A New Species of Bachia Gray, 1845 (Squamata: Gymnophthalmidae) from the Western Brazilian Amazonia

Home , Bachia, Bachia bicolor, Bachia bresslaui, Bachia dorbignyi, Bachia flavescens, Bachia panoplia

Zootaxa 3636 (3): 401–420 ISSN 1175-5326 (print edition) www.mapress.com/zootaxa/ Article ZOOTAXA Copyright © 2013 Magnolia Press ISSN 1175-5334 (online edition) http://dx.doi.org/10.11646/zootaxa.3636.3.1 http://zoobank.org/urn:lsid:zoobank.org:pub:13E09793-A6C9-4CF2-9ED8-55A2A9F04541 A new species of Bachia Gray, 1845 (Squamata: Gymnophthalmidae) from the western Brazilian Amazonia

MAURO TEIXEIRA JR1,2, FRANCISCO DAL VECHIO1, PEDRO M. SALES NUNES1, ANTONIO MOLLO NETO3, LUCIANA MOREIRA LOBO4, LUIS FERNANDO STORTI5, RENATO AUGUSTO JUNQUEIRA GAIGA6, PEDRO HENRIQUE FREIRE DIAS6, MIGUEL TREFAUT RODRIGUES1 1Laboratório de Herpetologia, Departamento de Zoologia, Instituto de Biociências, Universidade de São Paulo, CEP 05508-090, São Paulo, SP, Brazil. 2E-mail: [email protected] 3Universidade Federal do ABC, Centro de Ciências Naturais e Humanas, Laboratório de Vertebrados. Av. dos Estados 5001, Bloco A, 6° andar DI624. CEP 09210-971. Santo André, SP, Brazil. 4Alimini Consultoria Cientifica LTDA, R. Manoel Vieira Sarmento 03, Chácara Santana, CEP 05831-150, São Paulo, SP, Brazil. 5Departamento de Biologia Animal e Vegetal, Museu de Zoologia, Universidade Estadual de Londrina, CEP 86051-990, Londrina, PR, Brazil. 6Biotropica Consultoria Ambiental LTDA, Av. Santo Antônio 571, Jd. Cascatinha, CEP 37701-036, Poços de Caldas, MG, Brazil.

Abstract

A new species of Bachia of the B. dorbignyi group, Bachia scaea sp. nov., is described from the left bank of the upper Madeira River, at Rondônia state, at the western Brazilian Amazonia. The new species resembles morphologically B. dorbignyi and B. peruana, and seems to be related with the former species based on molecular data (16S and c-mos sequences). Nonetheless the presence of a first temporal separating parietal and supralabial scales and the absence of clawed fingers in the new species, can promptly distinguish it from their close relatives. This description ends with several-decades of stasis in the taxonomy of the Bachia dorbignyi group from Amazonian lowlands, and also presents new evidence that supports the Madeira River as a vicariant barrier.

Key words: Bachia scaea sp. nov., fossorial habits, Amazon Forest

Introduction

Bachia species of the B. dorbignyi group are widely distributed over the western Amazon Forest and the Andean slopes (Dixon 1973). The first described species was the in this group originally placed in the genus Chalcides by Duméril and Bibron (1839) (C. dorbignyi, from Santa Cruz, Bolivia). Later, Gray (1845) recognized Duméril and Bibron’s species as belonging to a distinct genus, and described Bachia to accommodate it. By the end of the XIX century, Cope (1868; 1896) described Heteroclonium bicolor and Ophiognomon trisanale, both now in the genus Bachia (Dixon 1973). During the first decades of the following century four more species were described, by Werner (1901), in the genus Cophias (C. peruanus), and also by Noble (1920) (B. intermedia), Ruthven (1925) (B. talpa), and Burt and Burt (1931) (B. barbouri). Finally, Dixon (1973) in his revision of the genus Bachia, described the last species, B. huallagana, and the B. dorbignyi group achieved its current content. Although other Bachia groups, such as the B. bresslaui one, have experienced a high number of descriptions in recent years (Castrillon & Strussmann 1998; Kizirian & McDiarmid 1998; Rodrigues et al. 2007, 2008; Freitas et al. 2011), the B. dorbignyi group has witnessed a long taxonomic stasis. This traditional arrangement of Bachia in species groups based on morphological features, as defined by Dixon (1973), has been recently challenged, as molecular phylogenetic approaches are showing that they may not represent natural arrangements, as they are not monophyletic (Kohlsdorf & Wagner 2006; Galis et al. 2010; Kohlsdorf et al. 2010). The species from B. dorbignyi group appear in distinct clades along the topology in different molecular studies, however as for the other groups, the monophyly is never recovered (Kohlsdorf &

Accepted by S. Carranza: 4 Mar. 2013; published: 5 Apr. 2013 401 Wagner 2006; Galis et al. 2010; Kohlsdorf et al. 2010). The topologies themselves are also not in agreement, and further studies need to be done to solve this question. Herein based on specimens from the left bank of upper Madeira River we recognize and describe a new species of Bachia, which can be assigned morphologically to B. dorbignyi group, along with information on its natural history, and a hypothesis of its phylogenetic position.

Material and methods

Sampling methods The field samplings were carried out during a three years period (2010–2012), with four surveys of 15 days every year (January, April, July and October). Three main areas (general coordinates: [1] 9°26' S, 64°49' W; [2] 9°35' S, 65° 3' W, and [3] 9°36' S, 65°24' W) were inventoried, at both banks of Madeira River, along Porto Velho municipality, Rondônia state, Brazil. Active samplings were performed by 12 persons each sampling period, and were complemented with pitfall traps; each trap consisting of seven aligned 100L buckets buried at ground level, connected with a 10m long and 50 cm high plastic vertical fence. The total effort was of 9,213 hours/man in the active search and 5,380 buckets/day. Additionally, a few more specimens found at the vicinities of Porto Velho city town were also included in our type series.

Morphology All morphological analyses were made after fixation. Length measurements were taken to the nearest 0.1mm with a Mitutoyo digital caliper; scale counts and the analysis of other morphological characters were performed with a stereomicroscope Zeiss STEMI SV6. Scale nomenclature follows Dixon (1973). Although the grouping arrangement proposed by Dixon (1973) may not represent monophyletic groups (Kohlsdorf & Wagner 2006; Galis et al. 2010; Kohlsdorf et al. 2010), the new species shows morphological features that fit the characterization of what Dixon (1973) defined as the B. dorbignyi group; thus for comparative purposes we compared the new species with the other groups as a whole, and with those within the B. dorbignyi group in more detail. Comparisons were made with specimens housed in the herpetological collection of the Museu de Zoologia da Universidade de São Paulo (MZUSP), Universidade Federal do Acre (UFAC) and Museu de Zoologia da Universidade Estadual de Campinas (ZUEC) (Appendix I), and with data from the literature for both external (Vanzolini 1961a,b; Dixon 1973; Avila-Pires 1995; Castrillon & Strussmann 1998) and hemipenial morphology (Cope 1896; Presch 1978; Nunes 2011). Meristic characters were: (SC) superciliaries; (TS) temporal scales; (FP) femoral pores; (PP) preanal pores; (PS) preanal shields; (SAB) scales around midbody; (DO) dorsals scales counted from parietals to the row over cloacal region; (VE) ventrals scales counted between interbrachials and preanal shields; (GU) gular scales, between mental plates and interbrachials ; (CA) caudal scales. The snout-vent length (SVL) and tail length (TL) were also measured, the last one only on intact tails. A hemipenis of one individual of the new species, and one of B. dorbignyi were prepared following the procedures described by Manzani and Abe (1988), modified by Pesantes (1994) and Zaher (1999). The retractor muscle was manually separated and the everted organ filled with stained petroleum jelly. The organ was immersed in an alcoholic solution of Alizarin Red for 24 hours in order to stain eventual calcified structures (e.g. spines or spicules), an adaptation of the procedures described by Uzzell (1973) and Harvey and Embert (2008). The terminology of hemipenial structures follows Dowling and Savage (1960), Savage (1997) and Myers and Donnelly (2001, 2008).

Distribution Distributional records of species of the Bachia dorbignyi group were compiled from the literature (Burt & Burt 1931; Vanzolini 1961a,b; Donosos-Barros 1968; Dixon 1973; Avila-Pires 1995; Castrillon & Strussmann 1998; Dirksen & De la Riva 1999; Silva 2005; Jerez & Tarazona 2009; Bernarde et al. 2011), online databases (GBIF 2012; SpeciesLink 2012, HerpNET [www.herpnet.org]) and from examined specimens (Appendix I).

Molecular phylogeny To infer the relationship of the new species with the remaining Bachia species, we used a molecular phylogenetic

402 · Zootaxa 3636 (3) © 2013 Magnolia Press TEIXEIRA JR. ET AL. approach. The DNA was extracted from tissue samples (from liver or tail muscle) preserved in 100% ethanol (Fetzner 1999). Two mitochondrial (16S and 12S) (Palumbi 1996) and one nuclear (c-mos) (Godinho et al. 2005) fragments were amplified using standard PCR protocols. The PCR cycle protocol consisted of one initial cycle of 94°C for 5 min followed by 35 cycles of 94°C for 40 sec, 51°C for 16S and 56°C for 12S and 52°C for C-mos for 40 sec, 72°C for 40 sec and 72°C elongation for 7 min. PCR products were directly purified with Exonuclease I and Shrimp Alkaline Phosphatase (USB or Fermentas). Automated sequencing was carried out using BigDye Terminator v3.1 Cycle Sequencing kit (Applied Biosystems), followed by analysis on ABI Prism 310, 3700 or 3170 Genetic Analyzer Sequencers (Applied Biosystems) at the Instituto de Química da Universidade de São Paulo (IQUSP, São Paulo, Brazil) and Instituto de Ciências Biomédicas da Universidade de São Paulo (ICB, São Paulo, Brazil). Resulting sequences were manually edited in CodonCode Aligner v.3.7.1.1. (http://www.codoncode.com). Three individuals of the new species were included along with 15 other Bachia taxa as ingroups, and Potamites ecpleopus (Cope, 1875), Placosoma glabellum (Peters, 1870), Cercosaura eigenmanni (Griffin, 1917) and C. oshaughnessyi (Boulenger, 1885), as outgroups (Appendix II). Although we sequenced the 12S gene, we noticed that the 12S sequences available at GeneBank (DQ383203–DQ383212), are identical among several Bachia species [B. barbouri, B. heteropa alleni (Barbour, 1914), B. huallagana, B. intermedia, B. m. monodactylus (Daudin, 1802), B. m. parkeri Ruthven, 1925, B. panoplia Thomas, 1965, B. peruana, B. scolecoides Vanzolini, 196l, B. trisanale], therefore we exclude them from our analysis. Each gene was aligned separately using ClustalX v.2. (Larkin et al. 2007) with default settings. The protein coding gene, c-mos, was translated into amino acids to check the alignment. The alignments were concatenated in a molecular dataset including 801 bp: 16S = 451 bp, with 135 variable sites, and c-mos = 350 bp, with 93 variable sites. A Bayesian phylogenetic analysis (BA) was performed using MrBayes v3.2. software (Ronquist et al. 2012) in on the concatenated matrix, including two partitions: one for the mitochondrial gene (16S); and a second one for the nuclear (c-mos), using the best-fit model of evolution for the data, selected through jModelTest v0.1.1 software (Posada 2008) using Akaike Information Criterion (AIC) (GTR+G+I for 16S, and GTR+G for c-mos). Two independent Bayesian runs were performed for each partition, starting with a random tree, with four Markov chains, for 10,000,000 generations with trees being sampled each 1,000 generations. Consensus tree and statistics were calculated after discarding 25% of the initial trees as burn-in samples. Convergence for independent runs was checked through the final standard deviation of split frequencies, and a stationary distribution was assumed if its value was under 0.01. Statistical support for each node was calculated using posterior probability, considered significant for values above 0.95. A Maximum Likelihood (ML) analysis was also performed in Treefinder (Jobb 2011), with default settings, using the same models identified by jModelTest v0.1.1 software (Posada 2008) described above. The statistical confidence was calculated with 1,000 bootstrap replicates (Felsenstein 1985), and considered high for values above 70%. Two separate BA and two ML analyses were also performed for each gene alone (16S and c-mos) using the same parameters as described above. The majority 50% consensus tree saved with posterior probabilities and bootstrap values on the nodes were visualized using FigTree 1.3.1 (http://tree.bio.ed.ac.uk/). Uncorrected genetic distances (p-distances) were calculated using MEGA 5 (Tamura et al. 2011), for each gene separately.

RESULTS

Taxon description

Bachia scaea sp. nov. (Figs. 1–2)

Bachia flavescens Marçal et al. (2011: 262)

Holotype: MZUSP 101586, an adult male from the left bank of Madeira River, (9°26'55.30"S, 64°50'0.51"W, 101 m a.s.l., SAD69), Porto Velho municipality, state of Rondônia, Brazil, collected by the authors on 30th September 2010. Field number H 828.

NEW SPECIES OF BACHIA Zootaxa 3636 (3) © 2013 Magnolia Press · 403 FIGURE 1. Lateral (A), dorsal (B) and ventral (C) views of the head of the holotype of Bachia scaea sp. nov. (MZUSP 101586). Scale bar = 5mm.

Paratypes: MZUSP 103415, adult male and MZUSP 103414, juvenile; field numbers H 3133, H 3132, respectively (9°35'42.30"S, 65° 3'54.11"W, 96 m a.s.l., SAD69). MZUSP 100631, adult female; field number H 510 (9°26'16.18"S, 64°49'58.22"W, 123 m a.s.l., SAD69). MZUSP 103408, adult female; field number H 2479 (9°35'4.53"S, 65° 4'9.40"W, 129 m a.s.l., SAD69). MZUSP 101735, MZUSP 101736, MZUSP 103413, adult females; field numbers SAME 1130, SAME 1661, MTR 21266 (approximately 09°07' S, 64° 30'' W). All from Porto Velho municipality, state of Rondônia, Brazil. Etymology: The specific epithet scaea is derived from the Greek word “skaios” which means “on the left”, referring to its geographical position regarding the Madeira River, as it is found only on its left bank, while its putative sister species, B. dorbignyi, is found only at its right bank. Diagnosis: (1) A median-sized species of Bachia (maximum SVL= 69 mm); (2) prefrontals absent; (3) 5 supralabials; (3) supraoculars absent and 2-2 supraciliars; (4) femoral pores absent, 1-1 preanal pores in males; (5) 24–26 scales around midbody; (6) 51–54 dorsal rows of scales; (7) 40–43 ventral rows of scales; (8) 6–7 gulars; (9)

NEW SPECIES OF BACHIA Zootaxa 3636 (3) © 2013 Magnolia Press · 405 4 preanal shields; (10) no supralabials in contact with parietal; (11) dorsal scales hexagonal, imbricate, smooth; (12) fore and hindlimbs without clawed-digits; (13) first temporal present, separating fifth supralabial and parietal; (14) second chin shield not reaching oral border; (15) frontonasal present, reduced. Description of the holotype: Body elongate, with a slight cervical constriction on head, snout rounded, tail longer than body. Rostral small, barely visible from above, contacting first supralabial, nasal and frontonasal. Viewed from above the rostral is about twice as wide as high; on lateral view it projects not far from the anterior margin of jaw. Frontonasal trapezoidal, as wide as long, wider posteriorly, contacting rostral, nasal and frontal. Prefrontals absent. Frontal roughly pentagonal, longer than wide, with anterior margin slightly convex, broader than, and in contact with frontonasal, and nasal; lateral margins straight to slightly concave, in contact with loreal, first and second superciliaries; posteriorly angulose, broadly contacting parietals. Frontoparietals and interparietal absent. Parietals large, longer than wide, slightly longer and slightly narrower than frontal, roughly pentagonal; their anterior margin deeply indented and in broad contact with frontal, contacting the second superciliary, three upper temporals and the dorsals. Posterior borders of parietals and dorsals coincides with a very slight transverse cervical constriction in the occipital region. Supraoculars absent. Two superciliaries of about the same size. Nasal large, longer than high. Nostril in the anterior lower margin of the nasal, invading the upper border of first supralabial. Loreal roughly squared, in contact with nasal, frontal, first superciliary, a small irregular anterior subocular and second and third supralabials. Frenocular absent. Six supralabials; third, fourth and fifth under the orbital region, fifth the highest and largest, not contacting parietal; fourth the smallest. Three suboculars; second longest; third one contacting second superciliary and an elongate anterior temporal. Postocular absent. Eyelid present with an undivided semitransparent disc. A small temporal scale between fifth and sixth supralabials and first and second temporal scales. Second upper temporal enlarged, longer than wide, diagonally disposed over a smaller scale contacting sixth supralabial. These two scales are followed by a similarly disposed third pair of temporal scales where the upper scale is the largest. Ear opening absent; its position marked by a longitudinal series of smaller granules. All head scales smooth and juxtaposed. Mental roughly trapezoidal, wider than long, just broader than the ventral surface of rostral. Postmental roughly heptagonal; longer than wide. Two pairs of chin shields, both contacting infralabials; the anterior pair smaller, in broad contact at the midline; second pair in narrower contact at the midline, followed by three pairs of symmetric flat elongate pregulars, inner ones the largest. Five infralabials; first, second and third with about the same size. Gulars smooth, imbricate, rounded posteriorly, in seven transversal rows; scales of gular rows increasing gradually in size toward interbrachial region. Interbrachial region with two central, longer than wide central scales, emarginated at each side by a pair of two smaller scales, the lower one larger. Lateral scales of neck subrectangular, smooth, imbricate, slightly rounded posteriorly and longer than wide, disposed in regular transverse rows and becoming gradually similar to adjacent dorsal or ventral scales. Collar fold absent. Dorsal scales imbricate and disposed in regular transversal rows; smooth, subrectangular and wider in occipital region, becoming progressively narrower, more elongate and rounded towards the level of the forelimbs and then on longer, hexagonal, lanceolate, smooth, with lateral sides almost juxtaposed. Fifty-three transverse rows between interparietal and the level of hind limbs. Lateral scales about the same size as the dorsals but smooth and less acuminate; those closer to ventrals slightly wider. A distinctive area with granular scales surrounds the area of forelimbs insertion and the posterior part of hindlimbs insertion. Twenty five scales around the midbody. Ventral scales smooth, longitudinally imbricate, laterally juxtaposed, almost squared just after the interbrachial row, becoming gradually longer than wide, rounded posteriorly, those after midbody narrower; forty transverse rows between interbrachials and preanals. Four preanal shields, divided in three longitudinal rows, central one with two scales aligned longitudinally; one small scale on each side of anal plate. One preanal pore, femoral pores absent. Scales of tail similar to midbody dorsals, smooth, lanceolate, strongly imbricated, with 121 transverse rows. Fore limbs stiliform, covered by smooth and imbricate scales, ending by three apical scales with no claws. Hind limbs also rudimentary, very reduced in size. Background color of dorsal and lateral surfaces of body and tail dark brown with a pair of dorsolateral yellowish stripes extending from the lateral edge of parietals to the tip of the tail, and two paravertebral faded cream lines beginning at the fifth dorsal row, coalescing just after the hindlimb level and fading. Ventral parts of body and tail brown.

406 · Zootaxa 3636 (3) © 2013 Magnolia Press TEIXEIRA JR. ET AL. FIGURE 3. Detail of the forelimb of (A) Bachia scaea sp. nov. (MZUSP 103414), (B) B. dorbignyi (MZUSP 2063); (C) B. barbouri (MZUSP 46274); (D) B. trisanale abendrothii (MZUSP 3334); (E) B. t. vermiforme (MZUSP 46275); (F) B. peruana (MZUSP 51640); (G) B. bicolor (MZUSP 44957) (H) B. intermedia (MZUSP 40675).

NEW SPECIES OF BACHIA Zootaxa 3636 (3) © 2013 Magnolia Press · 407 FIGURE 4. Dorsal pattern in (A) Bachia barbouri, (B) B. bicolor, (C) B. dorbignyi, (D) B. huallagana, (E) B. intermedia, (F) B. peruana, (G) B. scaea sp. nov. and (H) B. trisanale abendrothii.

408 · Zootaxa 3636 (3) © 2013 Magnolia Press TEIXEIRA JR. ET AL. FIGURE 5. Detail of the forelimb of (A) Bachia scaea sp. nov. (MZUSP 103414), (B) B. dorbignyi (MZUSP 2063); (C) B. barbouri (MZUSP 46274); (D) B. trisanale abendrothii (MZUSP 3334); (E) B. t. vermiforme (MZUSP 46275); (F) B. peruana (MZUSP 51640); (G) B. bicolor (MZUSP 44957) (H) B. intermedia (MZUSP 40675).

NEW SPECIES OF BACHIA Zootaxa 3636 (3) © 2013 Magnolia Press · 409 Measurements of the holotype (mm): SVL=62.2 mm; TL=107.4 mm Variation: the sample is fairly homogeneous in scale counts and dorsal color pattern, however some variation in head scalation can be observed; in three specimens (MZUSP 103408, 103414–15,) the first temporal is followed directly by the second temporal, whereas in the five other specimens (MZUSP 100631, 101586, 101735–36, 103413), there is a small scale between the first and second temporals and the supralabials. Ontogenetic variation on dorsal pattern can also be observed, with the two paravertebral cream lines merging at the cloacal level and entering the tail, fading posteriorly in juveniles; in adults after merging at the cloacal level the line disappears, rendering the dorsal surface of tail dark brown. Comparison with other species (data from species in comparison are given in parentheses): the new species is a member of the Bachia dorbignyi group by having hexagonal, smooth, imbricate dorsal scales; quadrangular, juxtaposed ventral scales; supraoculars absent; interparietal and prefrontal absent; no femoral pores; 1-1 preanal pores; hindlimb stiliform and forelimbs ending in three fingers. These characters promptly distinguish B. scaea sp. nov. from all species of the B. bresslaui group (all with keeled dorsal scales, and supraoculars present), from the B. heteropa group (four digits on limbs and interparietal present) and from the B. flavescens group (quadrangular dorsal scales, supraoculars present). Among the species of the Bachia dorbignyi group (diagnostic features for species of B. dorbignyi group are summarized in Table 1), B. scaea sp. nov. can be promptly distinguished from all species by the absence of clawed fingers (Fig. 3) (2–4 clawed fingers in combination for all other species). Additionally it differs from B. bicolor, B. huallagana, B. peruana, B. talpa and B. trisanale by having 6 supralabials (5) and from B. huallagana and B. trisanale by having 5 infralabials (4). By the absence of contact between supralabial and parietal it differs from B. dorbignyi (in the 5th), B. huallagana (3rd), B. peruana (4th), B. talpa (4th) and B. trisanale (4th). It also differs from B. dorbignyi, B. huallagana, B. peruana and B. trisanale by the presence of a first temporal (absent). By the absence of interparietal it differs from B. barbouri (present) and by the presence of a frontonasal it differs from B. trisanale (absent). The lower number of gulars, 6–7, distinguishes it from B. barbouri (8–10), B. bicolor, B. intermedia (both 7–9), B. talpa (9) and B. trisanale (7–8). By the lower number of scales around the body, 24–26, it can also be distinguished from B. barbouri (26–31), B. bicolor (27–31) and B. intermedia (28–35). By the number of dorsal scales, 51–54, it can also be distinguished from B. talpa (47–51). By the higher number of ventral scales, 42–45, it can be distinguished from B. barbouri (34–39), B. bicolor (34–40), B. intermedia (33–38) and B. talpa (36–38). Although information on the number of caudal scales is missing for most of the species, it can also be distinguished from B. dorbignyi by a higher number of rows, 111–121 (88–108). It can be further distinguished from B. bicolor, B. barbouri and B. talpa by the second chin shield not reaching the oral border (second chin shield reaches the oral border). It can also be distinguished from B. bicolor, B. talpa and B. trisanale by the presence of four preanal shields (three). Also differences in the dorsal color pattern distinguish it from all other Bachia species of the B. dorbignyi group (see Fig. 4 for a summary of dorsal color patterns in the group, excluding B. talpa, for which we did not have data on its dorsal color). Hemipenial morphology: The left hemipenis of MZUSP 103414 (Fig. 5A) is unilobed with the distal tip of the retractor muscle divided. The organ is relatively small, extending along approximately four subcaudal rows (4 mm). The hemipenial body is cylindrical, slightly divided in the distal tip, indicating possibly vestigial lobes. The sulcus spermaticus is a relatively broad channel, originating at the central region of the base of the organ, and proceeding in a straight line towards the apex. At the distal tip of the hemipenial body the sulcus is divided in two shallow branches. On the lobes, these branches run centrifugally ending at the external face of the tip. The full hemipenis is completely nude, with no evident plicae, papillae, ridges, calyces, mineralized spines or spinules in both faces (sulcate and asulcate), even after their immersion on Alizarin Red solution for 24 hours. The complete absence of evident hemipenial ornamentation on the genus Bachia is a common condition shared with other species, such as Bachia trisanale, B. intermedia (Presch 1978), B. oxyrhina (Rodrigues et al. 2008), B. heteropa alleni and B. bresslaui (Nunes 2011). The organ of B. dorbignyi (Fig. 5B) is relatively similar with that of described above for B. scaea sp. nov., differing only in the shape of the hemipenial body, which is more globular, and in the width of sulcus spermaticus, wider than observed in the specimen of B. scaea sp. nov. Distribution and Natural history: despite our large sampling effort in both banks of Madeira River, specimens of Bachia scaea sp. nov. were found only on its left bank (Fig. 6). The additional specimens from Porto Velho municipality were also found at the left bank. In the literature specimens referred to as B. gr. dorbignyi

410 · Zootaxa 3636 (3) © 2013 Magnolia Press TEIXEIRA JR. ET AL. (Avila-Pires 2009) are recorded from a close locality, Guajará-Mirim, at the right bank of Madeira River. We have not examined these specimens, however the dorsal coloration matches that of B. dorbignyi, nonetheless it will be important to examine these specimens to attest if they belong to the new one or B. dorbignyi.

TABLE 1. Summary of diagnostic features among Bachia species of the B. dorbignyi group, derived from examined specimens (see Appendix I) and data from the literature (Cope 1896; Dixon 1973; Presch 1978; Nunes 2011). SAB=scales around body; SL=number of supralabials; SL-P=the supralabial scale contacting parietal; Chin-oral=second chin shield reaching oral border. B. scaea B. B. B. B. B. B. B. B. sp. nov. barbouri bicolor dorbignyi huallagana intermedia peruana talpa trisanale Max SVL 69 mm 68 mm 75 mm 80 mm 73 mm 67 mm 107 mm 65 mm 79 mm Claws none 2 4 3 4 3 3 3 3 Frontonasal present present present present present present present present absent SAB 24–26 26–31 27–31 21–28 24–30 28–35 23–26 25–27 23–30 Dorsal 51–54 46–52 46–53 48–55 47–54 46–52 53–57 47–51 47–55 Ventral 40–43 34–39 34–40 37–42 35–42 33–38 41–45 36–38 34–42 Gular 6–7 8–10 7–9 6–7 7 7–9 6–7 9 7–8 Caudals 111–121 ? ? 88–108 ? ? ? ? ? SL 66565 6555 IL 55554 5554 SL-P none 5th none 5th 4th none 4th 4th 4th 1st temporal present absent present absent absent present absent absent absent Interparietal absent present absent absent absent absent absent absent absent Chin-oral no yes yes no no no no yes no Preanal shields 44344 4433 Hemipenis nude ? flounced nude ? nude ? ? nude

Although no specimen was found active, one individual found during the day in a trail was supposed to be moving under the leaves. One individual was captured in a pitfall trap, while all others were found among the leaf litter and under rotten trunks and fallen logs, through active search. Although we managed to gather six specimens from the sampling areas, given the large effort, it seems to be rare. The environment where B. scaea sp. nov. is found is dominated by a varzea forest (seasonally flooded forest by white-water rivers, such as the Madeira River), in some places with 35m tall trees, with dense leaf litter, crossed by small streams (Fig. 7), with the canopy frequently open due clearings. The altitude is about 90–100m a.s.l., however some higher grounds are also found, reaching up to about 300m a.s.l. Almost nothing is known about its biology, but a pregnant female (MZUSP 101735) carrying two eggs, observed through ventral skin, was found during the rainy season (December or January). Phylogenetic relationships: The final standard deviation of split frequencies on the BA on the concatenate matrix had a very low value (0.002), and also for 16S (0.004) and c-mos (0.006) indicating stationarity. The concatenate dataset recovered Bachia scaea sp. nov. as sister to B. dorbignyi, but the support was not significantly high, in the 16S results they are still recovered as sisters but although higher the support are also not significant; for the c-mos results B. scaea sp. nov. is recovered in a polytomy together with B. intermedia, B. m. monodactylus, B. m. parkeri, B. bicolor, B. barbouri, B. peruana, B. trisanale and B. panoplia, but the support for this polytomy is also low. In fact, the overall support for most of the branches in all topologies are low, although some well supported structure is also observed, such as in the lineage that includes B. huallagana, B. scolecoides and B. heteropa alleni (Fig. 8).

NEW SPECIES OF BACHIA Zootaxa 3636 (3) © 2013 Magnolia Press · 411 FIGURE 6. Distributional records of Bachia species of B. dorbignyi group, including a candidate species, over western- Amazonian Forests and the Andes. Symbols white-outlined represent localities of examined specimens (see Appendix I); black dots represent type localities.

412 · Zootaxa 3636 (3) © 2013 Magnolia Press TEIXEIRA JR. ET AL. FIGURE 7. The varzea forest of the left bank of upper Madeira River, the habitat of Bachia scaea sp. nov. (A), an interior view of the higher lands (B), and its external general view (C).

NEW SPECIES OF BACHIA Zootaxa 3636 (3) © 2013 Magnolia Press · 413 FIGURE 8. Molecular phylogenetic hypotheses for relationship among Bachia species including B. scaea sp. nov. (highlighted in red), based (A) on the concatenate matrix, (B) on the 16S mitochondrial gene and (C) c-mos nuclear gene c- mos). Values on nodes are Bayesian posterior probabilities and Maximum Likelihood bootstraps, respectively. Scale bar indicates substitution per site.

414 · Zootaxa 3636 (3) © 2013 Magnolia Press TEIXEIRA JR. ET AL. Uncorrected p-distances ranged from 1 to 6% for c-mos, and 4 to 12% for 16S among Bachia species (Table 2). Genetic distances between B. dorbignyi and B. scaea sp. nov. were 4% for 16S, and 2% for c-mos.

TABLE 2. Pairwise genetic distances for 16S gene (above) and c-mos (below) among sequences of Bachia analyzed. Values highlighted in bold represented those for the new species. 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 1) B. barbouri 0.00 0.11 0.08 0.06 0.06 0.01 0.07 0.01 0.05 0.02 0.05 0.01 0.08 0.08 0.08 0.08 0.07 2) B. bicolor 0.01 0.11 0.08 0.06 0.06 0.01 0.07 0.02 0.05 0.01 0.05 0.01 0.08 0.08 0.08 0.07 0.06 3) B. bresslaui 0.03 0.03 0.11 0.12 0.12 0.12 0.12 0.12 0.11 0.12 0.11 0.11 0.12 0.12 0.12 0.13 0.12 4) B. dorbignyi 0.02 0.02 0.03 0.10 0.03 0.09 0.03 0.09 0.05 0.09 0.05 0.07 0.04 0.04 0.04 0.04 0.04 5) B. flavescens 0.06 0.05 0.05 0.06 0.09 0.06 0.10 0.06 0.08 0.06 0.09 0.07 0.09 0.09 0.09 0.10 0.10 B. heteropa 6) 0.05 0.05 0.05 0.05 0.04 0.08 0.01 0.08 0.03 0.07 0.03 0.05 0.05 0.05 0.05 0.01 0.03 alleni B. heteropa 7) 0.05 0.05 0.05 0.05 0.04 0.00 0.08 0.00 0.06 0.00 0.06 0.02 0.09 0.09 0.09 0.09 0.08 trinitatis 8) B. huallagana 0.06 0.06 0.05 0.06 0.04 0.00 0.00 0.08 0.03 0.08 0.04 0.06 0.06 0.06 0.06 0.00 0.03 9) B. intermedia 0.00 0.01 0.03 0.02 0.05 0.05 0.05 0.05 0.06 0.01 0.06 0.03 0.09 0.09 0.09 0.09 0.08 B. m. 10) 0.02 0.02 0.03 0.03 0.04 0.04 0.04 0.04 0.01 0.06 0.02 0.04 0.05 0.05 0.05 0.04 0.02 monodactylus 11) B. m. parkerii 0.00 0.01 0.03 0.02 0.05 0.05 0.05 0.05 0.00 0.01 0.06 0.02 0.09 0.09 0.09 0.09 0.08 12) B. panoplia 0.00 0.01 0.03 0.02 0.05 0.05 0.05 0.05 0.00 0.01 0.00 0.04 0.05 0.05 0.05 0.04 0.03 13) B. peruana 0.01 0.01 0.03 0.02 0.05 0.05 0.05 0.06 0.01 0.02 0.01 0.01 0.07 0.07 0.07 0.06 0.06 B. scaea 14) 0.02 0.02 0.04 0.02 0.06 0.06 0.06 0.06 0.01 0.03 0.01 0.01 0.01 0.00 0.00 0.06 0.05 sp. nov. (H1119) B. scaea 15) 0.02 0.02 0.04 0.02 0.06 0.06 0.06 0.06 0.01 0.03 0.01 0.01 0.01 0.00 0.00 0.06 0.05 sp. nov. (H1978) B. scaea 16) 0.02 0.02 0.04 0.02 0.06 0.06 0.06 0.06 0.01 0.03 0.01 0.01 0.01 0.00 0.00 0.06 0.05 sp. nov. (H510) 17) B. scolecoides 0.05 0.05 0.05 0.05 0.04 0.00 0.00 0.00 0.05 0.04 0.05 0.05 0.05 0.06 0.06 0.06 0.04 18) B. trisanale 0.00 0.01 0.03 0.02 0.05 0.05 0.05 0.05 0.00 0.01 0.00 0.00 0.01 0.01 0.01 0.01 0.05

Discussion

The description of Bachia scaea sp. nov. herein ends a long term stasis on the taxonomy of Amazonian Bachia species, especially in the B. dorbignyi group, which had its last species described by Dixon (1973) four decades ago. Avila-Pires (1995) in her comprehensive work on the lizards from the Brazilian Amazonia commented that some Bachia species deserved a more careful examination. Regarding B. peruana, she argued that in a series of specimens from northern Acre state (Cruzeiro do Sul region), Brazil, although some features were coincident with the typical form, there were also other features that overlapped with characters of other species, such as the contact between fifth supralabial and parietal, similar to B. dorbignyi, the absence of frontonasal, similar to B. trisanale (Avila-Pires 1995); and also the absence of clawed fingers, which would distinguish it from other known species of the B. dorbignyi group. Nonetheless, as she examined only a few specimens from that region, she could not evaluate this issue. We have also examined two of three specimens she mentioned, and compared them to the examined B. peruana and Dixon’s (1973) data, who analysed nearlly 40 specimens of this species, and we are positive that this Cruzeiro do Sul population do not represent B. peruana. Indeed they are morphologically intermediate between B. dorbignyi and B. scaea sp. nov. However, as only two specimens (one sub-adult and one juvenile) are available, and variation between them is considerable, such as the presence of frontoparietal in one (ZUEC 436), while it is absent in the other (ZUEC 435) we prefer to wait for additional material in order to decide on their taxonomic status. At that time it should be also important to examine the previously referred specimens

NEW SPECIES OF BACHIA Zootaxa 3636 (3) © 2013 Magnolia Press · 415 from this area (Avila-Pires 2009; Bernarde et al. 2011; SpeciesLink 2011), all referred to as Bachia sp. in our map (Fig. 6). A few individuals from southern Acre (Rio Branco region) were also made available to us. This sample fully matched the diagnostic features of B. trisanale; a lower number of dorsal scales, frontonasal absent, presence of clawed-fingers, anal plate with three shields and the 4th supralabial touching parietal. Although the list of Brazilian reptiles (Bernils & Costa 2012) and IUCN Red List (Lehr & Doan 2010) indicate the presence of B. trisanale in Brazil, they do not have specific data on its occurrence (R. Bernils and E. Lehr per. comm.), thus to our knowledge this is the first unequivocal record of B. trisanale in Brazil. The reported absence of clawed fingers in the northern Acre populations is also observed in all specimens of Bachia scaea sp. nov. In fact, clawed fingers are widespread among species of the B. dorbignyi group, thus its absence in B. scaea sp. nov. distinguishes it from all other species (Fig. 3), although its overall external morphology still resembles B. dorbignyi and B. peruana. Nonetheless, the molecular data is not conclusive on Bachia scaea sp. nov. relationship, as the mitochondrial data suggests a close relationship with B. dorbignyi and the nuclear data place it in a polytomy with several species; thus only further analyses using additional genetic markers may elucidate this question. If confirmed, its relationship with B. dorbignyi will lead to an interesting biogeographical scenario, as both are geographically separated by the Madeira River. This river has already been recognized as a geographical barrier among closely related lineages within several groups (Cracraft 1985; Ayres & Cluttonbrock 1992; Haffer 1992; Avila-Pires 1995; Fernandes et al. 2012; Ribas et al. 2012; Tsuji-Nishikido et al. 2012), and could be associated with the break on the distributional range of the ancestor of B. scaea sp. nov. and B. dorbignyi, leading to their differentiation. Nonetheless a more comprehensive phylogenetic work is still needed to address this matter. The upper Madeira River at the state of Rondônia crosses a lowland area that has been under a massive anthropic pressure. The varzea forests found at its right bank has been progressively anthropized, with the increasing of several destructive activities such as mining and pasturelands for cattle. Moreover, the installation of two hydroelectric dams, currently being built at the Madeira River, will probably transform drastically the landscape in the next few years. Fortunately, the left bank of Madeira River, where B. scaea sp. nov. is found, harbors more preserved forests, and also encompasses a National Park (Parque Nacional do Mapinguari) and higher lands, that will not be reached by the reservoirs, and hopefully some populations will not be affected. The results presented here indicates that the diversity of Bachia at the Brazilian Amazonia is far from being comprehensively known, and it is likely to change in the future through further taxonomic studies on widespread species, and more surveys on poorly sampled areas, which will certainly find that its richness is even higher than currently known.

Acknowledgements

We are grateful to H. Zaher and C. Castro-Mello for access to specimens at MZUSP, Moisés B. de Souza at UFAC and Paulo Manzani at ZUEC. We also wish to thank A. Brito, A.C. Falcão, A.C.M. Leonel, D. Santana, F. Delfim, H.N. Pinto, I. Prates, J. Cassimiro, J. Rodrigues, J.M.B. Ghellere, L.G. Rizatti, L.R.V. Alencar, N.P. Ferreira, M. Gaiarsa, M.A. Sena, M.G. Esteves, P.M.S. Martínez, R.C. Loiola, R. Moretti, R.S. Recoder, R. Graboski and W. Matiazzi for help during the fieldwork, R.C. Amaro for help with molecular matters, Energia Sustentável do Brasil (ESBR) and Arcadis Logos S.A. for support during field work, and for the publication of this manuscript, as this study was part of ‘Programa de Conservação da Fauna Silvestre’, in the 'Plano Básico Ambiental da UHE Jirau'. We are also grateful to I. Prates for the help with the English. IBAMA for the authorization to work in the region (permit CGFAP 260/2010) and Fundação de Amparo à Pesquisa do Estado de São Paulo (FAPESP) and Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq) for financial support.

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Specimens examined Bachia barbouri: PERU: Amazonas: 3 km S junction Bagua Chico & Bagua Grande road, MZUSP 46274. Bachia bicolor: COLOMBIA: Santander: Socorro, MZUSP 44957. Bachia dorbignyi: BRAZIL: Mato Grosso: Aripuanã, MZUSP 81604; Juruena, MZUSP 82421-82424; UHE Guaporé MZUSP 97912–97921. BOLIVIA: Santa Cruz: Buena Vista, MZUSP 2063 (topotype). Bachia huallagana: PERU: Huanuco: Leoncio Prado, Tingo Maria R-147440 (paratype) (photos). Bachia intermedia: PERU: Cajamarca: Pericos, MCZ R-14630 (holotype) (photos), MZUSP 40675. Bachia peruana: PERU: Ayacucho, Rio Santa Rosas entre Pataccocha e San José, MZUSP 51640. Bachia trisanale: BRAZIL: Acre: Rio Branco, UFAC 0026–28; PERU: Madre de Dios: PN de Manu - Est. Biol. Cocha Cashu, MZUSP 46000. Ucayali: Alto Purus, Igarapé Champuia, Alto Curanja, MZUSP 3334. Loreto: Mishana, MZUSP 46275. Bachia sp: BRAZIL: Acre: Cruzeiro do Sul, ZUEC 435–36.

APPENDIX II

Accession number at GenBank for the sequences used in this study.

Species 16S 12S c-mos Bachia scaea sp. nov. (H1119) KC597262 KC597259 KC597265 Bachia scaea sp. nov. (H1978) KC597261 KC597258 KC597264 Bachia scaea sp. nov. (H510) KC597260 KC597257 KC597263 Bachia bresslaui AF420755 - AF420860 Bachia dorbignyi AF420754 - * Bachia flavescens AF420753 - AF420859 Bachia huallagana DQ383224 - DQ383201 Bachia heteropa DQ383222 - DQ383199 Bachia heteropa DQ383213 - DQ383190 Bachia monodactylus parkerii DQ383214 - DQ383197 Bachia barbouri DQ383216 - DQ383193 Bachia peruana DQ383218 - DQ383195 Bachia scolecoides DQ383223 - DQ383200 Bachia monodactylus DQ383220 - DQ383191 Bachia trisanale DQ383221 - DQ383198 Bachia panoplia DQ383219 - DQ383196 Bachia bicolor DQ383217 - DQ383194 Bachia intermedia DQ383215 - DQ383192 Potamites ecpleopus AF420748 - AF420829 Placosoma glabellum AF420742 - AF420833 Cercosaura eigenmanni AF420728 - AF420828 Cercosaura oshaughnessyi AF420750 - AF420852

*Sequence available on Galis et al. (2010).

420 · Zootaxa 3636 (3) © 2013 Magnolia Press TEIXEIRA JR. ET AL. Zootaxa 3646 (2): 200–200 ISSN 1175-5326 (print edition) www.mapress.com/zootaxa/ Erratum ZOOTAXA Copyright © 2013 Magnolia Press ISSN 1175-5334 (online edition) http://dx.doi.org/10.11646/zootaxa.3646.2.10 http://zoobank.org/urn:lsid:zoobank.org:pub:21F9C8C5-55EB-40BE-94FC-E41346DD3DA2

MAURO TEIXEIRA JR, FRANCISCO DAL VECHIO, PEDRO M. SALES NUNES, ANTONIO MOLLO NETO, LUCIANA MOREIRA LOBO, LUIS FERNANDO STORTI, RENATO AUGUSTO JUNQUEIRA GAIGA, PEDRO HENRIQUE FREIRE DIAS, MIGUEL TREFAUT RODRIGUES (2013) A new species of Bachia Gray, 1845 (Squamata: Gymnophthalmidae) from the western Brazilian Amazonia. Zootaxa, 3636(3): 401–420.

Figure 5 caption to read as:

FIGURE 5. Sulcate and asulcate faces of (A) left hemipenis of Bachia scaea sp. nov. (MZUSP 103414). and (B) right hemipenis of B. dorbignyi (MZUSP 97914). Scale bar=1mm.

200 Published: 2 May 2013 Licensed under a Creative Commons Attribution License http://creativecommons.org/licenses/by/3.0