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Apostolepis ambiniger (PETERS, 1869)

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Higher TaxaColubridae (Dipsadinae), Colubroidea, Caenophidia, Alethinophidia, Serpentes, Squamata (snakes)
Common NamesE: Paraguayan Blackhead
Portuguese: Falsa-Coral 
SynonymRhynchonyx ambiniger PETERS 1869: 438
Apostolepis ambinigra — BOULENGER 1896: 237 (unjustified emendation)
Apostolepis ambiniger — PETERS & OREJAS-MIRANDA 1970: 22
Apostolepis ambinigra — BAUER et al. 1995: 74
Apostolepis ambiniger — DE LEMA 2002
Apostolepis ambinigra — WALLACH et al. 2014: 50
Apostolepis ambiniger — NOGUEIRA et al. 2019 
DistributionW Brazil, Bolivia, Paraguay

Type locality: “angeblich aus Paraguay”  
TypesHolotype: ZMB 6450 
DiagnosisDiagnosis. A species of Apostolepis with the following characters: (1) 15/15/15 dorsal scales; (2) preocular present, contacting nasal; (3) loreal absent; (4) temporals 1+0 or absent; (5) supralabials six, 2nd–3rd entering orbit; (6) infralabials 6–7, 1st–4th in contact with first pair of chinshields; (7) ventrals 219–244 (219–244 in males, 230–242 in females); (8) subcaudals 21–33 (28–33 in males, 21–26 in females), cloacal plate divided; (9) dorsal ground coloration uniformly red or orange with black markings; (10) ventral pattern uniformly yellow or red; (11) first 3–6 vertebrals black, nuchal collars and dorsal stripes absent; (12) distal black caudal blotch overlapping last 9–15 vertebrals and 7–11 subcaudals; (13) terminal scale white or bicolored black dorsally, white ventrally; (14) supralabial blotch large or small, up to four supralabial scales wide; (15) white blotch on rostral vestigial or absent; (16) SVL 390–600 mm, tail length 33–60 mm, tail is 8–10% of total length and SVL is 90–92% of total length (Entiauspe-Neto et al. 2021).

Comparisons. Apostolepis ambiniger occurs near to A. dimidiata (characters in parentheses), another red species of Apostolepis that inhabits the Cerrado and Gran Chaco. Apostolepis ambiniger can be distinguished from A. dimidiata by its uniformly red dorsal pattern (two lateral black stripes over a red background), 3–6 black dorsal rows in the nuchal area (up to 3 rows), a white or bicolored terminal scale (bicolored or black), rostral white blotch absent or vestigial (present, small, reaching nasal), and slightly bilobed hemipenis, with expanded lips and a distinct sulcus spermaticus (unilobed, inconspicuous sulcus spermaticus).
From Apostolepis goiasensis Prado, 1942, A. ambiniger can be distinguished by its uniformly colored dorsum (three black dorsal stripes), 3–6 black dorsal rows in the nuchal area (0–3 scales), rostral blotch absent or vestigial (large, white, reaching prefrontals), fourth supralabial separated from parietal (in contact), larger size (up to 600 mm in A. ambiniger compared to a maximum known SVL = 392 mm), and a calyculate hemipenial apex (spinulate apex).
Several relatively rare congeners may occur in sympatry with A. ambiniger in some parts of its range but are unlikely to be confused with it. Unlike A. breviceps Harvey, Gonzales and Scrocchi, 2001, A. ambiniger has six supralabials with the second and third entering the orbit (five supralabials, third only entering orbit), two pairs of chinshields (one pair), and infralabials 1–4 contacting the first chinshield (1–3). Its uniformly red dorsum makes A. ambiniger readily distinguished from A. lineata (striped), as well as its 1–4 infralabials contacting the first chinshield (1–3). Additionally, unlike A. intermedia Koslowsky, 1898, female A. ambiniger have 21–26 subcaudals (28–31) and unlike both A. vittata (Cope, 1887) and A. christineae Lema, 2002, male A. ambiniger have 28–33 subcaudals (24–28).
Various extralimital or non-sympatric species, such as Apostolepis adhara França, Barbo, Silva-Jr, Silva and Zaher, 2008, A. albicollaris Lema, 2002, A. arenaria Rodrigues, 1993, A. assimilis (Reinhardt, 1861), A. borelli Peracca, 1904, A. cearensis Gomes, 1915, A. cerradoensis Lema, 2003, A. dorbignyi (Schlegel, 1837), A. flavotorquata (Duméril, Bibrón and Duméril, 1854), A. gaboi Rodrigues, 1993, A. kikoi Santos, Entiauspe-Neto, Araújo, Souza, Lema, Strüssmann and Albuquerque, 2018, A. multicincta Harvey, 1999, A. nelsonjorgei Lema and Renner, 2004, A. phillipsi Harvey, 1999, A. quirogai Giraudo and Scrocchi, 1998, A. sanctaeritae Werner, 1924, A. tenuis Ruthven, 1927, and A. thalesdelemai Borges-Nojosa, Lima, Bezerra and James, 2016 have both white or black nuchal collars present, whereas A. ambiniger lacks any trace of white nuchal pigment and lacks collars. Furthermore A. longicaudata Gomes in Amaral, 1921, A. niceforoi Amaral, 1935, A. nigrolineata (Peters, 1869), A. pymi Boulenger, 1903, A. serrana Lema and Renner, 2006, and A. striata Lema, 2004, have longitudinal dorsal stripes whereas A. ambiniger has a uniformly-colored dorsum (Entiauspe-Neto et al. 2021).

Coloration in life. Dorsal head uniformly black, without rostral blotches or nuchal collars; lateral head uniformly black, or with small to vestigial white supralabial blotch covering no more than two scales. Infralabials uniformly black, or with white blotch covering 2–3 scales. Ventral portion of head black, with diffuse white pigmentation on posterior chinshields. Rostral lacking white blotch. Dorsal background coloration red or orange (Figs. 4–5); first 3–6 dorsal scale rows black. Ventral coloration yellow or light red. Distal black caudal blotch overlapping last 9–15 vertebrals and 7–11 subcaudals. Terminal scale white or bicolored black dorsally, white ventrally (Entiauspe-Neto et al. 2021).

Sexual dimorphism. Males of Apostolepis ambiniger have relatively longer tails (F7,6 = 17.02, P = 0.002), fewer ventrals (t10,8 = 2.77, P = 0.014), and more subcaudals (t11,8 = 4.19, P = 0.001) than females (Fig. 7). In males, the tail is 8–10% (9 ± 0, n = 7) of total length and SVL is 90–92% (91 ± 0, n = 7) of total length, whereas, in females, the tail is 6–9% (7 ± 0, n = 6) of total length and SVL is 91–94% (93 ± 0, n = 6) of total length. Males have 218–244 (229 ± 8, n = 10) ventrals and 27–34 (30 ± 2, n = 11) subcaudals, whereas females have 230–251 (239 ± 6, n = 8) ventrals and 23–31 (25 ± 3, n = 8) subcaudals (Entiauspe-Neto et al. 2021). 
CommentSynonymy: Apostolepis tenuis RUTHVEN 1927 and Rhynchonyx ambiniger vittatus COPE 1887 have been considered as synonyms of ambinigra by PETERS & OREJAS-MIRANDA 1970.

Distribution: see map in Entiauspe-Neto et al. 2021: 246 (Fig. 12). 
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