Corresponding author: Thaís B. Guedes ( thaisbguedes@yahoo.com.br ) Academic editor: Uwe Fritz
© 2021 Omar M. Entiauspe-Neto, Claudia Koch, Michael B. Harvey, Guarino R. Colli, Thaís B. Guedes.
This is an open access article distributed under the terms of the CC0 Public Domain Dedication.
Citation:
Entiauspe-Neto OM, Koch C, Harvey MB, Colli GR, Guedes TB (2021) Redescription of Apostolepis ambiniger (Peters, 1869) (Serpentes: Dipsadidae: Elapomorphini). Vertebrate Zoology 71: 231-251. https://doi.org/10.3897/vz.71.e65097
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Apostolepis is a diverse genus of dipsadid snakes, currently comprising 34 species occurring in most cis-Andean South America. The taxonomy of the group is highly unstable. Upon discovering its type series, we redescribe the rare species A. ambiniger (Peters, 1869) and provide an account of its geographic distribution and morphological variation in pholidosis, osteology, and hemipenial characters. We also discuss some aspects of the taxonomy of Apostolepis.
Geographic distribution, micro-CT, morphology, Neotropical, osteology, Squamata, taxonomy
The tribe Elapomorphini contains approximately 50 species of fossorial and cryptozoic snakes, widely distributed across cis-Andean South America (
The genera of Elapomorphini can usually be differentiated from one another by scalation and coloration features, which show varying degrees of reduction associated with fossorial and cryptozoic habits (
Apostolepis is the most diverse of the elapomorphine genera, with 34 species occurring in open and forested biomes from southern Colombia and the Guyana Shield to southern Brazil and northern Argentina (
Like many congeners, Apostolepis ambiniger (Peters, 1869) has an interesting and complicated taxonomic history. It was described as Rhynchonyx ambiniger, a new genus and species, based on a single specimen “Bought; allegedly from Paraguay” (“Gekauft; angeblich aus Paraguay”) (
Koslowsly (1898) mentioned the presence of Apostolepis ambiniger in Brazil, based on specimens collected by C. Bach in Miranda, Mato Grosso do Sul. Unfortunately, all specimens collected by Julio G. Koswlosly, deposited at Museo La Plata, are presumably missing (Thales de Lema, personal communication).
A great deal of this twisty taxonomic history is due to the limited number of specimens at hand in previous studies. This condition limited a proper assessment of intraspecific variation and robust species delimitation through comparisons with congeneric species. Recently, we found the type specimen of Apostolepis ambiniger. Here we redescribe A. ambiniger, report morphological variation, and describe the skull and hemipenis to promote stabilization in this group’s taxonomy. We also present comparisons between congeners and comments on the taxonomy of Apostolepis.
We examined a total of 697 specimens of Apostolepis (Appendix
We describe osteology of the skull of the holotype of A. ambiniger (
We used the verified point record database of A. ambiniger (Table 1 in the Supplementary Table) to draw the map of its distribution and calculate its extension of occurrence (EOO) by using QGIS version 3.16 (QGIS Development Team 2021).
According to the International Code of Zoological Nomenclature (ICZN 1999, hereafter referred to as the Code), a species name (should it be in Latin, or a nominative singular particle or adjective after latinization) should agree in its gender with the generic name of its combination (Art. 31.2). Previously, we mentioned that
Rhynchonyx ambiniger
Peters, 1869:438. Holotype male (ZMB 6450; Figs
Apostolepis ambinigra
(Peters, 1869):
Apostolepis dimidiata
(Jan, 1862):
Apostolepis dorbignyi
(Schlegel, 1837):
Apostolepis vittata
(Cope, 1887):
Apostolepis tenuis
Ruthven, 1927:
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.
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.
Among congeners, the skull has only been described for three species: A. assimilis, A. cearensis, and A. sanctaeritae. The skull of Apostolepis ambiniger differs from congeners by having edentulous pterygoids (pterygoid teeth 2–4) and only 5–6 dentary teeth (dentary teeth 7–8 in A. assimilis [n = 12] and in A. cearensis and 8–10 in A. sanctaeritae [n = 5]) and by exhibiting a rectangular-shaped fronto-parietal suture in dorsal view (U-shaped in A. assimilis and A. sanctaeritae and W-shaped with an antero-median parietal indentation in A. cearensis).
It can be further distinguished from A. assimilis and A. sanctaeritae by having 4 palatine teeth (5); a strongly enlarged premaxilla with a very broad ascending process, which covers the lateral processes in dorsal view (premaxilla and ascending process less strongly developed and lateral processes visible in dorsal view in A. assimilis and premaxilla and ascending process less strongly developed in A. sanctaeritae); nasal and prefrontal dorsally in contact (separated from each other); maxillae with an edentulous and pointed anterior process (anterior part of maxillae blunt and toothed); ectopterygoid partly covered by parietal (not covered by any other bone and therefore completely visible in dorsal view); pterygoids almost entirely covered by roofing skull bones in dorsal view, except for posterior tips (anterolateral region of pterygoids visible and not covered by parietal in dorsal view in A. assimilis and lateral edges of pterygoid visible and not covered by parietal in dorsal view in A. sanctaeritae); parietal ridges about parallel to each other, almost reaching to a point where parietal forms a right-angled suture with prootic vs parietal ridges approach each other and almost reach to suture with supraoccipital in A. assimilis and ridges merge in the posterior fifth of parietal and reach to suture with supraoccipital in A. sanctaeritae; rectangular frontals in dorsal view (slightly circular [both together] in A. assimilis and irregular shaped and not rectangular in A. sanctaeritae); slender palatines with a straight medial edge (less slender and with medial edge slightly curved in A. assimilis and robust in A. sanctaeritae); parasphenoid rostrum anteriorly surpassing anterior border of frontals and in contact with ventral surface of posteromedial region of nasals (parasphenoid rostrum not surpassing frontals and not contacting nasals); absence of symmetrical bulges medially and backwards pointing projection on each side of the outer lateral edges on the basioccipital (presence of short, blunt, and backwards pointing projection in A. assimilis and presence of symmetrical bulges and backwards pointing projection in A. sanctaeritae); slender dentaries and pterygoids (both distinctly broader); similarly high prearticular and surangular crests (prearticular crest is slightly higher than surangular crest).
It further differs from A. assimilis by having an unfused parabasisphenoid and basioccipital (fused); anterior end of supratemporal overlapping posterior half of prootic and posterior end not reaching posterior end of exoccipital (anterior end of supratemporal only overlaps posterior part of prootic and posterior end protrudes posteriorly beyond posterior end of exoccipital and is thus the most posteriorly protruding bone of the skull roof); a short retroarticular process of the compound bone (longer); vomerine processes of premaxilla robust and blunt (sharply pointed); exoccipital components of occipital condyle brought together on its dorsal surface (being separated by the basioccipital).
Adult male. Total length 395 mm; SVL 356 mm; tail length 39 mm (9.8% of total length, 10.9 % of SVL). Head length 10.32 mm (2.61 % of total length, 2.89 % of SVL); head width 4.71 mm (45 % of head length); interorbital distance 2.4 mm (51% of head width); rostro-orbital distance 5 mm; naso-orbital distance 1.5 mm (14% of head length). Cervical constriction slightly distinct; head slightly distinct from neck, triangular in dorsal view, narrow anteriorly, arched in lateral view. Pupil sub-elliptical. Rostral conical, 1.81 mm wide, strongly projected over lower jaw, length of portion visible in dorsal view slightly larger than distance to external anterior edge of frontal. Internasals absent. Prefrontals paired, square, 2.15 mm long, 1.81 mm wide; each prefrontal contacting rostral, nasal, preocular, supraocular, and frontal. Frontal hexagonal, 3.12 mm long, 2.28 mm wide, contacting prefrontal, supraoculars, and parietals. Supraocular trapezoidal, longer than wider, in contact with posterior edges of prefrontal, lateral edges of frontal, superior edge of preocular, superior edge of postocular, and anterior edges of parietals. Parietals paired; right parietal 3.96 mm long, 2.25 mm at its largest width, contacting frontal, supraocular, postocular, fourth through sixth supralabials, anterior temporal, occipital, and interoccipitals. Occipitals square-shaped, enlarged; each occipital contacts a single interoccipital and dorsals. Interoccipitals three, slightly smaller than vertebral and paravertebral rows of dorsals. Nasal triangular, undivided, longer than wide; contacting rostral, prefrontal, first and second supralabials, and preocular. Nostril in anterior third of nasal, slightly visible from above. Preocular pentagonal; postocular pentagonal, contacting third through fifth supralabials, supraocular and parietal. Temporals 1 + 0, contacting sixth supralabial, latero-posterior edge of parietal, dorsals, and a single interoccipital. Six supralabials, 2–3 entering orbit; first contacting nasal, second contacting nasal and preocular, third and fourth contacting postocular, fifth contacting postocular and parietal, sixth contacting parietal; sixth and fifth supralabials largest and equal in length. Mental triangular, as long as wide. Two pairs of chinshields, second pair longer. Seven infralabials, 1–4 in contact with anterior chinshields, 5–7 in contact with posterior chinshields, first pair in contact with each other behind mental; fourth and fifth infralabials largest, equal in size. Dorsals smooth, in 15/15/15 rows. Pre-cloacal scale divided. Ventrals 219; preventrals 5. Subcaudals paired, 34/34. Terminal scale acuminate, curved ventrally.
Head uniformly dark brown dorsally and laterally, dark brown pigmentation covering up to five dorsal rows. Rostral uniformly dark brown on dorsal surface, with diffuse white pigmentation on its ventral surface and anterior edges. Supralabial blotch single and large, covering inferior edges of supralabials 1–5. White and black nuchal collars absent. Diffuse dark brown pigmentation on outer edges of infralabials and chinshields. Dark brown gular collar complete, covering up to four rows of gular scales. Dorsal background coloration light tan (discolored and dehydrated across ventrals 153–164). Dorsal stripes absent. Ventral and subcaudal coloration uniformly light tan. Caudal blotch dark brown, extending proximally 15 vertebrals and seven subcaudals. Terminal scale bicolored, dark brown dorsally, white ventrally.
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
In preservative, black coloration turns dark brown, red coloration gradually becomes light tan, and yellow ventral coloration becomes uniformly tan (Figs
Hemipenis slightly bilobed, noncapitate, semicalyculate (Fig.
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.
The snout complex is composed of the premaxillae, nasals, septomaxillae, vomers, and prefrontals (Figs
The paired nasals are about rectangular in dorsal view, about twice as long as broad, convex, in medial contact along a straight suture, the front edges form a broad V-shaped anterior region, that frames the ascending process of the premaxilla; the lateral edges are curved downwards; posteriorly the nasals share a loose transverse suture with the frontals; posterolaterally is a small contact region with the prefrontals; the posteroventral process of the nasal contacts the anteroventral process of the frontal posteriorly and the anterior tip of the dorsally oriented process of the parasphenoid rostrum ventrally; the vertical lamina of the nasals laterally contacting the medial edge of the septomaxillae.
The paired septomaxillae are about 2.6 times longer than broad, separated from each other by the vertical laminae of the nasals; each with a broad ascending conchal process, freely extending laterally, marginally visible beyond the lateral edge of the nasal in dorsal view, but not reaching the height of the lateral nasal edge; the anteromedial process of the septomaxilla is about one fifth the length of the septomaxilla, curved and anterolaterally oriented, each reaching over about half the depth of the premaxilla and contacting the posterior lamina of the lateral wing of the premaxillary ascending process, the anterior tip of the process is hardly visible in dorsal view; the posteromedial process is long and thin, makes up about 40% the length of the entire septomaxilla, medially contacting the vertical lamina of the nasal, the rounded posterior tip contacting the anteroventral tip of the frontal, directly lateral to the ventral nasal‒frontal contact region; the septomaxillary body is a complex structure made of thin bone material with a rounded indentation in its posterior view; the anterior region of the septomaxillary body is medially contacting the dorsal surface of the premaxillary vomerine process; the posterior region of the septomaxillary body and the ventral surface of the anterior two thirds of the posteromedial process of the septomaxilla contact the anterior and anterodorsal region of the vomer.
The paired vomers are complex structures, about 1.3 times longer than broad, approaching, but not contacting each other medially, and with the anterior and posterior region diverging; anteriorly and anterodorsally contacting the septomaxilla, laterally approaching but not contacting the anterior region of the palatine; the body of the vomer is made of thin bone material and globular with an anteriorly oriented opening; the bifurcate vertical posteromedial laminae are diverging dorsally and ventrally, framing but not contacting the choanal process of the palatine anteriorly.
The paired prefrontals are 1.5 times longer than broad, greatly separated from another, oriented slightly oblique, forming the anterior margin of the orbits; the anterolateral margin is undulated; the posterolateral margin is slightly concave; the dorsal edge contacts the anterolateral edge of the frontal along a slightly curved suture; anterodorsally there is a small contact region with the posterolateral edge of the nasal; the ventral edge contacts the dorsal surface of the maxilla and approaches the maxillary process of the palatine, without contacting it; in rear view, a triangular medially directed process is visible at about mid-height of the prefrontal and a large slightly oval lacrimal foramen is visible in the ventromedial region.
The paired frontals are rectangular, about 1.8 times longer than broad, almost similar in size as the nasals, slightly convex, in contact medially with a straight suture, with undulated lateral margins (Fig.
Micro-CT images of the skull without mandibles of the holotype of Apostolepis ambiniger (
The single parietal is elongate, about 1.5 times longer than broad, has a slightly convex dorsal surface and is almost entirely rectangular, except for the anterolateral processes; the anterolateral processes are moderately long and robust, framing the lateral borders of the frontals and forming the posterior and posterodorsal margin of the orbit; the anterior border is almost straight, but with a medial elongated notch, so that the parietal is not in contact with the frontal bones in this area; the anterolateral processes extend straight anteriorly, run approximately parallel to each other and form a right angle to the anteromedial edge of the parietal, resulting in a rectangular-shaped fronto-parietal suture in dorsal view; in dorsal view, the parietal covers the pterygoids completely; a ridge extends dorsally on both sides from the anterolateral process in posterior direction, almost reaching to a point where the parietal forms a right-angled suture with the prootic, both ridges are about parallel to each other; lateral to the dorsal ridges, the parietal slopes downwards with a slightly convex surface to meet ventrally the posterior half of the parasphenoid rostrum and about the anterior half of the basisphenoid portion of the parabasisphenoid; posterolaterally the parietal contacts the anterior margin of each prootic and forms the anterior limit of the foramen for the maxillary branch of the trigeminal nerve at the lateral suture with the prootic; the posterodorsal region of the parietal forms an almost straight suture with the supraoccipital. Postorbitals are absent.
The single supraoccipital is ovaloid, 1.5 times broader than long, slightly concave, and the anterior region is slightly elevated; anteriorly it contacts the parietal, anterolaterally the prootics, and posteriorly the exoccipitals; it is distinctly separated from the supratemporals; laterally the supraoccipital extends downwards (internal) to contribute to the dorsomedial walls of the otic capsules.
The paired exoccipitals are irregularly shaped, and in medial contact along a straight suture; each has a little pronounced, oblique, dorsolateral ridge, which is parallel to the inner margin of the supratemporals; anterodorsally each exoccipital contacts the supraoccipital with a slightly curved suture, anterolaterally the prootic, ventrally the basioccipital and dorsolaterally the supratemporals; the fenestra ovalis is situated at the suture between the prootic and the exoccipital, and the exoccipital forms the posterior margin of the fenestra; posterior to the fenestra ovalis halfway to the foramen magnum is a recess, bearing tow (left exoccipital) or three (right exoccipital) foramina; posteroventrally, the exoccipital components of the occipital condyle are in close contact, excluding the posterior process of the basioccipital from participation in the foramen magnum; the exoccipital contributes to the formation of the posteroventral, posteromedial and posterolateral wall of the otic capsule; the exoccipital is the most posteriorly protruding bone of the skull roof.
The single basioccipital is almost hexagonal shaped, 1.2 times longer than broad, convex; it contacts the parabasisphenoid complex anteriorly, the prootics anterolaterally, and the exoccipitals posterolaterally; posterodorsally the basioccipital is overlain by the exoccipital components of the occipital condyle, preventing its participation in the formation of the foramen magnum; the posterior end of the basioccipital (part of the occipital condyle), is rectangular-shaped in ventral view and quadrant-shaped in posterior view; the widest part of the basioccipital is at the sutures with the prootics and exoccipitals; the surface of the basioccipital is relatively smooth and the lateral edges are straight, without backwards pointing projections; a small foramen is present medially in the posterior region, at about where the basioccipital part of the occipital condyle begins.
The paired prootics are ovaloid in lateral view, 1.25 times higher than long; each prootic contacts the parietal anteriorly, the supraoccipital dorsally, the exoccipital posteriorly, the parabasisphenoid complex anteroventrally, the basioccipital posteroventrally, and the anterior part of the supratemporal on its dorsal surface; at the suture with the parietal the prootic forms the posterior border of the foramen for the maxillary branch of the trigeminal nerve and at the suture with the exoccipital it forms the anterior margin of the fenestra ovalis; the foramen for the mandibular branch of the trigeminal nerve is posterolaterally oriented and situated slightly posterior and slightly below the central part of the prootic in lateral view; there are three further foramina on the lateral surface of the prootic: a medium-sized foramen is situated in between both trigeminal nerve foramina, a larger foramen in the anteroventral region of the prootic, at the suture with the parabasisphenoid, and a small foramen is placed in between the posterior trigeminal nerve foramen and the contact region of the prootic with the suture of the parabasisphenoid and the basioccipital; in dorsal view, the prootic bears a depression in the posterior region in which the anterior tip of the supratemporal rests; the prootic contributes to the formation of the anteroventral, anteromedial and anterolateral wall of the otic capsule; few small foramina pierce the medial laminae of each exoccipital.
The unpaired parasphenoid and basisphenoid are fused to form the parabasisphenoid, forming an elongate structure, about 2.2 times longer than broad, that occupies most of the skull floor, and is concave in the parasphenoid region and convex in the basisphenoid region; the basisphenoid portion is each roughly rounded, and the parasphenoid rostrum is lanceolate with a pointed anterior tip; the anterior half of the parasphenoid rostrum bears a dorsally oriented process, which protrudes into the gap between the two vertical lamina of the frontals, and contacts them with its lateral edges; the anterior tip of the parasphenoid rostrum surpasses the anterior border of the frontals in ventral view; the anteriormost tip of the parasphenoid rostrum is freely extending, but slightly posterior to the tip, the anteriormost part of the dorsal process of the parasphenoid rostrum contacts the ventral surface of the posteromedial region of the nasals; the parasphenoid rostrum is distinctly separated from the choanal process of the palatine, the posterior ending of the vomers, and the septomaxillae; dorsolaterally the parabasisphenoid contacts the parietal, and the prootics posterior to it, and reaches its greatest width at the sutures between the parietal and the prootics; a foramen is located on each side in the posterolateral part of the basisphenoid portion, at the suture with the prootics; in dorsal view the parabasisphenoid is pierced by some foramina in the lateral region of the bone.
The palatomaxillary arch is composed of the maxillae, ectopterygoids, palatines, and pterygoids (Fig.
Micro-CT images of the skull without mandibles of the holotype of Apostolepis ambiniger (
The paired ectopterygoids are divining rod-shaped, deeply bifurcated anteriorly, with a longer medial process than lateral process; in dorsal view of the skull, the lateral process and the anterior half of the medial process are visible, whereas the rest of the ectopterygoid is covered by the parietal; the two anterior processes are directed anterolaterally and frame the posterior end of the maxilla laterally and medially without touching it, forming an almost closed maxilo-ectopterygoid fenestra; the ventral surface of the flattened posterior process has a facet, where it firmly contacts the dorsal surface of the anterolateral portion of the pterygoid.
The paired pterygoids are edentulous, flattened, elongate and slender, about 9.7 times longer than broad, corresponding to approximately half the length of the skull; except for the firm contact with the ectopterygoid, the pterygoid does not contact any of the other skull bones; in dorsal view of the skull, only small portions of the posterior tips are visible and not completely covered by the supratemporals, whereas the rest of the pterygoids are completely covered by the roofing skull bones; the anteromedial tip of the pterygoid dorsally overlaps only marginally the posteromedial tip of the palatine without touching it; in ventral view, the lateral border of the pterygoid is slightly curved posterolaterally; the medial borders of both pterygoids are nearly parallel to each other in the anterior third, with the smallest distance between them at their anterior tips; the medial border of the last two-thirds of the bone gradually tapers posterolaterally, resulting in the greatest distance between both pterygoids at their posterior tips; the posterior end of the pterygoid approaches medially the ventromedial process of the quadrate without touching it.
The paired palatines are elongate and slender, about 7.3 times longer than wide, when not considering the choanal and maxillary processes, and correspond to 24% of the length of the skull; their medial edge is straight, and both palatines are about parallel to each other; the ventral surface has four tooth loci; the teeth are subequal, curved, and rear facing; of the cranial bones the palatine only contacts the medioventral region of the prefrontal on the dorsal surface of its maxillary process; the anterior portion of the palatine, anterior to the tooth line is almost quadrangular and approaches the ventrolateral part of the vomer dorsally, without contacting it; dorsomedially, a long, thin choanal process rises and curves downwards in a semicircle, approaching but remaining distinctly separated from its counterpart medially; the short, about triangular maxillary process is situated on the lateral surface of the palatine at the level of the first tooth, directed anterolaterally, approaching but not contacting the palatine process of the maxilla; the posterior part of the palatine behind the tooth line is bifurcated, with a slightly shorter ventrolateral process and a slightly longer dorsomedial process, both tapering towards the posterior end, and the anterior part of the pterygoid extends into the gap between the two processes without touching them.
The suspensorium is composed of the supratemporals and the quadrates. Each mandible is composed of the dentary, splenial, angular, and compound bone (Figs
Micro-CT images of the skull of the holotype of Apostolepis ambiniger (
Micro-CT images of the mandible of the holotype of Apostolepis ambiniger (
The quadrates are flattened and broad dorsally, tapering dorsoventrally in lateral view, but gradually increasing in width in rear view; they are oriented oblique, from anterodorsally to posteroventrally; the posterodorsal part approaches the posterolateral region of the supratemporal medially; the medial part has a short process, which corresponds to the contact region with the columella auris, but the columella auris is not visible; the ventral part is bifurcated, with the medial branch being broader than the lateral branch and both together spanning the glenoid cavity of the retroarticular process of the mandible; the quadrate does not exceed the posterior limit of the skull roof.
The dentaries are elongate and slender, making up about 40% the length of the mandible, and are slightly curved anteromedially; the dorsal surface bears five (left dentary) or six (right dentary) tooth loci; the teeth are subequal, curved and rear facing; the lateral face is slightly convex with a mental foramen located at about the level of the fifth tooth, almost in the medial region of the dentary; at about the level of the last tooth, the dentary branches into a shorter dorsal process, which overlays the anterior part of the compound bone, and a longer lanceolate ventral process; the gap between the dorsal and ventral processes is mainly filled by the splenial, whereas the posteriormost part of the ventral process contacts the anteroventral part of the angular; the ventral process runs with its dorsal surface parallel and close along the anterior part of the medioventral region of the compound bone without touching it; in medial view, the dorsal process is bifurcated in its posterior region, distinctly behind the last tooth, with a longer dorsolateral branch and a very short ventromedial branch.
The splenials are elongate, triangular, tapered anteriorly, about 4.2 times longer than high, and represent the smallest of the mandibular bones, making up about one-fifth of the mandibular length; the anterior mylohyoid foramen is centrally positioned in the posterior part of the bone at the level of its greatest height; the posterior edge of the splenial firmly contacts the anterior region of the angular; the dorsal edge is notched in the posterior region, just before the summit of the triangle.
The angulars are elongate, triangular, tapered posteriorly, almost five times longer than high, and represent the second smallest of the mandibular bones, making up about one-fourth of the mandibular length; each angular contacts the splenial anteriorly, the compound bone laterally and dorsally, the posteriormost tip of the medial process of the dentary, and the posteromedial surface of the ventral process of the dentary; the posterior mylohyoid foramen is on the lateral surface in the anterior third of the bone.
The compound bones are elongate and slender, about 8.6 times longer than high, and represent the largest of the mandibular bones, making up about two-thirds of the length of the mandible; the prearticular and surangular crests are about similar in height, and therefore neither of them is visible in lateral or medial view; in lateral view, the compound bone tapers anteriorly, loosely fitting between the dorsal and ventral processes of the dentary; an anterodorsally oriented foramen is present in the anterior region on the lateral surface of the compound bone; the retroarticular process is moderately long, slightly medially directed, and does not surpass beyond the posterior end of the exoccipital.
This species has been historically recorded in Argentina, Brazil, Bolivia, and Paraguay (
Geographic distribution of A. ambiniger. Green circles = verified point records of A. ambiniger; red circles = wrong (or from missing specimens) point records previously attributed to A. ambiniger (see Table 1 in the Supplementary Table); Polygon = extension of occurrence (7,626.941 km²).
The identity of Apostolepis ambiniger has been subject to historical controversy.
The documented geographic distribution showed that A. ambiniger seems to be restricted to open areas ecoregions of Humid Chaco and and a small portion in the east of the Alto Paraná Atlantic Forest (Dinerstein et al. 2017) mostly in Paraguay; previous extralimital records are shown to be cases of misidentifications or mistaken localities. In its geographic distribution, A. ambiniger is sympatric with A. dimidiata, A. intermedia, and possibly with A. assimilis, A. breviceps, A. christineae, A. lineata, A. phillipsi, A. kikoi, and A. vittata. Both A. ambiniger and A. dimidiata are morphologically similar, with substantial overlap in diagnostic characters (ventrals, subcaudals, supralabials, infralabials, temporals, nasal-preocular contact, number of infralabials contacting chinshields) which has contributed to misidentifications and taxonomic instability of both species; Apostolepis dimidiata presents a remarkably wide morphological variation, that still needs to be formally assessed (see
The skull of A. ambiniger conspicuously differs from congeners, yet skulls have been previously described only for A. assimilis, A. cearensis, A. sanctaeritae. The absence of pterygoid teeth may be unique among its congeners, and possibly constitutes an autapomorphy for A. ambiniger, although this requires further comparisons among its congeners. Its distinctive morphology was highlighted by
We are deeply indebted to Uwe Fritz (Editor), Gunther Köhler, Uri García-Vázquez, and an anonymous reviewer, who kindly reviewed and provided constructive criticism to our work. OME-N thanks CNPQ for a PIBIC grant (136628/2016-8). We thank Arthur Tiutenko (Arthur Tiutenko Nature Images) for illustrating a specimen of Apostolepis ambiniger in life and hemipenis. We also thank Frank Tillack (
Material examined
Coordinates are given in SIRGAS2000.
Apostolepis aff. niceforoi (n = 1). BRAZIL: Roraima: Caracaraí, Vila de Caicubi, Rio Jufari, 1.797409°S, 61.143789°W (MZUSP 19625).
Apostolepis albicollaris (n = 2). BRAZIL: Distrito Federal: Brasília, 15.775247°S, 47.922950°W (USNM 148789, 148790).
Apostolepis arenaria (n = 5). BRAZIL: Bahia: Originally given as “Alagoado”, herein corrected to Casa Nova municipality, 9.161121°S, 40.983073°W (MZUSP 10027, 10028, 10029, 10030, 10289).
Apostolepis assimilis (n = 144). BRAZIL: Bahia: Barreiras, 12.143650°S, 45.003059°W (UMMZ 20411); Distrito Federal: Brasília, 15.775247°S, 47.922950°W (CHUNB 24456, 24474, IBSP 20566, 28734, USNM 148790); Goiás: border with Tocantins, Ilha do Bananal, Santa Isabel, 11.080152°S, 50.636316°W (IBSP 12324); Jataí, 17.899968°S, 51.730803°W (MZUSP 3783); Mineiros, 17.558692°S, 52.552554°W (IBSP 55495); Rio Verde, 17.787982°S, 50.937775°W (IBSP 10326, 12945, MZUSP 3194), Uruaçu, Cana Brava, 14.519870°S, 49.150580°W (IBSP 9154); Minas Gerais: Cabo Verde, 21.475110°S, 46.397057°W (IBSP 29448); Cambuí, 22.614916°S, 46.056915°W (IBSP 44222), Capão dos Porcos, Mariana, 20.365709°S, 43.407326°W (ZMUC 63806, holotype of Apostolepis assimilis), Caxambu, 21.98007°S, 44.932633°W (IBSP 816), Conceição dos Ouros, 22.412537°S, 45.800055°W (IBSP 33206), Entre Rios de Minas, 20.680355°S, 44.089915°W (FUNED 691), Gonçalves, 22.658901°S, 45.856063°W (IBSP 49666), Ibirité, 20.012663°S, 44.081203°W (FUNED 603), Itajubá, 22.434017°S, 45.467651°W (IBSP 9115, 9407, 9592), Itamonte, 22.284305°S, 44.874096°W (IBSP 22405), Itatiaiuçu, 20.204767°S, 44.462560°W (FUNED 510), Jaíba, 15.345628°S, 43.686232°W (FUNED 1465), Maria da Fé, 22.307992°S, 45.378278°W (IBSP 5597), Moeda, 20.330538°S, 44.055868°W (FUNED 02), Nova Lima, 20.004714°S, 43.871471°W (FUNED 550), Ouro Fino, 22.278276°S, 46.374551°W (IBSP 34306), Munhoz, 22.278276°S, 46.374551°W (IBSP 66376), Passa Quatro, 22.389193°S, 44.972256°W (IBSP 3264, 3274, 34306), Poços de Caldas, 21.812489°S, 46.588499°W (IBSP 45737, 23985, 14256), Pouso Alegre, 22.246732°S, 45.927618°W (IBSP 42162, 44597, 49942), Santa Rosa da Serra, 19.530423°S, 45.967855°W (IBSP 46088), Serra do Cipó, 19.370279°S, 43.585560°W (MZUSP 7595), Uberabinha, border with São Paulo, 20.016022°S, 47.793120°W (IBSP 888), Uberlândia, 19.133733°S, 48.333231°W (IBSP 3841, 3845, 6388, 3841), Vespasiano, 19.734960°S, 43.939672°W (FUNED 04); Mato Grosso do Sul: Amambaí, 23.111955°S, 55.230187°W (IBSP 41163), Campo Grande, 20.494551°S, 54.610826°W (IBSP 41163, 42978, 57222, MHNCI 6719, MZUSP 10155), Nova Andradina, 22.002500°S, 53.491960°W (IBSP 27489, 27489), Paranaíba, 19.671986°S, 51.191663°W (IBSP 45615), Ponta Porã, 22.512971°S, 55.713606°W (IBSP 44065); Mato Grosso: Buriti, 17.973681°S, 53.554398°W (IBSP 5346), Cuiabá, 15.426759°S, 55.943251°W (MNRJ 2031); Paraná: Londrina, 23.322525°S, 51.176065°W (IBSP 37462, 40008); Santa Catarina: Florianópolis (BGSS5344, Locality possibly in error, see Entiauspe-Neto et al. 2020d); São Paulo: Araçariguama, 23.441586°S, 47.069288°W (IBSP 83132), Barueri, 23.515768°S, 46.882181°W (IBSP 23206), Bauru, 22.327818°S, 49.107726°W (MHNCI 4790), Cabreúva, 23.316678°S, 47.082624°W (IBSP 26565), Caieiras, 23.374820°S, 46.735134°W (IBSP 40320), Caixa d’Água, coordinates unknown (IBSP 6659), Carapicuíba, 23.540539°S, 46.846607°W (IBSP 87769, 82260), Cajamar, 23.348228°S, 46.877875°W (IBSP 30408, 87083, 3186), Campo Largo, 25.463235°S, 49.537514°W (IBSP 4498), Campo Limpo, 23.635070°S, 46.754990°W (IBSP 6532), Campos do Jordão, 22.735380°S, 45.583899°W (IBSP 26796, UMMZ 204112), Carapicuíba, 23.54919°S, 46.84454795918558°W (IBSP 72970), Cotia, 23.636020°S, 46.956576°W (IBSP 24588), Ibiúna, 23.662839°S, 47.213951°W (IBSP 32672, 79312, 78900), Itapevi, 23.554159°S, 46.978074°W (IBSP 30436, 86908, 79489), Itatiba, 22.993786°S, 46.824113°W (IBSP 5703), Itu, 23.297557°S, 47.301031°W (IBSP 4180, 6606, 82230, , MHNCI 6969, MZUSP 4180, 6606), Jaguara, 23.512151°S, 46.742029°W (IBSP 70356), Jandira, 23.543093°S, 46.900625°W (IBSP 31694, 40493), Jarinu, 23.116928°S, 46.716730°W (IBSP 30019), Jundiaí, 23.203436°S, 46.939945°W (IBSP 16688), Mairinque, 23.539455°S, 47.185860°W (IBSP 41065, 89049), Osasco, 23.540895°S, 46.795779°W (IBSP 23889, 40480, 6141, 62362, 78442, MCP 64), Pirituba, 23.474910°S, 46.743881°W (IBSP 70351, 78948), Rio Grande, coordinates unknown (IBSP 40008), Santana de Parnaíba, 23.457582°S, 46.918621°W (IBSP 61761, 81066,), São Caetano do Sul, 23.626138°S, 46.565080°W (IBSP 81238), São Paulo, 23.534933°S, 46.609078°W (IBSP 318, 348, 6401, 6558, 8040, 8945, 21993, 22221, 24180, 24548, 24873, 27598, 30153, 30586, 31716, 32441, 33316, 84949, 78948), São Roque, 23.528545°S, 47.141170°W (IBSP 23548, 78641, , 79658, MHNCI 4495, 6970), Sorocaba, 23.481770°S, 47.455341°W (IBSP 15760, 40008); PARAGUAY: Trinidad: Unknown locality (MZUM 108810).
Apostolepis ambiniger (n = 20). PARAGUAY: Unknown locality (
Apostolepis cearensis (n = 140). BRAZIL: Alagoas: Piranhas, 9.607967°S, 37.768092°W (CHUFS 3217, 3365, MUFAL 1315); Bahia: Brumado, 14.207125°S, 41.675605°W (IBSP 33651, 33685), Camaçari, 12.706750°S, 38.331172°W (MZUEFS 371), Capim Grosso, 11.381030°S, 40.010385°W (MZUEFS 294), Feira de Santana, 12.223937°S, 38.990827°W (MZUEFS 12, 19, 70, 71, 74, 86, 130, 162, 166, 203, 277, 310, 315, 429, 434, 463, 464, 505, 515, 615, 624, 637, 669, 672, 689, 771, 804, 836, 841, 895, 927, 1007, 1040, 1053, 1067, 1069, 1070, 1071, 1077, 1080, 1110, 1146, 1157, 1158, 1195, 1196, 1208, 1209, 1210, 1236, 1240, 1241, 1244, 1260, 1302, 1310, 1313, 1369, 1377, 1405, 1445, 1446, 1477, 1478, 1479, 1499, 1539, 1559, 1570, 1587, 1604, 1611, 1622, 1629, 1645, 1673, 1674), Poções, 14.526737°S, 40.366499°W (MZUFBA 1595, 1796, 1805, 1813, 1826, 1827), São Gonçalo dos Campos, 12.435848°S, 38.952627°W (MZUEFS 73, 825), Jaguarari, 10.258500°S, 40.194672°W (IBSP 26203); Ceará: Aquiraz, 3.910332°S, 38.383874°W (CHUFC 1185), Beberibe, 4.183106°S, 38.131271°W (CHUFC 1628), Crateús, 5.174963°S, 40.675624°W (CHUFC 2238), Crato, 7.244635°S, 39.448914°W (IBSP 20385), Fortaleza, 3.76337°S, 38.527876°W (CHUFC 208, 826, 1240, 1242, 1243, 1524, 1525, 1526, 1527, 1528, 1529, 1531, 1539, 1620, 1621, 1622, 1623, 1623, 1624, 1625, 1626, 1627, 1629, 2001, 2236, 2287, 2243, 2633, IBSP 20020, 40262, 55318, 18219, 18220), Icó, 6.400202°S, 38.860449°W (IBSP 12106), Juazeiro do Norte, 7.228342°S, 39.314310°W (IBSP 20164), Limoeiro do Norte, 5.140528°S, 38.071232°W (IBSP 12775), Maranguape, 3.889587°S, 38.678660°W (CHUFC 2235), Quixadá, 4.969875°S, 39.017092°W (CHUFC 1221), São Benedito, 4.045633°S, 40.865595°W (CHUFC 2114, 2147), Tianguá, 3.727411°S, 40.997394°W (IBSP 77109), Ubajara, 3.865421°S, 40.980188°W (IBSP 75855, 77101), Viçosa do Ceará, 3.566272°S, 41.110413°W (IBSP 77509); Paraíba: Cabaceiras, 7.490668°S, 36.288036°W (MZUSP 9013), Campina Grande, 7.230571°S, 35.892641°W (IBSP 9050), Lagoa de Dentro, 6.673537°S, 35.377482°W (MNRJ 17055); Piauí: Teresina, 5.052687°S, 42.764437°W (IBSP 49743), Redenção do Gurguéia, 9.488261°S, 44.583311°W (IBSP 80942).
Apostolepis cerradoensis (n = 1). BRAZIL: Goiás: Minaçu, Cana Brava UHE, 13.510069°S, 48.209950°W (MCP 15219, holotype of Apostolepis cerradoensis).
Apostolepis christineae (n = 2). BOLIVIA: Santa Cruz: Puerto Suarez, German Busch, 18.970739°S, 57.816239°W (BMNH 1907.10.31.62). BRAZIL: Mato Grosso: Cáceres, 16.076427°S, 57.675286°W (MCP 12515, holotype of Apostolepis christineae).
Apostolepis dimidiata (n = 12). BRAZIL: Mato Grosso: Unknown locality (AMNH 62192); São Paulo: Unknown locality (AMNH 7245, 102252, FMNH 69934, MCZ 27661, USNM 76369, 76370, ZUEC 936, 2277, 0884, 947).
Apostolepis dorbignyi (n = 2). “AMÉRIQUE MÉRIDIONALE”: Unknown locality (MNHN 3664, holotype of Apostolepis dorbignyi), BOLIVIA: Tarija (MZUT 963).
Apostolepis flavotorquata (n = 5). BRAZIL: Bahia:Unknown locality (UMMZ 108808); Mato Grosso: Unknown locality (AMNH 93559, 935690, 93561, SMS voucher unavailable).
Apostolepis gaboi (n = 34). BRAZIL: Bahia: Ibiraba, Barra, 10.787131°S, 42.823709°W (MZUFBA 1673, 1674, 1675, 1676, 1677, 1678, 1679, 1680, 1681, 1682, 1683, 1684, 1685, 1686, 1687, 1688, 1689, 1690, 1691, 1692, 1693, 1694, 1695, 1696, 1697, 1698, 1699, 1700, 1701, 1702, 1702, 1704), Icatú-Barra, 10.787140°S, 42.823790°W (MZUEFS 981), Queimadas, 11.042418°S, 39.699621°W (MZUSP 10290).
Apostolepis goiasensis (n = 1). BRAZIL: Minas Gerais: Três Lagoas, 20.788371°S, 51.706916°W (CHFURG 1344).
Apostolepis intermedia (n = 2). PARAGUAY: San Pedro: Laguna Blanca, 23.808604°S, 56.283731°W (MHNP 11533, 11636).
Apostolepis kikoi (n = 5). BRAZIL: Mato Grosso: APM Manso, Chapada dos Guimarães, 15.090012°S, 55.712671°W (MCP 12096, holotype of Apostolepis kikoi, 14524, 14525, 11372, UFMTR 1933, paratypes of Apostolepis kikoi).
Apostolepis longicaudata (n = 1). BRAZIL: Tocantins: Estação Ecológica Serra Geral, 10.856603°S, 46.696670°W (MZUSP 14122).
Apostolepis lineata (n = 1). BRAZIL: Mato Grosso: Chapada dos Guimarães, 15.090012°S, 55.712671°W (ANSP 11211, syntype of Apostolepis lineata).
Apostolepis multicincta (n = 5). BOLIVIA: Santa Cruz: San Juan, 20.900597°S, 67.765903°W (ZFMK 66375, paratype of Apostolepis multicincta), Florida, 19.450080°S, 65.450257°W (ZFMK 75025, 75026), Santa Cruz, 17.792752°S, 63.162745°W (MNKR 729, holotype of Apostolepis multicincta, 878, paratype of Apostolepis multicincta).
Apostolepis nelsonjorgei (n = 2). BRAZIL: Goiás: Campinaçu, 13.787851°S, 48.571293°W (MZUSP 20636); Tocantins: Estação Ecológica Serra Geral, 10.856603°S, 46.696670°W (MZUSP 17615).
Apostolepis nigrolineata (n = 217). SOUTH AMERICA (
Apostolepis quinquelineata (n = 4). GUYANA: Georgetown, 6.797002°S, 58.156512°W (BMNH 89.9.30.12, holotype of Apostolepis quinquelineata). BRAZIL: Amazonas: Presidente Figueiredo, 2.031079°S, 60.025275°W (INPAH 31440); Rondônia: Porto Velho, 8.758575°S, 63.883062°W (UFROH 228, 229).
Apostolepis niceforoi (n = 1). COLOMBIA: Caquetá: Florencia, 1.614836°S, 75.608047°W (ICN 10422).
Apostolepis nigroterminata (n = 13). BOLIVIA: Unknown locality (USNM 280371); Santa Cruz: Unknown locality (BMNH 1927.8.1.180, 1927.8.1.181, 1927.8.1.182, CM 2909, MNKR 472, 942, UMMZ 60773, 67962, 67963, UTA 44687); BRAZIL: Mato Grosso: Comodoro, 13.657177°S, 59.795470°W (MPEG 26500), Vila Bela da Santíssima Trindade, 15.005081°S, 59.948396°W (MZUSP 6408); PERU: Cayaria (herein considered as Pucallpa), 8.360497°S, 74.585841°W (BMNH 1946.1.9.77, holotype).
Apostolepis quirogai (n = 1). BRAZIL: Rio Grande do Sul: Santo Ângelo, Campus URI, 28.277008°S, 54.270656°W (MCP 12185).
Apostolepis sanctaeritae (n = 31). BRAZIL: Bahia: Unknown locality (MCP 8442), Cocos, 14.181859°S, 44.537064°W (IBSP 61525, CHUNB 51360, CHUNB 23715), Correntina, 13.571514°S, 45.303100°W (CHUNB 39079), Santa Rita de Cássia, Ibipetuba, 10.866069°S, 44.610886°W (NMW 23452, holotype of Apostolepis sanctaeritae, MZUFBA 728, topotype); Distrito Federal: Brasília, 15.775247°S, 47.922950°W (IBSP 49363); Goiás: Unknown locality (IBSP 15723); Minaçu, 13.510069°S, 48.209950°W (IBSP 40478), São Domingos, 13.401192°S, 46.322214°W (IBSP 62593, IBSP 67392); Mato Grosso: Nova Xavantina, 14.662802°S, 52.362981°W (MCP 8002), Ribeirão Cascalheira, 12.939072°S, 51.828372°W (MCP 19481), São Félix do Araguaia, 11.617427°S, 50.666608°W (IBSP 15723); Minas Gerais: Betim, 19.925915°S, 44.222527°W (FUNED 03), Curvelo, 18.762770°S, 44.440999°W (IBSP 22410), Pirapora, 17.353695°S, 44.889140°W (MPEG 18347), Serra do Cipó, 19.370279°S, 43.585560°W (MZUSP 7595); Três Marias, 18.218011°S, 45.239204°W (FUNED, Without voucher); Vazante, 17.989087°S, 46.899214°W (IBSP 48041); Tocantins: Gurupi, 11.729458°S, 49.074318°W (MZUSP 8007); Lajeado, 9.839862°S, 48.321855°W (IBSP 64533, IBSP 64534, IBSP 65571, IBSP 65680, IBSP 65681), Palmas, 10.283001°S, 48.342440°W (IBSP 65267, holotype of Apostolepis ammodites), Porto Nacional, 10.683005°S, 48.377269°W (IBSP 65682, IBSP 65683, IBSP 66166), Santa Isabel, 11.384446°S, 48.073970°W (IBSP 12324).
Apostolepis tenuis (n = 2). BOLIVIA: Beni: Guayaramirim, 10.827529°S, 65.363502°W (USNM 123973); Santa Cruz: Buena Vista, Ichilo, 16.888852°S, 64.189732°W (UMMZ 64436, holotype of Apostolepis tenuis).
Apostolepis thalesdelemai (n = 39). BRAZIL: Ceará: Guaramiranga, 4.261648°S, 38.932367°W (CHUFC 1950, 2067, 2353, 2371); Ibiapina, 3.925965°S, 40.888903°W (CHUFC 2337, 2340, 2342, 2343, 2351, 2437); Maranguape, 4.008095°S, 38.818735°W (CHUFC 2102, 2208, 2212, 2213, 2218, 2339, 2347, IBSP 80734); Pacoti, 4.226994°S, 38.921911°W (CHUFC 2344, 2346, 2463, 2731, 2841); São Benedito, 4.047003°S, 40.866281°W (CHUFC 2338); Ubajara, 3.859255°S, 40.926629°W (CHUFC 1349, 2085, 2110, 2137, 2154, 2341, 2350, 2769, 2954, IBSP 80735, ZUEC 3384).
Apostolepis vittata (n = 4). BRAZIL: Mato Grosso: Chapada dos Guimarães, 15.090012°S, 55.712671°W (ANSP 11293, holotype of Rhynchonyx ambiniger vittatus; CHUNB 30656), Parque Nacional Chapada dos Guimarães, 15.090012°S, 55.712671°W (UFMTR 12259), Rio da Casca, 15.359774°S, 55.458283°W (MCP 13283).