Erythrolamprus aenig­ma can be distinguished from E. reginae based on the basis of its dorsal scale counts (17/17/15), ventral pattern (black square-shaped marks), and dorsal pattern in life (uniformly olive green) (Fig. 8E).
It is possible to distinguish it from E. aenigma based on its dorsal scale counts (17/17/15), ventral pattern (black square-shaped marks), and dorsal pattern in life (uniformly olive green or greyish brown) (Fig. 8C).
Both E. zweifeli and E. reginae can be differentiated from E. aenigma based on the dorsal scale counts (17/17/15), dorsal coloration in life (green, yellow or orange), and ventral pattern (large, black square-shaped marks).
It is also possible to distinguish E. dorso­ corallinus from E. aenigma based on its dorsal scale counts (17/17/15), dorsal coloration in life (blue, yellow, or orange), ventral pattern (black square-shaped markings), and higher subcaudal counts (62–80).
Erythrolamprus oligolepis [...] can be readily distinguished from E. aenigma based on its lower dorsal scale counts (15/15/15), dorsal coloration (uniformly brown or light green), and ventral pattern in life (uniformly cream or white) (Fig. 8D).
Erythrolamprus rochai can be distinguished from E. aenigma based on its dorsal pattern (body stripes and variegated) and ventral pattern (uniformly white).
From Erythrolamprus epinephelus (Cope, 1862), including its subspecies (E. e. bimaculatus (Cope, 1899); E. e. epinephelus (Cope, 1862), E. e. juvenalis (Dunn, 1937), E. e. kogiorum (Bernal-Carlo, 1994), E. e. opistothaeni­ us (Boulenger, 1908), and E. e. pseudocobella (Peracca, 1914), E. aenigma can be distinguished by its dorsal scale counts (17/17/15), dorsal pattern (black crossbands, dark brown stripes or uniformly grey, light brown, red, or green) and ventral pattern (uniformly yellow, cream, or alternating black squares) (Dixon 1983a). The same diagnosis applies to taxa formerly assigned to E. epinephelus, these being E. albiventris (Jan, 1863), E. fraseri (Boulenger, 1894), and E. lamonae (Dunn, 1944) (Entiauspe-Neto et al. 2021). 
Comment 
EtymologyThe specific epithet, aenigma, is a Latin singular noun in nominative case, meaning a “mystery” or “riddle”, and refers to the noteworthy absence of males in our examined sample of specimens. 
References
  • Entiauspe-Neto, Omar M.; Arthur D. Abegg, Claudia Koch, Leroy P. Nuñez, Weverton S. Azevedo, Leandro J. C. L. Moraes, Arthur Tiutenko, Tatiane S. Bialves & Daniel Loebmann 2021. A new species of Erythrolamprus (Serpentes: Dipsadidae: Xenodontini) from the savannas of northern South America. Salamandra 57 (2): 196–218
 
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