Research Article |
Corresponding author: Germán Chávez ( vampflack@yahoo.com ) Academic editor: Oliver Hawlitschek
© 2023 Germán Chávez, Luis A. García-Ayachi, Alessandro Catenazzi.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Chávez G, García-Ayachi LA, Catenazzi A (2023) A new species of Microteiid Lizard (Gymnophtalmidae, Cercosaurini, Selvasaura) from a remote area in the Peruvian Andes. Evolutionary Systematics 7(1): 123-132. https://doi.org/10.3897/evolsyst.7.99118
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We describe a new species of Selvasaura lizard from the western slopes of the eastern Andes of central Peru. Among other characters, the new species differs from congeners in having keeled dorsal scales and more transverse rows of scales on dorsum. We present a phylogeny as additional evidence supporting delimitation of the new species.
New species, eastern Andes, central Peru, phylogeny
Since the beginning of this century, molecular phylogenetic studies have brought some light into the taxonomy of the neotropical gymnophthalmid lizards of the subfamily Cercosaurinae (
However, more unnamed species of Selvasaura may occur than those previously reported. The western side of the eastern Andes in northern Peru have been poorly explored. Thus, explorers of these mountains often discover new species and report new distribution records (
We collected specimens in the morning while conducting opportunistic visual surveys in the upper Chontayacu River, central Peru, in October 2018. We euthanized specimens with T61 (Embrutamide), fixed them in 10% formalin, and stored them in 70% ethanol. Before fixation, we obtained tissue samples (liver) from all individuals and stored them in absolute ethanol. We fixed everted hemipenes of the holotype and paratype CORBIDI 21866 using hot water, following
The format of the description and terminology of the morphological characters follow
We analysed DNA sequences of the new species to confirm generic placement and to investigate its phylogenetic affinities. We obtained fragments of the nuclear oocyte maturation factor gene (c-mos), and the three mitochondrial genes: small subunit rRNA (12S), large subunit rRNA (16S), and NADH dehydrogenase subunit 4 (ND4) from the holotype (CORBIDI 21865) and adult paratype (CORBIDI 21866). We extracted DNA from samples of muscle tissue preserved in 90% ethanol. We used the primers and protocols for the polymerase chain reactions described in
The electronic version of this article in Portable Document Format (PDF) will represent a published work according to the International Commission on Zoological Nomenclature (ICZ), and hence the new name contained in the electronic version is effectively published under that Code from the electronic edition alone. This published work and the nomenclatural acts it contains have been registered in ZooBank, the online registration system for the ICZN. The ZooBank LSIDs (Life Science Identifiers) can be resolved and the associated information viewed through any standard web browser by appending the LSID to the prefix http://zoobank.org/. The LSID for this publication is: urn:lsid:zoobank.org:pub: F76580A7-0F79-4109-8F0F-4047EDC2A223.
The phylogeny using three mitochondrial and one nuclear gene fragments confirms the generic placement of the new species within Selvasaura (Fig.
Phylogenetic tree of Selvasaura. Maximum Likelihood analysis based on a concatenated dataset of three mitochondrial (12S, 16S, ND4) and one (c-mos) nuclear gene fragments, showing the relationships among Selvasaura candesi sp. nov. (in bold), its congeners, and species of Proctoporus, Cercosaura, and Petracola (Gymnophthalmidae). ML bootstrap values are indicated at each node.
Morphologically, the new species can be differentiated from species of the genus Andinosaura, Euspondylus, Gelanesaurus, Oreosaurus, Petracola, Riama, most Anadia and Placosoma by having an undivided lower palpebral disc (vs. divided), and from Pholidobolus because the lower palpebral disc is transparent (vs. opaque). Furthermore, the new species is distinguished from species of Centrosaura, Echinosaura, Gelanesaurus, Kataphractosaurus, Neusticurus, and Potamites by having homogeneous rectangular dorsal scales (vs. heterogeneous, granular and tuberculate dorsal scales); from Cercosaura by the absence of enlarged gular scales (vs. medial gulars distinctly enlarged forming longitudinal rows); from Dendrosauridion and Wilsonosaura by having keeled dorsal scales (vs. smooth or slightly keeled respectively); from Macropholidus by bearing a great reduction in the size of the lateral scales (vs. no reduction); and from Proctoporus by bearing lateral scales adjacents to ventrals non granular (vs. granular).
Holotype.
Peru • Adult male; Huánuco Department, Marañón Province, 11 km SW San Pedro de Chonta, on the road to Antaquero Community; 8°42'59.6"S, 76°57'22.3"W; 2,458 m; 15 Oct. 2018; G. Chávez leg.; CORBIDI 21865 (Figs
Dorsal (left) and ventral (right) views of (A, B) the male holotype (CORBIDI 21865, SVL = 49.5 mm), (C, D) male paratype (CORBIDI 21866, SVL = 49.1 mm), and (E, F) juvenile paratype (CORBIDI 21867, SVL = 28.3 mm) of Selvasaura candesi sp. nov.
Heads in lateral (left) and dorsal (right) view of adult males of Selvasaura candesi sp nov. A, B. Holotype, CORBIDI 21865, SVL=49.5 mm; C, D. Paratype, CORBIDI 21866, SVL=49.1 mm. Red arrows show variation in the condition of anteriormost supraocular: A, B. Not in contact with superciliaries; C, D. Fused to anteriormost superciliary.
Left hemipenis of the holotype (CORBIDI 21865) showing: A. Asulcate; B. Left; C. Sulcate, and D. Right side of the organ. Scale bar: 2.5 mm.
Paratypes. Peru • 1 ♂ adult, 1 juvenile, collected with the holotype; G. Chávez leg.; CORBIDI 21866 (Figs
A medium sized lizard (adult males SVL 28.3–49.5 mm, n=2) characterized by the following combination of morphological features: 1) body slender, dorsoventrally depressed in males, females unknown; 2) head slightly short, pointed, about 1.6 times longer than wide; 3) ear opening distinct, moderately recessed; 4) nasals separated by an undivided frontonasal; 5) prefrontals, frontal, frontoparietals, parietals, interparietal, and postparietals present; 6) parietals polygonal, slightly longer than wide; 7) supraoculars four, anteriormost fused (2 individuals) or not (1 individual) with anteriormost superciliary; 8) superciliary series complete, consisting in four scales; 9) nasal plate divided posterior to nostril; 10) loreal present, in contact with second supralabial; 11) supralabials seven; 12) genials in four pairs, first and second pairs in contact; 13) collar present, containing 9–10 enlarged scales; 14) dorsals in 40–41 transverse rows, rectangular, nearly twice as long as wide, subimbricate, keeled; 15) ventrals in 24–25 transverse rows; square to rectangular, juxtaposed, smooth; 17) scales around midbody 38–43; 18) lateral scales at midbody reduced in size, in 8–9 rows; 19) limbs pentadactyl, all digits clawed;20) forelimb reaching anteriorly the third supralabial; 21) subdigital lamellae under Finger IV 14–16; 22) subdigital lamellae under Toe IV 19–21; 23) femoral pores per thigh 9–10 in males; 24) rectangular preanal scales large, four in number; 25) tail about 0.4 times longer than body; 26) caudals larger than wide, subimbricate, rugose dorsally, smooth ventrally; 27) lower palpebral disc transparent, undivided; 28) dorsal surface of head, body and limbs brown with black speckling, dorsal surface of tail pale brown to reddish brown; a thick yellowish brown vertebral stripe on dorsum, a row of 1–2 faded black rings on each flank; throat creamy white with minute black spots or blotches within each scale; belly yellow (creamy white in juveniles) with minute black spots within each scale; ventral surfaces of limbs yellow or yellowish orange; anal area saffron yellow or reddish yellow; tail red or reddish orange in males (pale red in juveniles) with fine black speckling; iris pale orange in males.
Selvasaura candesi sp. nov. differs from S. brava by having keeled dorsal scales in adults (vs rugose), a larger number of dorsal rows 40–41 (vs 33–36), a higher number of lateral scales 8–9 (vs 6–7), a larger number of scales around midbody 38–43 (vs 32–34), a yellow belly in adult males (vs creamy white), and black minute spots or small blotches within every ventral scale (vs grey speckles). The new species differs from S. almendarizae (
Body slender; legs moderately long, tail complete; head length 24.0% of SVL, head width 15.7% of SVL; snout pointed, moderately long, eye-nose distance 29.6% of HL; neck distinct, collar present; head scales smooth; rostral scale wider than long, slightly higher than adjacent supralabials, in contact with frontonasal, nasals, and first supralabials; frontonasal pentagonal, slightly wider than long; prefrontals present, in wide contact medially; frontal longer than wide, in contact with second and third supraoculars; frontoparietals pentagonal, longer than wide, in contact with third and fourth supraoculars, parietals and interparietal; supraoculars four, not in contact with superciliars; superciliary series complete, consisting of four shields; anteriormost superciliar not fused with anteriormost supraocular, in contact with prefrontal and loreal anteriorly; parietals in contact with frontoparietals, fourth supraocular, dorsalmost postocular, one temporal and two postparietals; interparietal longer than wide, in contact with three postparietals posteriorly; postparietals five; nasal shield divided posterior to nostril, in contact with first and second supralabial; frenocular triangular, in contact with loreal anteriorly, anteriormost superciliar dorsally, second and third (at one point) supralabial ventrally, suboculars posteriorly, on both sides; palpebral disc oval, translucent, undivided; postoculars three; temporals polygonal, supratympanic temporal one; supralabials six, fourth below the centre of eye; infralabials six; mental wider than long, in contact with first infralabials; postmental single, pentagonal, in contact with first and second infralabials; genials in four pairs, first and second pair in contact medially, first pair in contact with second and third infralabials, second pair in contact with third and fourth infralabials, third pair in contact with fourth and fifth infralabials, fourth pair in contact with fifth and sixth infralabials; gulars 14; plates along collar 10; dorsal scales homogenous, rectangular, longer than wide, keeled, in 40 transverse rows; dorsals (enlarged scales) around body at fifth transverse ventral scale row 10, at 10th transverse ventral scale row 13, at 15th transverse ventral scale row 15; laterals (smaller than dorsals) at fifth transverse ventral scale row 9–10, at 10th transverse ventral scale row 9–8, at 15th transverse ventral scale row 10–10; ventrals squared to rectangular, juxtaposed, in 24 transverse rows; longitudinal rows of ventrals at midbody 10; scales around midbody 43; anterior preanal plate scales two; posterior preanal plate scales four; scales on tail rectangular, subimbricate, slightly keeled dorsally at tail base, smooth and juxtaposed ventrally; subdigital lamellae under Finger IV 15/15 (4/5 distal lamellae single and smooth, remaining lamellae divided in two subconical segments); subdigital lamellae under Toe IV 22/21 (4/4 distal lamellae single and smooth, remaining lamellae divided in two subconical segments); femoral pores 9/10.
Measurements of the holotype (in mm): SVL 49.5; TL 38.5; HL 12.3; HW 7.7; HD 5.4; EN 3.6; FLL 12.0; HLL 19.7; AGD 24.8.
(Fig.
General colouration pattern is as described for the holotype in life. The dorsal colouration on head, body and tail is dark brown with black markings. The throat is creamy yellow, belly greyish white, tail pinkish yellow. Ventral surfaces of limbs are yellow.
The completely everted, left hemipenis is a bicapitate organ measuring about 5.5 mm (Fig.
General scutellation data of the type series is given in Table
Characters | Selvasaura candesi sp. nov. (n = 3) | Selvasaura brava (n = 10) | Selvasaura evasa (n = 7) | Selvasaura almendarizae (n = 3) |
---|---|---|---|---|
Maximum SVL in Adult males | 49.5 | 45.9 | 46.1 | 39.7 |
Supralabials | 7 | 7 | 7–8 | – |
Scales in collar | 9–10 | 9–11 | 9–10 | 7–19 |
Transverse rows of dorsals | 40–41 | 33–36 | 33–38 | 25–32 |
Laterals at midbody | 8–9 | 6–7 | 0–3 | 5 |
Scales around midbody | 38–43 | 32–34 | 32–34 | – |
Transverse rows of ventrals | 24–25 | 22–25 | 20–25 | 22–23 |
Ventrals across belly | 10–10 | 10–10 | 10–12 | 8–10 |
Preanal plate scales | 4 | 4 | 4–5 | 4 |
Lamellae under Finger IV | 15–16 | 14–16 | 14–19 | 12–16 |
Lamellae under Toe IV | 21–19 | 19–22 | 17–24 | 18–20 |
Femoral Pores in Adult males (per leg) | 9–10 | 7–9 | 9–12 | 9–12 |
The specific epithet “candesi” refers to the acronym CANDES (Consultores Asociados en Naturaleza y Desarrollo) in recognition of their efforts supporting the herpetological research in Peruvian territory.
This species is known only from the type locality, a montane forest in the upper basin of Chontayacu River at 2,400 m a.s.l. in the western slopes of the eastern Andes of central Peru (Fig.
Map showing the distribution of the Selvasaura candesi sp. nov. (red star), Selvasaura almendarizae (fucsia circles), Selvasaura brava (yellow circle), Selvasaura evasa (blue circles) and an unnamed Ecuadorian population of Selvasaura (green circle) mentioned by
Photographs of the area where Selvasaura candesi sp. nov. was found. A. Panoramic view of the mountains and the cloud forests at the type locality in the western Andes of central Peru; B. Habitat and microhabitat (shown by red arrow) of S. candesi alongside the Chontayacu River, Huánuco Department, Peru.
We present morphological and genetic evidence that supports our recognition of a new species of Selvasaura from Peru. Regarding scutellation, we noted that condition of supraoculars is variable in the new species (anteriormost supraocular fused or not to the anteriormost superciliar) as is noted in S. evasa, whereas it appears stable in S. brava (anteriormost supraocular fused to anteriormost superciliary). Despite the condition of supraoculars as a diagnostic character in S. brava (
Regarding hemipenial morphology, S. candesi sp. nov., has unilobed hemipenis like S. evasa (
Our phylogenetic analysis of four concatenated gene fragments (three mitochondrial and one nuclear) agrees with previous findings suggesting that specimen QCAZ 12891 from El Pangui, Zamora-Chinchipe Province, Ecuador may represent a new taxon (
Selvasaura lizards are known from a handful of places from the Andes of central Peru to northern Ecuador (Fig.
Arboreal habits previously reported for S. brava (
We thank Omar Torres-Carvajal and Pablo J. Venegas for their insightful feedback that helped us to improve considerably this manuscript. Our genetic analyses were performed thanks to the support of the Global Genome Initiative Awards Program of the Global Genome Biodiversity Network (GGBN). This research would not have been possible without the exceptional support of Carlos Garnica and Hatzel Ortiz, Consultores Asociados en Naturaleza y Desarrollo´s staff. GC thanks Paola Martinez, Lourdes Durand, and Clever Llagas for their enthusiastic help in the field. We also thank Pablo J. Venegas for kindly providing photographs in life of Selvasaura evasa for comparison.
Material examined.
Selvasaura evasa. Peru: SAN MARTIN: Huicungo CORBIDI 15115–20.
Selvasaura almendarizae. Ecuador: PASTAZA: Centro Ecológico Zanjarajuno QCAZR9140.
Selvasaura almendarizae. Ecuador: NAPO: Wildsumaco Wildlife Sanctuary QCAZR5073, QCAZR12798.
GenBank accession numbers for the taxa and genes sampled in this study.
Species | Voucher | 12S | 16S | ND4 | c-mos |
---|---|---|---|---|---|
S. almendarizae | QCAZ 5073 | MW204462 | MW204460 | MW655619 | MW655617 |
S. almendarizae | QCAZ 9140 | MW204463 | MW204461 | NA | MW655618 |
S. almendarizae | QCAZ 12798 | KU902206 | KU902281 | KU902362 | KU902125 |
S. brava | MUSM 32718 | MH579609 | MH579645 | NA | MH579700 |
S. brava | MUSM 32738 | MH579612 | MH579648 | NA | MH579703 |
S. brava | NMP6V 75653 | MH579611 | MH579647 | NA | MH579702 |
S. brava | NMP6V 75654 | MH579613 | MH579649 | NA | MH579704 |
S. brava | NMP6V 75655 | MH579610 | MH579646 | NA | MH579701 |
S. candesi | CORBIDI 21865 | OQ848614 | OQ848616 | OQ851467 | OQ851469 |
S. candesi | CORBIDI 21866 | OQ848615 | OQ848617 | OQ851468 | OQ851470 |
S. evasa | CORBIDI 15117 | KU902203 | KU902278 | KU902359 | KU902122 |
S. evasa | CORBIDI 15118 | KU902204 | KU902279 | KU902360 | KU902123 |
S. evasa | CORBIDI 15119 | KU902205 | KU902280 | KU902361 | KU902124 |
Selvasaura sp. | QCAZ 12891 | KU902207 | NA | KU902363 | KU902126 |
Cercosaura argulus | QCAZ 4888 | KP874738 | KP874790 | KP874900 | KP874842 |
Cercosaura pacha | MUBI 14515 | MT531388 | MT524456 | MT522847 | MT512510 |
Proctoporus carabaya | CORBIDI 14710 | KU902164 | KU902245 | KU902321 | KU902083 |
Proctoporus pachyurus | CORBIDI_11811 | KU902178 | KU902255 | KU902335 | KU902097 |
Petracola ventrimaculata | CORBIDI 10482 | KP874775 | KP874827 | KP874937 | NA |