Zootaxa 4000 (4): 401–427 ISSN 1175-5326 (print edition) www.mapress.com/zootaxa/ Article ZOOTAXA Copyright © 2015 Magnolia Press ISSN 1175-5334 (online edition) http://dx.doi.org/10.11646/zootaxa.4000.4.1 http://zoobank.org/urn:lsid:zoobank.org:pub:9D8F0DD1-B28B-4E43-8817-0E165467D68B Description and phylogenetic relationships of a new and two new of from Brazilian Amazonia, with nomenclatural comments on the of (Reptilia: )

GUARINO R. COLLI1,8, MARINUS S. HOOGMOED2, DAVID C. CANNATELLA3, JOSÉ CASSIMIRO4, JERRIANE OLIVEIRA GOMES2, JOSÉ MÁRIO GHELLERE4, PEDRO M. SALES NUNES5, KÁTIA C. M. PELLEGRINO6, PATRICIA SALERNO7, SERGIO MARQUES DE SOUZA4 & MIGUEL TREFAUT RODRIGUES4 1Departamento de Zoologia, Universidade de Brasília, 70910-900 Brasília, DF, 2Museu Paraense Emílio Goeldi/CZO, Caixa Postal 399, 66017-970 Belém, PA, Brazil 3The University of Texas at Austin, Section of Integrative Biology, 1 University Station C0930, 78712, Austin, Texas, USA 4Universidade de São Paulo, Instituto de Biociências, Departamento de Zoologia, Caixa Postal 11.461, 05422-970, São Paulo, SP, Brazil 5Universidade Federal de Pernambuco, Centro de Ciências Biológicas, Departamento de Zoologia, Av. Professor Moraes Rego, s/n. Cidade Universitária CEP 50670-901, Recife, PE, Brazil 6Universidade Federal de São Paulo, Departamento de Ciências Biológicas, Rua Prof. Artur Riedel, 275, 09972-270, Diadema, SP, Brazil 7Department of Biology, Colorado State University, 80523, Fort Collins, CO, USA 8Corresponding author. E-mail: [email protected]

Abstract

We describe a new genus and two new species of gymnophthalmid lizards based on specimens collected from Brazilian Amazonia, mostly in the "arc of deforestation". The new genus is easily distinguished from other Gymnophthalmidae by having very wide, smooth, and imbricate nuchals, arranged in two longitudinal and 6–10 transverse rows from nape to brachium level, followed by much narrower, strongly keeled, lanceolate, and mucronate scales. It also differs from all oth- er Gymnophthalmidae, except , by the presence of two longitudinal rows of ventrals. The new genus differs from Iphisa by having two pairs of enlarged chinshields (one in Iphisa); posterior dorsal scales lanceolate, strongly keeled and not arranged in longitudinal rows (dorsals broad, smooth and forming two longitudinal rows), and lateral scales keeled (smooth). Maximum parsimony, maximum likelihood, and Bayesian phylogenetic analyses based on morphological and molecular data indicate the new species form a clade that is most closely related to Iphisa. We also address several no- menclatural issues and present a revised classification of Gymnophthalmidae.

Key words: , phylogeny, , forest, Conservation, Amazon,

Introduction

The ongoing biodiversity crisis, primarily driven by human activities, could lead to a mass extinction event comparable to the "Big Five" (Bambach 2006; Benton 1995; Raup & Sepkoski 1982) in just a few centuries (Barnosky et al. 2011; Glavin 2007; Leakey & Lewin 1995). Even worse, current estimates of extinction rates may well be seriously biased because a large number of species still awaits formal description (Costello et al. 2013; Dirzo & Raven 2003; May 2011). Therefore, discovering and describing species is a fundamental and necessary step towards biodiversity conservation. Previous analyses indicate that most undescribed species are likely cryptozoic, small-bodied, with small geographic ranges, lower abundance and from less-explored regions, including threatened biodiversity hotspots (Mora et al. 2011; Scheffers et al. 2012). Gymnophthalmid lizards are an exemplary case of a group that may harbour many undescribed species. They

Accepted by A. Bauer: 23 Jun. 2015; published: 18 Aug. 2015 401 are often cryptic, with many species having fossorial or semi-fossorial habits (Colli et al. 1998; Mesquita et al. 2006; Vitt et al. 2003). All species are small-bodied (e.g., Avila-Pires 1995; Vitt & Caldwell 2009) and many have small geographic ranges, such as the sand dunes in the São Francisco Basin in Brazil (Rodrigues 1996). Finally, several species have recently been described from the Atlantic Forest (Rodrigues & Borges 1997; Rodrigues et al. 2005, 2009b, 2013) and Cerrado biodiversity hotspots (Rodrigues et al. 2007a, 2008, 2009a; Teixeira et al. 2013b), as well as from the Amazon basin (Peloso et al. 2011; Rodrigues & Avila-Pires 2005; Teixeira et al. 2013a). Among the 246 recognized species of Gymnophthalmidae, 63 (26%) were described since 2000 (Uetz et al. 2007). Furthermore, additional undetected diversity should be described in the near future, as revealed by recent studies evidencing deep divergences in single widespread species (Nunes et al. 2012; Pellegrino et al. 2011). Herein we describe a new genus and two new species of gymnophthalmid lizards from Brazilian Amazonia. We assess their phylogenetic relationships with other gymnophthalmid genera based on morphological and molecular characters and examine distribution records in light of current trends of deforestation in Brazilian Amazonia. We also address several nomenclatural issues to correct misuses of family-group names within Gymnophthalmidae.

Material and methods

Morphological data. All specimens we examined are deposited in CHUNB (Coleção Herpetológica da Universidade de Brasília), MPEG (Museu Paraense Emílio Goeldi, Belém), and MZUSP (Museu de Zoologia da Universidade de São Paulo). We recorded the snout-vent length (SVL) and tail length of all specimens with a ruler (1 mm precision). We also recorded 77 morphological characters (Appendix S1) pertaining to external morphology, scalation, hemipenis morphology, and osteology following Rodrigues et al. (2005, 2007b, 2009b). We obtained hemipenial characters from the left hemipenis of CHUNB 18738 and CHUNB 50548 and osteological characters from CHUNB 23454 (cleared-and-stained). We prepared the hemipenis following the procedures described by Manzani & Abe (1988), modified by Pesantes (1994) and Zaher (1999). We manually severed the retractor muscle and filled the everted organ with stained petroleum jelly. We stained calcareous hemipenial structures in an alcoholic solution of alizarin red following Uzzell (1973) and Nunes et al. (2012). Terminology of hemipenial morphology follows Dowling & Savage (1960), Savage (1997) and Nunes et al. (2012). We determined the sex of all individuals based on the presence of femoral pores. DNA extraction, amplification and sequencing. We obtained sequence data for one nuclear (c-mos) and three mitochondrial (12S, 16S, ND4) genes of four specimens of the first newly described species (below), from two different localities (CHUNB 18738–18739, 50568–50569). Tissue samples of the second newly described species were not available. We extracted genomic DNA of liver samples with the Viogene Blood and Tissue Genomic DNA Extraction Kit. We used primers and polymerase chain reaction protocols for all genes following Pellegrino et al. (2001). We purified products using the Viogene Gel-M Extraction Kit and performed sequencing in the Core Sequencing Facility at the University of Texas at Austin. We edited sequences using Sequencher 4.8, constructed initial alignments with MUSCLE 3.5 (Edgar 2004) and slightly manually adjusted alignments with MacClade 4.08 (Maddison & Maddison 2005). We excluded 26 bases from the alignment due to homology uncertainty, and after trimming sequence ends used 1884 bases for the combined DNA dataset. Phylogenetic analyses. We downloaded sequences of 69 gymnophthalmids from GenBank representing all major lineages of Gymnophthalmidae (Appendix S2). We chose sequences so as to minimize missing data and selected loci that had been found useful in previous studies (e.g., Castoe et al. 2004; Pellegrino et al. 2001). The ingroup consisted of Gymnophthalminae and the outgroup taxa consisted of atriventris and Ptychoglossus (Alopoglossinae), following Pellegrino et al. (2001). We used morphological data (77 characters) from Rodrigues et al. (2005; 2007b; 2009b) and collected data from the new genus. This dataset included ingroup taxa only from Gymnophthalminae. The outgroup taxa were and Rhachisaurus brachylepis. We performed phylogenetic analyses on three datasets: DNA only, morphology only, and a pruned DNA matrix plus morphology. In the last dataset, we pruned the DNA matrix to retain only taxa for which morphological data were available. The best-fitting model of evolution for the DNA dataset was inferred under the Akaike Information Criterion (AIC) in MrModeltest (Nylander 2004): GTR+I+G for each mitochondrial gene and GTR+G

402 · Zootaxa 4000 (4) © 2015 Magnolia Press COLLI ET AL. for the nuclear gene. We analysed the DNA dataset under five different partitioning schemes: single partition, partition by gene, partition by gene and codon position for ND4 only, partition by gene and codon position for c- mos only, partition by gene and by codon position for both ND4 and c-mos. Using MrBayes 3.1 (Ronquist & Huelsenbeck 2003), we obtained the highest Bayes factor for the partition by gene and codon position for both ND4 and c-mos, and subsequently used it in all analyses (including the combined DNA/morphology datasets). The Bayesian phylogenetic analysis of the DNA dataset was performed using MrBayes 3.1 (10 million generations, sampled every 1,000, with burn-in of 1,000 samples); the partitions and models of evolution were as described above. Support for the tree was assessed using Bayesian posterior probabilities. While fine-tuning the Bayesian analyses using Tracer (Rambaut & Drummond 2007), we found a strong correlation between the proportion of invariable sites (I) and the gamma distribution (G) parameters, as well as low effective sample sizes for both, which prompted us to eliminate parameter I from subsequent analyses. We also found that the Bayesian search overestimated the total tree length relative to maximum likelihood length, which is an occasional problem resulting from an inappropriate use of the branch length priors (Brown et al. 2010; Marshall 2010) and thus we changed the mean of the exponential prior from the default 0.1 to 0.01. We conducted a second set of Bayesian analyses in MrBayes 3.2 using "nst = mixed" rather than "nst = 6" to estimate the rate matrix parameters. This option samples from all the possible reversible substitution models, so it is not necessary to specify the model a priori. Otherwise the analysis parameters were the same. The three resulting rate matrices with the highest posterior probabilities had either four or five different parameters, compared to six in the MrBayes tree estimated using GTR. Thus, the GTR tree was only slightly over-parameterized. The highest posterior density (HPD) tree topology from the nst = mixed analysis was the same as the GTR Bayesian tree except for two poorly supported nodes. We used the GTR tree as the source of posterior probability support for further discussion of results. We conducted maximum likelihood analyses of the DNA matrix with RAxML (Stamatakis 2006) using the models and partitioning schemes as above. Nonparametric bootstrapping (1,000 replicates) was used to assess support. We analyzed the morphological dataset in PAUP* (Swofford 2002) under maximum parsimony with 100 random-tree searches and 1,000 bootstrap repetitions, and in MrBayes 3.1 (Ronquist & Huelsenbeck 2003) under the “standard discrete model with unordered characters (5 million generations, sampled every 1000, burn-in of 500 samples). We used the following parameter values: symdirihyperpr = fixed (infinity), ratepr = variable, rates = gamma, and coding = all. For the combined DNA/morphology datasets (both with all taxa and reduced), we determined the best parsimony tree in PAUP* using 100 searches starting with a random tree, and calculated nonparametric bootstrap support (1,000 bootstrap repetitions). Using the same partitions and parameter values as described above, we performed a MrBayes analysis of the combined DNA/morphology dataset.

Results

Nomenclatural issues with family-group names in Gymnophthalmidae

We reviewed the family-group names in current use for Gymnophthalmidae by reference to the International Code of Zoological Nomenclature (International Commission on International Nomenclature 1999), hereafter the Code. Alopoglossinae. This name was proposed by Pellegrino et al. (2001) to accommodate Alopoglossus Boulenger, 1885 but was not explicitly diagnosed by a list of characters (Article 13.1.1 of the Code). Boulenger (1885) diagnosed Alopoglossus, but did not designate a type species. Burt & Burt (1931) designated buckleyi O'Shaughnessy, 1881 as the type species, without any justification. Later, Peters and Donoso-Barros (1970) indicated A. copii as the type species, also providing no justification. Therefore, Leposoma buckleyi O'Shaughnessy 1881 is the type species of Alopoglossus, by subsequent designation (Article 69.1). Rhachisaurinae. This name was proposed by Pellegrino et al. (2001) to include the monotypic genus Rhachisaurus, but was not explicitly diagnosed by a list of characters (Article 13.1.1). Under Article 13.1.2, a diagnosis may consist of "a bibliographic reference to such a published statement." Although Rhachisaurinae was not explicitly diagnosed in words, Pellegrino et al. (2001) stated "Content: Rhachisaurus, new genus for

NEW LIZARDS FROM BRAZILIAN AMAZONIA Zootaxa 4000 (4) © 2015 Magnolia Press · 403 brachylepis Dixon, 1974." Article 13.5 allows the combined description of a new family name and new genus name as long as the type species is fixed. Thus, Rhachisaurinae is available, with type genus Rhachisaurus, and type species Anotosaura brachylepis Dixon, 1974, the latter by monotypy (Article 68.3). Bachini. This name was proposed by Castoe et al. (2004) to accommodate species of , exclusively. However, the name was published without a statement in words of the characters that purported to differentiate the taxon, i.e. the diagnosis, nor reference to such a publication (Article 13.1). The citation of Gray (1845) after the name Bachia in Castoe et al. (2004, Table 5) does not constitute a reference to a diagnosis, since the latter authors made no reference to statements by Gray (1845) purporting to diagnose the new taxon. In any case, Gray (1845) did not propose a new family-group taxon for Bachia. Recommendation 13C of the Code further enlightens the issue: "Each new nominal taxon should be differentiated from other taxa at the same rank." Therefore, the text of Castoe et al. (2004) cannot be regarded as a statement that purports to provide characters differentiating Bachini (Article 13.1.1) nor do they refer to such a diagnosis (Article 13.1.2). Moreover, no type genus was explicitly designated for Bachini (Article 16.2). Listing Bachia as the sole genus under Bachini does not constitute designation of the type genus, because the formation of a family-group name from an available generic name by "indication" (Article 12.2.4) does not make the name available (Article 13.6.1). Thus, Bachini is a nomen nudum under Article 13 (13.1.1 and 13.1.2) and is, ipso facto, unavailable. The same applies to Bachiinae (Pyron et al. 2013). We diagnose and name this group below, noting that the correct formation of the name is Bachiini or Bachiinae. After our examination of several Greek and Latin lexicons, the etymology of Bachia is unclear. Therefore, under the provisions of Article 29.3.3, we consider Bachi- to be the stem of Bachia. Ecpleopodini. was named by Duméril & Bibron (1839), by the combination of ecpleos (Latin transliteration of the Greek ἔκπλεος), which means complete or full (Liddell & Scott 1889), and pous (Latin transliteration of the Greek πούς), which means foot (Liddell & Scott 1889). Fitzinger (1843) formed the family name Ecpleopoda and subsequent uses and spellings included Ecpleopoda (e.g., Tschudi 1847), Ecpleopidae (Cope 1864), Ecpleopodidae (Cope 1868;1875), Ecpleopinae (Castoe et al. 2004; Pyron et al. 2013), Ecpleopini (Pellegrino et al. 2001), Ecpleopodinae (Rodrigues et al. 2009b) and Ecpleopodini (Peloso et al. 2011). Rodrigues et al. (2009b) credited Frost with pointing out the correct formation of the name Ecpleopodinae, without further justification. The stem of a family name should be based on the name of its type genus, Ecpleopus, after deleting the case ending of the genitive singular (Article 29.3), which in the case of πούς (pous; nominative) is πούς (podos; genitive) (Liddell & Scott 1889). Therefore, Ecpleopoda Fitzinger (1843) should be emended to Ecpleopodidae and the names Ecpleopodinae and Ecpleopodini are correctly formed. Heterodactylini. This name was proposed by Pellegrino et al. (2001) to include Amaral 1933, Boulenger 1887, Spix 1825, Iphisa Gray 1851, and provisionally Boulenger 1888. Rodrigues et al. (2009b) restricted Heterodactylini to Caparaonia, Colobodactylus and Heterodactylus. However, Heterodactylini is not the valid name for this group. Gray (1838) named Chirocolidae to allocate Chirocolus Wagler 1830, a genus created for H. imbricatus Spix 1825. The combination Ch. imbricatus was also used as valid by Schinz (1833), Gray (1845) and Fitzinger (1867), and was considered to be a junior synonym of H. imbricatus by Duméril & Bibron (1839) and Boulenger (1885). In fact, because the type species of Chirocolus, H. imbricatus, is also the type species of Heterodactylus, Chirocolus is a junior objective synonym of Heterodactylus (Article 61.3.3). Even though Chirocolidae was based on a genus name that is a junior synonym, it is available because use of the stem of the genus name is sufficient evidence that the author considered the generic name as valid (Article 11.7.1.1). Even though the genus name Heterodactylus is older than Chirocolus, Chirocolini has priority over Heterodactylini (Article 40.1); therefore, the valid family-group name for the clade including Heterodactylus is Chirocolini Gray 1838. Iphisiini. This name was proposed to include Acratosaura, Alexandresaurus, Colobosaura, Iphisa, and Stenolepis (Rodrigues et al. 2009b), following the splitting of the Heterodactylini (sensu Pellegrino et al. 2001). Iphisiini (Rodrigues et al. 2009b) was published without a statement in words of the characters that purported to differentiate the taxon, i.e. a diagnosis (Article 13.1.1), nor reference to such a publication (Article 13.1.2). Furthermore, the name was incorrectly formed according to Article 29.1. The correct form is Iphisini, because the stem of Iphisa is Iphis-, a name attributed to several individuals in Greek mythology; one of the most popular, according to Ovid's Metamorphoses, was the daughter of Telethusa and Ligdus: raised by her mother as a man to conceal her gender from her father, Iphis was later transformed into a man by the goddess Isis and married Ianthe (More 1922). Authorship of Iphisini should be credited to Gray (1851), who described and diagnosed Iphisadae as

404 · Zootaxa 4000 (4) © 2015 Magnolia Press COLLI ET AL. a family, when describing the genus Iphisa. However, Iphisadae is incorrectly formed and must be corrected to Iphisidae (Article 32.5.3), because the spelling Iphisadae is not in prevailing usage (Article 29.5). Therefore, the correct family-group for the tribe is Iphisini Gray 1851.

Taxonomic descriptions

Genus Rondonops, gen. nov.

Type species. Rondonops biscutatus, sp. nov.

Etymology. Rondon (in homage to Cândido Mariano da Silva Rondon) + ops (from the Latin: power, might, strength, ability). The genus-group name is masculine, according to Article 30.1.4.3. The name refers to the accomplishments of Marshal Rondon (May 5, 1865–January 19, 1958), a Brazilian military officer who dedicated his life to the exploration and integration of remote areas of the Brazilian territory, especially of southwestern Amazonia, and to the peaceful integration of indigenous peoples of Brazil (Conselho Editorial do Senado Federal 2003). In 1910, he was nominated the first director of Brazil's Indian Protection Bureau (SPI/FUNAI) and, in 1914, conducted, with Theodore Roosevelt, a scientific expedition to explore the River of Doubt, now Roosevelt River (Diacon 2004; Roosevelt 1914). The Brazilian state of Rondônia, where we collected the first specimens of Rondonops, was also named after him. Content. Rondonops biscutatus, sp. nov. and R. xanthomystax, sp. nov. Definition and diagnosis. Size moderate, 65 mm maximum SVL; intact tail long, up to about three times SVL. Limbs slender, pentadactyl; first toe reduced, lacking a claw. Ear openings and eyelids distinct. Frontonasal single; prefrontals, frontoparietals, parietals, and interparietal present; parietals longer than wide; three supraoculars. Collar fold absent; three pairs of chinshields, posteriormost reduced. Nuchals very wide, smooth, imbricate, in two longitudinal rows from nape to arm level. Occipitals absent. Dorsal scales posterior to arm level narrower than nuchals, lanceolate, imbricate, strongly keeled, and mucronate. Ventrals identical to nuchals, smooth, imbricate, wider than long, forming two longitudinal rows. Males with a continuous series of pores, with no gap between preanal and femoral pores; females without femoral pores. Rondonops differs from all other genera of Gymnophthalmidae by having an anterior series of smooth, imbricate, and extremely wide nuchals forming two longitudinal and 6–10 transverse rows, followed, posterior of the arm level, by much narrower, strongly keeled, lanceolate, and mucronate scales. It also differs from all other gymnophthalmid genera, except Iphisa, by the presence of only two longitudinal rows of ventrals. The following morphological characters ally Rondonops to Alexandresaurus, Iphisa, Colobosaura, Acratosaura, Stenolepis, Caparaonia, Colobodactylus, and Heterodactylus: interclavicle cruciform, with central area extremely reduced; lateral process of interclavicle long, straight, and pointed; glossohyal fused to basihyal; nasals in narrow contact with premaxilla, wide, divergent, and in contact at midline, but broadly separated anteriorly by the subtriangular lamina of premaxillary; first finger reduced (lost in Heterodactylus and Colobodactylus), without a claw (Rodrigues et al. 2009a). As currently defined, the Chirocolini (former Heterodactylini) contains Caparaonia, Colobodactylus, and Heterodactylus, whereas Alexandresaurus, Acratosaura, Colobosaura, Iphisa, and Stenolepis belong to the Iphisini (Rodrigues et al. 2009a). Our phylogenetic analyses (below) clearly indicate that Rondonops is part of the clade Iphisini. The following unique characters of Iphisini are present in Rondonops: lateral expansions of the parietal and shape of postfrontal. Rondonops and the other Iphisini differ from the Chirocolini by the shape and size of parietal scale, condition of supratemporal fenestra and postorbital width (Fig. 4). In addition to molecular characters, the Iphisini and Chirocolini differ by a more elongate body and higher degree of limb reduction in the latter. Rondonops and the other Iphisini differ from the Gymnophthalmini by possessing eyelids, which are absent in all Gymnophthalmini, except (Avila-Pires 1995; Rodrigues 1991). Among Iphisini, Rondonops differs from Iphisa by having two pairs of enlarged chinshields (one in Iphisa); dorsal trunk scales lanceolate, strongly keeled and not arranged in longitudinal rows (dorsal trunk scales broad, smooth and forming two longitudinal rows), and lateral scales keeled (smooth). It differs from other Iphisini (Alexandresaurus, Colobosaura, Acratosaura, and Stenolepis) and all Chirocolini by the presence of very wide ventrals disposed in two longitudinal rows (ventrals narrower, in four or six longitudinal rows), and 6–10

NEW LIZARDS FROM BRAZILIAN AMAZONIA Zootaxa 4000 (4) © 2015 Magnolia Press · 405 transverse series of smooth, wide nuchals disposed in two longitudinal rows (nuchals narrower, not forming two longitudinal rows). Rondonops further differs from Stenolepis by having prefrontals (absent), and from Acratosaura, by having only two pairs of enlarged chinshields (three). Among Chirocolini, Rondonops differs from Colobodactylus and Heterodactylus by having a distinct first finger (absent), prefrontal scales (absent), and parietals longer than wide (wider than long). From Heterodactylus it differs by having an enlarged interparietal (absent or vestigial), a distinct ear opening (absent), and a typical lacertiform body (extremely elongate). Rondonops differs from Caparaonia by having only two pairs of enlarged chinshields (three).

Rondonops biscutatus, sp. nov. (Figs. 1–2)

Gymnophthalmidae sp.—Gainsbury and Colli (2003):509. Colobosaura sp. nov.—Hoogmoed et al. (2007):147, 151, 152. Gymnophthalmidae sp.—Garda et al. (2013):247, 248.

Holotype. CHUNB 18739 (field number GRCOLLI 06106); adult male; from Pimenta Bueno (11°35’S, 61°10’W), RONDÔNIA, BRAZIL; leg. G. R. Colli, F. G. R. França, A. M. Gainsbury, A. A. Garda and H. C. Wiederhecker; 1 August 2000. Paratypes. BRAZIL: MATO GROSSO: Alta Floresta: Parque Estadual do Cristalino (9°34'38.77"S, 55°55'18.49"W): CHUNB 47042, leg. J. P. Caldwell, G. R. Colli, F. G. R. França, D. L. P. Leite, D. B. Shepard, M. M. Vasconcellos and L. J. Vitt, 11 November 2005. Pará: Tapajós region, Jacareacanga-Itaituba: MPEG 31102, leg. Team UFPA/Herpetologia; Itaituba, APA Tapajós, Mina do Tocantinzinho (6°02'45.41"S, 56°18'13.38"W): MPEG 28555, leg. A. Lima, A. Araújo and S. R. dos Anjos, 23 October 2010; (6°4'58.68"S, 56°15'14.34"W): MPEG 28556, leg. J.O. Gomes and F. Chagas, 13 August 2010; (6°3'48.00"S, 56°16'58.68"W), MPEG 28557, leg. J. O. Gomes and F. Chagas, 9 August 2010; Mina do Palito (6°18'48.9"S, 55°47'02.7"W): MPEG 28558, leg. J. Gomes and A. D'Angiolella, 19 November 2010; (05°13'49.18"S, 56°55'91"W): MPEG 31095, leg. Team UFPA/ Herpetologia, 28 October 2013; (5°17'57.01"S, 56°58'52.57"W) MPEG 31096, leg. Team UFPA/Herpetologia, 11 January 2013; (5°22'41.99"S, 56°54'50.65"W): MPEG 31098, leg Team UFPA/Herpetologia, 16 August 2013; Jacareacanga (05°45'59.9”S, 57°1714.6”W): MPEG 31097, 31099, 31100-01, all leg. Team UFPA/Herpetologia, 23 January, 21–22 August 2013; Novo Progresso (7°08.061'S, 55°24.888'W): MPEG 24127–24130, leg. M. S. Hoogmoed, M. A. Ribeiro Jr., C. O. Araujo and D. G. Nascimento, 22–26 November 2005. RONDÔNIA: Alta Floresta d’Oeste, Parque Estadual do Corumbiara (12°54'21.10"S, 62°4'6.90"W): CHUNB 52868, leg. R. J. Bosque and G. R. Colli, 9 March 2008; Cerejeiras, Parque Estadual do Corumbiara (13°6'47.87"S, 61°28'37.65"W): CHUNB 50539–50569, leg. R. J. Bosque and G. R. Colli, August 2007; Guajará-Mirim (10°46’S, 65°20’W): CHUNB 23454–23458, leg. G. R. Colli, G. C. Costa, A. M. Gainsbury, A. A. Garda, F. P. Werneck and H. C. Wiederhecker, December 2000–January 2001; Pimenta Bueno (11°35’S, 61á10’W): CHUNB 18738, leg. F. G. R. França, A. M. Gainsbury, A. A. Garda and H. C. Wiederhecker, 31 July 2000. Etymology. The specific epithet is an adjective formed from Bi (from the Latin: two) and scutatus (from the Latin: shield-shaped). The name refers to the arrangement of nuchals in two longitudinal rows, characteristic of the genus, but first noted in this species. Diagnosis. Body robust; tail up to 3 times longer than body. Limbs pentadactyl, slender; first finger lacking claw. Ear openings and eyelids distinct. Frontonasal single; prefrontals, frontal, frontoparietals, parietals and interparietal present; parietals longer than wide. Collar fold absent. Three pairs of chinshields; anteriormost two enlarged, posteriormost reduced. Three supraoculars, anteriormost smallest. Dorsals (including nuchals) in 26–32 rows; anteriorly smooth, wide, imbricate, with rounded posterior margins, in two longitudinal and 6–10 transverse regular rows; posteriorly to arm level becoming progressively narrower, mucronate, with broad and flat keels, and then lanceolate, strongly keeled, imbricate and mucronate. Occipitals absent. Ventrals very wide, smooth, imbricate, in two regular longitudinal and 14–20 transverse rows, identical in size and shape to nuchals. Scales around midbody 23–30; subdigital lamellae under finger IV and toe IV, respectively 11–15 and 16–20. Fingers and toes short and robust. Males with a continuous series of 17–22 pores, with no gap between preanal and femoral pores; femoral and preanal pores absent in females. Description of holotype (Fig. 1). Adult male, in good state of preservation. Snout-vent length 56 mm, intact

406 · Zootaxa 4000 (4) © 2015 Magnolia Press COLLI ET AL. tail length 170 mm. Rostral broad, wider than high, contacting first supralabials, nasals, and frontonasal. Frontonasal pentagonal, wider than long, contacting rostral, nasals, and prefrontals. Prefrontals wider than long, in contact at midline and contacting frontonasal, nasals, loreals, first supraoculars (and second supraocular on the right side) and frontal. Frontal hexagonal, with parallel lateral margins, longer than wide, anteriorly indenting prefrontals and, posteriorly, frontoparietals. Frontoparietals roughly pentagonal, as wide as prefrontals, in broad contact, strongly indented by interparietal, in contact with second and third supraoculars and parietals. Interparietal longer than wide, longer and narrower than frontal, as long as and narrower than parietals. Parietals heptagonal; bordered laterally by three enlarged, longer than wide, temporals; bordered anteriorly by third supraocular and frontoparietal, medially by interparietal, and posteriorly by first dorsals. Posterior margin of parietals and interparietal contacting first row of nuchals. Three supraoculars; first smallest; second largest, with longest suture with frontal, narrowly contacting frontoparietals and right prefrontal; third about the same size as frontoparietals, rounded posteriorly, in broad contact with frontoparietal, parietal, and temporal. Nasal dorsal to first supralabial, large, slightly longer than high; nostril in the middle of lower part of nasal, indenting suture with first labial. Loreal posterior to nasal, higher than long, diagonally oriented; contacting nasal, prefrontal, first supraocular, first superciliary, preocular, frenocular, and first and second supralabials. Frenocular small, below preocular, followed posteriorly by four suboculars; second and third suboculars elongate; fourth longer, followed by slightly smaller postoculars. Seven supralabials; suture between third and fourth below the center of eye; fifth largest, contacting third and fourth subocular; seventh smallest, contacting granules surrounding anterior margin of ear. Three superciliaries; first largest, higher anteriorly, longer than first supraocular, contacting first and second supraoculars, loreal, preocular, second superciliary and upper eyelid; second superciliary smallest, contacting second supraocular. Enlarged quadrangular scale posterior to third superciliary and contacts postocular. Central part of lower eyelid with semitransparent, undivided disc surrounded by small, slightly pigmented, granular, smooth scales and eleven moderately pigmented palpebrals. Lower eyelid with eleven moderately pigmented palpebrals. Temporals smooth, juxtaposed, irregular in size and shape, largest about the same size of sixth supralabial. Ear opening surrounded by series of very small, juxtaposed, rounded granules; external auditory meatus shallow; tympanum distinct, subovoid. Scales on sides of neck in approximately 12 irregularly transverse series between ear and arm level; those next to ear small, smooth, rhomboid, becoming gradually more elongate, mucronate, striated and imbricate near arm. All head scales smooth and juxtaposed, with many scattered sensorial pits.

FIGURE 1. Rondonops biscutatus, holotype, CHUNB 18739, adult male, SVL 56 mm. (A) Dorsal view of head; (B) lateral view of the head; (C) ventral view of the head; (D) precloacal plate and femoral pores at each side.

NEW LIZARDS FROM BRAZILIAN AMAZONIA Zootaxa 4000 (4) © 2015 Magnolia Press · 407 FIGURE 2. (A) Flooded forest in Alta Floresta d'Oeste, Rondônia, Brazil, depicting termite nests on tree trunks, used as shelter by Rondonops biscutatus during the wet season. (B) Flooded forest in Cerejeiras, Rondônia, Brazil, during the dry season, depicting array of pitfall traps with drift fences, used to capture R. biscutatus. (C) Rondonops biscutatus, adult female. (D) Rondonops biscutatus, adult male.

Mental broad, wider than long. Postmental heptagonal, wider than long, contacting first and second infralabials. Two pairs of enlarged chinshields, in broad contact along midline; first contacting second to fourth infralabials; second largest, contacting fourth and fifth infralabials. Third pair of chinshields reduced, narrow, chevron-like, laterally contacting a scale slightly longer than wide, about the same size as fifth supralabial, separated from sixth infralabial by elongate scale. Six infralabials; third, fifth, and sixth largest, subequal in size. Gulars enlarged, wider than long, smooth, imbricate, most rounded posteriorly and a few with irregular margins, in two longitudinal and seven transverse rows; scales in first row smaller. Interbrachial region distinct, with seven smooth, strongly imbricate scales; central scale subtriangular, contacting larger scales laterally; lateral interbrachials smaller, longer than wide. Collar fold absent. Nuchals large, wider than long, smooth, imbricate, rounded posteriorly, in two longitudinal and nine transverse, regular rows between parietals/interparietal and arm level. Occipitals absent. Dorsals becoming progressively narrower, imbricate, mucronate, lanceolate, and strongly keeled posteriorly of forelimb level; keels appearing gradually, broad and flat anteriorly, becoming thin, high and sharp posteriorly. Twenty-nine transverse rows of dorsals between parietals/interparietal and posterior level of hind limbs. Scales on flanks smaller and more diagonally arranged than dorsals; smooth, wide, imbricate, and posteriorly rounded in the row bordering ventrals, becoming progressively narrow, elongate, keeled, and mucronate dorsally. Distinctive area with small, smooth, and rounded granules around arm insertion. Twenty-four scales around midbody. Ventrals smooth, imbricate, wider than long, rounded posteriorly, in two longitudinal and 17 transverse rows from interbrachials to preanals. Five preanals; central one rhomboidal (posterior part wider), not reaching preanal border, which is formed by two enlarged scales in broad median contact and two smaller external paramedials. Pores 20 (total), opening in centre of scales, continuous; no gap between femoral and preanal pores. Dorsal scales on tail elongate, lanceolate, keeled, strongly imbricated, smaller than midbody dorsals; ventral scales on tail smooth, rounded and enlarged near preanal area, becoming gradually keeled, elongate, lanceolate; dorsals and ventrals undifferentiated near tail tip.

408 · Zootaxa 4000 (4) © 2015 Magnolia Press COLLI ET AL. FIGURE 3. (A) Sulcate, (B) lateral and (C) asulcate faces of the right hemipenis of Rondonops biscutatus (CHUNB 18738).

Forelimbs with large, smooth, imbricate scales; scales on ventral part of upper arm smaller, rounded, rhomboid, juxtaposed; scales on ventral part of forearm almost granular. Anterior and ventral parts of hind limbs with irregularly large, smooth, imbricate scales, identical to those on corresponding parts of forelimbs, except for some on posterior dorsal part that are keeled. Scales on posterior part of hind limbs granular, juxtaposed, becoming larger, imbricate, and keeled on anterior part of tibia and femur. Carpal and tarsal scales large, imbricate, smooth; supradigital lamellae smooth, imbricate. Palmar and plantar surfaces with smooth, small, tuberculate granules. Fingers and toes relatively short and robust. First finger cylindrical, distally only slightly compressed laterally. First two-thirds to three-quarters of the 14 subdigital lamellae of finger IV single, bulbous, rounded, sometimes separated by a longitudinally divided rounded tubercle from the most distal scales, the latter being mostly single and smooth. Toe IV laterally compressed over its entire length, 19 subdigital lamellae, proximal six or seven bulbous, followed by seven or eight pairs of smaller bulbous scales that are arranged in zigzag pairs, distal scales single, smooth, narrow with a median keel. Toes and fingers with claws, except finger I; relative sizes: 1 < 2 = 5 < 3 = 4 and 1 <2 < 5 < 3 < 4, respectively. Dorsal surfaces of body and tail and lateral part of tail dark brown with irregularly distributed dark brown dots, generally concentrated on anterior part of some dorsal scales (Fig. 2C–D). Flanks dark brown due to more intense pigmentation in anterior part of scales, becoming more conspicuous laterally. This darker pattern strongly mottled with scattered, cream yellow spots concentrated on central part of lateral scales, and extending to hind limb level. Lateral parts of head with similar pattern, with irregular dark brown blotches concentrated in central parts of supra- and infralabials, with yellow dominating in sutures. Ventral parts of body cream-yellow, immaculate. Ventral part of tail dark-brown near its tip. Tail dark brown dorsally, lighter ventrally. Limbs dark brown dorsally, irregularly mottled with cream yellow pattern similar to flanks; ventrally cream yellow, immaculate. Description of hemipenis (Fig. 3). We prepared the left hemipenis of two paratypes (CHUNB 50548, 18738). In normal position (retracted), the organ is up to 7 mm long, extending for about six subcaudal rows. Hemipenial body roughly globular, with a slight median constriction and clearly bilobate, ending in two pronounced lobes with approximately one-third of total length of the organ. Lobes noncapitated; apex ornamented with a few small folds. Sulcus spermaticus in midline of sulcate face, extending straight from base of organ to lobes. At distal part of

NEW LIZARDS FROM BRAZILIAN AMAZONIA Zootaxa 4000 (4) © 2015 Magnolia Press · 409 hemipenial body, the sulcus is divided by a small fleshy fold at base of a lobular crotch in two branches, each running on medial surface of lobes and ending in their tip among lobular folds. Two large naked areas parallel to sulcus spermaticus in sulcate face of hemipenial body. Each naked area bordered externally by isolated longitudinal ornamented area composed by single spines or series of spicules, arranged in about 45 transverse rows from base of organ to about second third of each lobe. Basal region of longitudinal ornamented area with about 10 single enlarged spines, the latter gradually giving way to more complex transverse rows with up to 10 small and bicuspidate spicules. Lateral face of hemipenis adorned by about 40 transverse rows of spicules; the more apical bi- or tricuspidate, basal ones longer, unicuspidate, enlarged. Lateral series of spicules separated from those on sulcate face by narrow longitudinal nude area extending from base to lobes of organ. Another bare area, curved and wider medially, separates the 20 more basal rows of lateral spicules of ornamented asulcate face of organ. Asulcate face of hemipenis adorned by two longitudinal ornamented areas, separated from each other by bare sagittal area running from base towards lobular crotch, and then bifurcating along medial area of lobes. This sagittal bare area is slightly wider in CHUNB 50548. Ornamented areas of asulcate face composed by about 30 rows of spines and/or spicules; spines of these rows bordering the bare area enlarged, hook-shaped, gradually decreasing in size towards sulcate face. Spicules of superior third of asulcate side bi- or tricuspidate. Osteological description (Figs. 4–5). Premaxillary as long as large, touching but not articulating with the maxillary laterally. Its dorsal lamina triangular posteriorly, long, covering the nasals slightly anteriorly the nasals and deeply indenting their suture, preventing their anterior contact. Thirteen conical premaxillary teeth. Nasals large, slightly longer than wide, wider anteriorly, diagonally arranged, widely separated anteriorly, in midline contact in posterior third, covering the frontal anteriorly. Frontal longer than wide, strongly constricted between orbits, wider posteriorly, covering parietal and articulating laterally with it by a pair of frontoparietal tabs. Parietal longer than wide, wider and concave posteriorly, covering occipital region laterally. Lateral expansion of parietal absent, leaving supratemporal fenestra open. Epipterygoid contacting superficially a descending epipterygoid process of parietal. Maxillary contacting nasal dorsally, parts of frontal and lacrimal laterally, but not overlapping, and extensively covering prefrontal and jugal; 24 maxillary teeth. Prefrontal large, its posterior process long but not reaching level of middle of orbit; in broad contact with frontal. Lacrimal small, rod shaped, very conspicuous, contacting prefrontal and maxillary along the inferoanterior part of orbit. Postfrontal and postorbital single. Postfrontal roughly triangular, contacting jugal, frontal, postorbital and parietal, closing the orbit posteriorly. Posterior part of postfrontal wider, longer, almost straight, preventing contact between frontal/parietal and postorbital and concealed marginally by postorbital. Postorbital long and wide, slightly expanded, contacting posteriorly squamosal but leaving supraorbital fenestra widely open. Squamosal long, posteriorly curved and articulating with dorsal end of quadrate. Supratemporal fenestra widely open. Supratemporal present, small, in close contact with posterior part of parietal and squamosal. Fifteen scleral ossicles. Vomer, palatine, pterygoid and ectopterygoid present. Vomer, palatine, premaxillary and maxillary in contact, restricting fenestra exochoanalis. Infraorbital fenestra large, bordered posteriorly by ectopterygoid and pterygoid. Pterygoid teeth present. Stapes rod-like, wider and rounded at the base. Sutures between supraoccipital, exoocipital, basioocipital and otic area of skull not clearly visible in articulated skeleton, as well as those between basioccipital and basisphenoid. Dentary, articular, splenial, angular, and supraangular distinct; 25 dentary teeth, conical anteriorly, bicuspid or tricuspid posteriorly. Glossohyal long, fused to basihyal. First ceratobranchial curved posteriorly; hypohyal and ceratohyal present. A second short pair ceratobranchials present and positioned parallel to anterior part of trachea. Anterior part of clavicle greatly enlarged, flattened, enclosing a fenestra. Interclavicle long, cruciform, with very long lateral processes reaching sternum but not sternal fenestra. Scapulocoracoid with coracoid, scapular and scapulocoracoid fenestra; suprascapula present. Sternum with large fenestra invaded by long sternal process; three sternal ribs; xiphisternum with two inscriptional ribs. Ilium, ischium and pubis present, the latter with a conspicuous pectinate apophysis. Hypoischium long, larger at the base, almost reaching preanal border; preischium small, elongate; prepubis small, quadrangular and ossified. Twenty-seven procelous presacral vertebrae, neural spines low, higher anteriorly hypapophyses present in first eight vertebrae; zygantrum-zygosphene present. Last presacral vertebra lacking ribs. Two sacral vertebrae. First four caudal vertebrae lacking autotomic processes, with long and wide transverse processes and wide and high neural spines. From fifth vertebra on, intravertebral autotomic septa present, transverse processes narrow and neural spines decreasing in height. Humerus and femur slightly longer than radius and ulna, and tibia and fibula, respectively. Remaining elements of forelimbs and hind limbs as in Fig. 5.

410 · Zootaxa 4000 (4) © 2015 Magnolia Press COLLI ET AL. FIGURE 4. (A) Dorsal and (B) ventral views of the skull of Rondonops biscutatus (CHUNB 23454). Scale bar = 1 mm.

NEW LIZARDS FROM BRAZILIAN AMAZONIA Zootaxa 4000 (4) © 2015 Magnolia Press · 411 FIGURE 5. (A) Right hand, (B) hyoid, (C) right foot, (D) pectoral, and (E) pelvic girdles of Rondonops biscutatus (CHUNB 23454). Scale bars = 1 mm.

Variation and sexual dimorphism. Variation in external morphology is summarized in Table 1. All specimens have three supraoculars and three superciliaries, except CHUNB 52868 where the first and second superciliaries are fused on the right side. Most specimens have 6 or 7 supralabials and infralabials; variation is due to fusions or subdivision of scales and frequently asymmetric. Likewise, most specimens have five suboculars, although the number may vary between three and seven, frequently asymmetrically in the same individual. Gulars also vary between six and eight. Other variations include: the loreal and frenocular are fused on the left side in MPEG 24128; four specimens (CHUNB 50551, 50545, 50553, 50563) have a small azygous scute behind frontal that separates frontoparietals (not in CHUNB 50563); the central scale of the preanal plate is absent in three specimens (CHUNB 50549, 50557, 50565); in CHUNB 50552 the semitransparent disc of the left lower eyelid is divided medially; in CHUNB 23458 prefrontals are not in contact; and CHUNB 50545 has anomalous chinshields. The sexes can be readily separated by the absence of preanal and femoral pores in females. Pores are highly conspicuous, placed in distinctively elevated scales, and aligned on each side without gaps between preanal and femoral. In addition to the presence of pores, there is sexual dimorphism in the number of ventrals, which are

412 · Zootaxa 4000 (4) © 2015 Magnolia Press COLLI ET AL. significantly more numerous in females (Table 1, t-test p < 0.05), and adult males have bright orange bellies. In one male (MPEG 24129), a distinct white ocellus is present above the insertion of the hind limb. Ventral parts in life are pale reddish orange, in one male (MPEG 24129) with vertical, pale reddish orange bands reaching dorsolateral area. Tongue is dark grey.

FIGURE 6. Geographic distribution of Rondonops biscutatus (triangles) and R. xanthomystax (circles). 1,2: Borba and Nova Olinda do Norte, 3: Novo Progresso, 4: Alta Floresta, 5: Guajará-Mirim, 6: Pimenta Bueno, 7: Alta Floresta d'Oeste, 8: Cerejeiras, 9: Itaituba, 10: Jacareacanga. AM: Amazonas, MT: Mato Grosso, PA: Pará, RO: Rondônia.

Distribution, and natural history. Rondonops biscutatus is known only from forests in southwestern Amazonia in the states of Rondônia, Mato Grosso and Pará (Fig. 6). This region is part of the "arc of deforestation" (Aldrich et al. 2012; Fearnside & Graça 2006; Ferreira et al. 2005). Specimens from Parque Estadual do Corumbiara (Cerejeiras and Alta Floresta d'Oeste) were collected in seasonally flooded forest (Fig. 2ab); specimens from Guajará-Mirim and Parque Estadual do Cristalino (Alta Floresta) were collected in terra firme forest. Specimens from Pimenta Bueno were collected in Cerrado enclaves within terra firme forest. All these individuals were collected in pitfall traps (Fig. 2b), consisting of four 30-liter buckets, arranged in a Y, separated by 6 m drift fences, except CHUNB 52868 that was hand collected at 13:23 h. This individual was in a flooded forest, on top of a termite nest at the side of a large tree, ca. 60 cm above the water (Fig. 2a). Specimens from Novo Progresso were collected in logged primary terra firme forest with many Bertholletia excelsa (Brazil nut) trees. One specimen from Itaituba (Mina do Tocantinzinho) was collected at the margin of a creek in an açaí (Euterpe oleracea) forest. The other specimens from Itaituba (Mina do Tocantinzinho and most material collected by the UFPA/Herpetologia Team) were collected in terra firme forest. MPEG 31101 from Jacareacanga and MPEG 31098 from Itaituba were collected in riparian plots. MPEG 24128–9 and 31095–31102 were collected in Y-shaped pitfall traps consisting of four 60-l iter buckets placed 10 m apart and connected by 50 cm high drift fences. MPEG 24127 and 24130 were collected by hand during active searching in leaf litter of terra firme forest at 10:20 and 11:25 h. MPEG 28555–6, 28558 were all collected by hand in leaf litter, between 9:00 and 13:50 h, MPEG 28557 also was collected by hand in leaf litter, but at 22:15 h. Thus, R. biscutatus is an inhabitant of forest floor leaf litter in terra firme forest, transitional areas between Amazon forest and Cerrado, açaí forest, riparian areas and seasonally flooded forest. Specimens seem to be mostly diurnal (9:00–13:50 h), but one specimen was caught while actively

NEW LIZARDS FROM BRAZILIAN AMAZONIA Zootaxa 4000 (4) © 2015 Magnolia Press · 413 moving in leaf litter at night (22:15 h), indicating some nocturnal activity as well. Three females (CHUNB 50544, 50562, 50563) from Cerejeiras, Rondônia, collected in August 2007 (dry season) contained one egg each, whereas none of the adult females collected during the wet season contained eggs. Presumably reproduction takes place during the dry season. Remarks. The species was first mentioned by Gainsbury and Colli (2003) as Gymnophthalmidae sp., from Cerrado enclaves on latosols and sandy soils in Pimenta Bueno, Rondônia. Hoogmoed et al. (2007) reported the species as Colobosaura sp. nov., but provided no further details except the locality of collection and its general habitat (terra firme forest). Garda et al. (2013) studied the effects of microhabitat variation on distribution in a terra firme forest in Guajará-Mirim, Rondônia. The species, reported as Gymnophthalmidae sp., is associated with sites distant from large trees, with few fallen logs and burrows, less canopy cover, thicker understory, thinner leaf litter, and numerous termite nests.

Rondonops xanthomystax, sp. nov. (Figs. 7–8)

Holotype. MZUSP 98085 (field number MTR 12977); adult female; from left bank of Rio Abacaxis, São Sebastião, Borba (4°18'32"S, 58°38'11"W), Amazonas, BRAZIL; leg. M. T. Rodrigues, J. Cassimiro, S. M. Souza and J. M. Ghellere, 12 January 2007. Paratypes. BRAZIL: AMAZONAS: Nova Olinda do Norte: Igarapé-Açu, right bank of Rio Abacaxis (4°20'39"S, 583°8'06"W): MZUSP 99243, 6 January 2007; Borba: São Sebastião, left bank of Rio Abacaxis (4°18'32"S, 58°38'11"W): MZUSP 99244–99246, 99248, 12–25 January 2007; Borba: Palhalzinho, left bank of Rio Abacaxis (4°18'06"S, 58°38'01"W): MZUSP 99247, 25 January 2007. All by the same collectors as for the holotype. PAR: Itaituba (5°27'19.22"S, 57°4'14.09"): MPEG 31109, leg. Team UFPA/Herpetologia, 31 August 2013; Jacareacanga (6°06'2241"S 57°36'19.91"W): MPEG 31103, 31107, 31110, leg. Team UFPA/Herpetologia, 30 September 2012, 27 January and 24 August 2013; (5°27'15.84"S, 57°4'41.02"W): MPEG 31104, leg. Team UFPA/Herpetologia, 13 October 2013; (5°48'03.1"S 57°24'23.4"); MPEG 31105–06, leg. Team UFPA/ Herpetologia, 8 October 2013; (5°44'18.7"S 57°21'18.8"W): MPEG 31108, leg. Team UFPA/Herpetologia, 11 January 2013. Etymology. The specific epithet is a noun derived from xanthos (Latin transliteration of the Greek ξανθός: yellow) and mustax (Latin transliteration of the Greek μύσταξ: upper lip or moustache) (Liddell & Scott 1889). The name refers to the yellow (in life) upper lip of this species, which strongly differs from that of its congener, where it is strongly mottled with dark brown. Diagnosis. Body robust; tail up to 2.7 times longer than body. Limbs pentadactyl, slender; first finger lacking claw. Ear openings and eyelids distinct. Frontonasal single; prefrontals, frontal, frontoparietals, parietals and interparietal present; parietals longer than wide. Collar fold absent. Three pairs of chin shields; third pair reduced. Three supraoculars, anteriormost smallest. Dorsals (including nuchals) in 26–30 rows; nuchals multistriate, wide, imbricate, disposed in two longitudinal and 5–8 transverse regular rows; posterior of forearm insertion becoming progressively narrower, mucronate, with broad and flat keels, and then lanceolate, strongly keeled, imbricate, and mucronate. Occipitals absent. Ventrals very wide, smooth, imbricate, in two regular longitudinal and 15–18 transverse rows, identical in size and shape to nuchals. Scales around midbody 25–28; subdigital lamellae of finger IV and toe IV, respectively 13–17 and 20–26. Males with a continuous series of 19–24 pores, with no gap between preanal and femoral pores; femoral and preanal pores absent in females. Rondonops xanthomystax differs from R. biscutatus by having two longitudinal rows of 5–8 multistriate nuchals (6–10, smooth); scales on sides of neck keeled (smooth); 13–17 and 20–26 infradigital lamellae under finger IV and toe IV, respectively (11–15 and 16– 20); relatively longer fingers and toes. In R. xanthomystax, a wide black stripe covers all of the lateral surface of the head above the upper part of supralabials and extends from nasal to the insertion of the forearm, where it merges with the flank colour; below the black stripe, a bright orange-yellow colour covers the larger part of supralabials, infralabials, and ventral parts of head and throat. In R. biscutatus, the lateral dark stripe is absent and supralabials are strongly mottled with dark brown. Description of the holotype (Fig. 7). Adult female, in good state of preservation. Snout-vent-length 67 mm; tail regenerated, total length 68 mm, length of intact portion 62 mm. Rostral broad, wider than high, contacting first supralabials, nasals and frontonasal. Frontonasal heptagonal, wider than long, contacting rostral, nasals, loreal, and

414 · Zootaxa 4000 (4) © 2015 Magnolia Press COLLI ET AL. FIGURE 7. Rondonops xanthomystax, holotype, MZUSP 98085, adult female, SVL 67 mm. (A) Lateral view of head; (B) ventral view of the head; (C) dorsal view of the head.

NEW LIZARDS FROM BRAZILIAN AMAZONIA Zootaxa 4000 (4) © 2015 Magnolia Press · 415 prefrontals. Prefrontals slightly wider than long, in broad contact at midline and contacting frontonasal, loreal, first and second supraoculars and frontal. Frontal hexagonal, with posteriorly, slightly convergent lateral margins, longer than wide, slightly wider anteriorly; anteriorly indenting prefrontal and, posteriorly, frontoparietals. Frontoparietals pentagonal, as wide as prefrontals, in broad contact, strongly indented by interparietal, in contact with second and third supraoculars and parietals. Interparietal longer than wide, slightly longer than and as wide as frontal, as long as and narrower than parietals. Parietals heptagonal; bordered laterally by three enlarged temporals (the second being largest), anteriorly by third supraocular and frontoparietal, medially by interparietal, and posteriorly by first row of nuchals. Posterior margin of parietals and interparietal rounded, contacting first row of nuchals. Three supraoculars; first smallest; second largest, forming a large suture with frontal, narrowly contacting frontoparietal and prefrontal; third supraocular larger than frontoparietals, in broad contact with frontoparietal, parietal and temporal. Nasal dorsal to first supralabial, large, slightly longer than high; nostril in middle of lower part of nasal, indenting suture with first labial. Loreal posterior to nasal, narrower and diagonally oriented; contacting nasal, frontonasal, prefrontal, first supraocular, first superciliary, preocular, frenocular, and first supralabial and second supralabials. Frenocular small, ventral to preocular, followed posteriorly by six suboculars, mostly elongate and about the same size; sixth subocular longest, almost square, followed by postocular. Seven supralabials; fourth below center of eye; fifth largest, contacting posterior suboculars; seventh supralabial smallest, contacting granules surrounding anterior margin of ear. Three superciliaries on right, four on left; first longest (right), deeper anteriorly, longer than first supraocular, contacting first and second supraoculars, loreal, preocular, second superciliary and upper eyelid; second superciliary smallest, contacting second supraocular (left) or second and third supraoculars (right). Enlarged quadrangular scale follows third superciliary and contacts postocular. Central part of lower eyelid with semitransparent, undivided disc surrounded by small, slightly pigmented, granular, smooth scales and twelve strongly pigmented palpebrals. Temporals smooth, juxtaposed, irregular in size and shape, largest temporal closer to ear and about the same size of sixth supralabial. Ear opening surrounded by series of very small, juxtaposed, rounded tuberculate granules; external auditory meatus shallow; tympanum distinct, subovoid. Scales on sides of neck in about 13 irregularly transverse series between ear and arm level; those next to ear small, smooth, rhomboid, almost juxtaposed, becoming gradually larger, more elongate, mucronate, keeled and imbricate near arm. Neck scales close to nuchals sharply keeled. All head scales smooth and juxtaposed, with many scattered sensorial pits. Mental broad, wider than long. Postmental heptagonal, wider than long, contacting first and second infralabials. Three pairs of enlarged chinshields, in broad contact along midline; first smaller than second, contacting second and third infralabials on right side; second chinshield largest, contacting third and fourth infralabials on right side; third pair reduced, small, narrow, chevron-like, laterally contacting a scale that is slightly longer than wide; third pair of chinshields slightly smaller than fifth supralabial, separated from fifth infralabial by elongate scale. Six infralabials; first and second fused on left side; on the right, third and fourth infralabials largest, about the same size. Gulars enlarged, wider than long, smooth, imbricate, rounded posteriorly, in two longitudinal and seven transverse rows; scales in first row smaller. Interbrachial region distinct, with seven smooth, strongly imbricate scales; central one subtriangular, laterally contacting larger scales; external interbrachials smaller, longer than wide. Collar fold absent. Nuchals large, wider than long, multistriate, imbricate, rounded posteriorly, in two identical longitudinal and six regular transverse rows between parietal area and just before insertion of the arm. Occipitals absent. Dorsals becoming progressively narrower, imbricate, mucronate, lanceolate, with strong central keel and several lateral striae, variable in position, just anterior to insertion of arm, continuing in this fashion to hind limbs; keels appearing gradually, broad and flat anteriorly, becoming thin, high and sharp posteriorly. Twenty-eight transverse rows of dorsals between parietals/interparietal and posterior level of hind limbs; six anterior rows corresponding to striate, enlarged series. Scales on flanks strongly keeled, mucronate and imbricate, slightly smaller, and more diagonally disposed than dorsals; flank scales from the row bordering ventrals smooth, wider, more imbricate and posteriorly rounded. Axilla with distinctive area with small, smooth and rounded granules. Twenty-five scales around midbody. Ventrals smooth, imbricate, wider than long, rounded posteriorly, in two longitudinal and 16 transverse rows from interbrachials (not included) to preanals. Five preanal scales; central scale rhomboidal (posterior part wider), not reaching preanal border, the latter being formed by two enlarged scales in broad median contact and two smaller external paramedials. Preanal and femoral pores absent. Caudal scales elongate, lanceolate, keeled, strongly imbricate, smaller than midbody dorsals; smaller on regenerated part of tail.

416 · Zootaxa 4000 (4) © 2015 Magnolia Press COLLI ET AL. Forelimbs with large, smooth, imbricate scales, except on ventral part of forearm where they are much smaller, rounded, slightly imbricate, and on ventral part of upper arm where they are smaller than scales on corresponding dorsal parts. Anterior and ventral parts of hind limbs with irregularly large, smooth, imbricate scales, identical to those on corresponding parts of forelimbs, except for some on posterior dorsal part that are keeled. Scales on posterior part of hind limbs granular, juxtaposed, becoming larger, imbricate and keeled on dorsal part of tibia and femur. Carpal and tarsal scales large, imbricate, smooth; supradigital lamellae smooth, imbricate. Palmar and plantar surfaces with smooth, small, tuberculate granules. Fingers and toes relatively long and slender. Proximal and middle part of finger IV cylindrical, distal part laterally compressed. Proximal two-thirds of the 17 subdigital lamellae of finger IV single, flat to slightly bulbous, broad; distal scales single and smooth; divided lamella rarely present. Toe IV laterally compressed over its entire length; 23 subdigital lamellae single, narrow, with blunt ventral keel, or proximally a pointed tubercle on each lamella; middle part of finger with irregularly divided, paired scales, the distal scales being single. Toes and fingers clawed, except finger I; relative sizes: 1 < 2 = 5 < 3 = 4 and 1< 2 < 5 < 3 < 4, respectively. Dorsal surfaces of body, flanks and tail dark brown; dorsal part of head paler, olive-brown with scattered dark brown punctuate spots (Fig. 8). Lateral parts of head with a wide black stripe covering the entire lateral surface above the upper part of supralabials and extending from the nasal to arm level, merging there with flank colour. Below it, a bright orange-yellow bright colour covering most parts of supralabials, infralabials and ventral parts of head and neck. Ventral parts of body cream-yellow, strongly mottled, with irregular black pigmentation concentrated in anterior and central part of scales. Ventral parts of tail dark brown near its tip. Tail dark brown dorsally, slightly lighter ventrally. Limbs dark brown dorsally, irregularly mottled with cream-yellow; ventrally, cream-yellow, immaculate. Variation and sexual dimorphism. Variation in morphology is summarized in Table 1. The holotype (67 mm) is the largest specimen in the type series. The sexes can be readily separated by the absence of preanal and femoral pores in females. In addition, females have higher numbers of ventrals than males (Table 1, t = 3.025, df = 11.816, P = 0.011).

FIGURE 8. (A) View of a flooded forest near Igarapé-Açu, right bank of Rio Abacaxis, Nova Olinda do Norte, Amazonas Brazil; (B) terra firme forest at Igarapé-Açu; (C) holotype of Rondonops xanthomystax in life. (D) Closer view of the same specimen.

NEW LIZARDS FROM BRAZILIAN AMAZONIA Zootaxa 4000 (4) © 2015 Magnolia Press · 417 — — 16.14 ± 0.95 0.95 ± 16.14 (15–18) 6.47 ± 0.92 0.92 ± 6.47 (4–8) 3.00 ± 0.00 0.00 ± 3.00 (3–3) 22.21 ± 1.53 1.53 ± 22.21 (20–26) 0.35 ± 6.87 (6–7) 6.80 ± 0.77 0.77 ± 6.80 (5–8) 16.00 ± 1.11 1.11 ± 16.00 (13–17) 3.13 ± 0.35 0.35 ± 3.13 (3–4) 28.13 ± 1.13 1.13 ± 28.13 (26–30) 0.83 ± 6.40 (5–8) 5.20 ± 0.41 0.41 ± 5.20 (5–6) 122.5 ± 46.99 46.99 ± 122.5 (52–170) 26.07 ± 1.00 1.00 ± 26.07 (25–28) 54.13 ± 13.77 13.77 ± 54.13 (27–67) 22.2 ± 2.05 2.05 ± 22.2 (19–24) 15.40 ± 0.55 0.55 ± 15.40 (15–16) 6.80 ± 0.45 0.45 ± 6.80 (6–7) 3.00 ± 0 ± 3.00 (3–3) 21.40 ± 1.34 1.34 ± 21.40 (20–23) 6.80 ± 0.45 0.45 ± 6.80 (6–7) 7.00 ± 1.22 1.22 ± 7.00 (5–8) 15.40 ± 1.52 1.52 ± 15.40 (13–17) 7.00 ± 0.71 0.71 ± 7.00 (6–8) 27.40 ± 1.14 1.14 ± 27.40 (26–29) 3.20 ± 0.45 0.45 ± 3.20 (3–4) 5.00 ± 0.00 0.00 ± 5.00 (5–5) 114.33 ± 53.2 53.2 ± 114.33 (64–170) 26.00 ± 1.00 1.00 ± 26.00 (25–27) 52.8 ± 16.33 16.33 ± 52.8 (30–66) — — 3.00 ± 0.00 0.00 ± 3.00 (3–3) 16.56 ± 0.88 0.88 ± 16.56 (15–18) 6.30 ± 1.06 1.06 ± 6.30 (4–8) 6.70 ± 0.48 0.48 ± 6.70 (6–7) 16.33 ± 0.71 0.71 ± 16.33 (15–17) 22.67 ± 1.50 1.50 ± 22.67 (21–26) 6.90 ± 0.32 0.32 ± 6.90 (6–7) 6.10 ± 0.74 0.74 ± 6.10 (5–7) 3.10 ± 0.32 0.32 ± 3.10 (3–4) 28.50 ± 0.97 0.97 ± 28.50 (27–30) 5.30 ± 0.48 0.48 ± 5.30 (5–6) 54.8 ± 13.23 13.23 ± 54.8 (27–67) 127.40 ± 48.67 48.67 ± 127.40 (52–165) 26.11 ± 1.05 1.05 ± 26.11 (25–28) Females (10) Females (5) Males (15) Total . Values. indicate sample size (in parentheses, top row), mean ± standard R. xanthomystax 25.80 ± 1.19 1.19 ± 25.80 — — 17.02 ± 1.69 1.69 ± 17.02 (14–20) 6.72 ± 0.49 0.49 ± 6.72 (6–8) 0.50 ± 6.43 (6–7) 3.00 ± 0.00 0.00 ± 3.00 (3–3) 17.81 ± 0.92 0.92 ± 17.81 (16–20) 6.98 ± 0.34 0.34 ± 6.98 (6–8) 12.95 ± 1.02 1.02 ± 12.95 (11–15) 8.15 ± 0.97 0.97 ± 8.15 (6–10) 2.98 ± 0.13 0.13 ± 2.98 (2–3) 29.60 ± .39 .39 ± 29.60 (26–32) (23–30) 0.56 ± 4.83 (3–7) 110.14 ± 34.44 34.44 ± 110.14 (47–170) 52.55 ± 8.16 8.16 ± 52.55 (26–66) and Rondonops biscutatus Rondonops 26.14 ± 1.29 1.29 ± 26.14 (23–30) 19.81 ± 1.28 1.28 ± 19.81 (17–22) 3.00 ± 0.00 0.00 ± 3.00 (3–3) 16.28 ± 1.28 1.28 ± 16.28 (14–18) 6.72 ± 0.51 0.51 ± 6.72 (6–8) 0.51 6.5 ± (6–7) 7.00 ± 0.24 0.24 ± 7.00 (6–8) 12.92 ± 1.05 1.05 ± 12.92 (11–15) 17.61 ± 0.80 0.80 ± 17.61 (16–19) 8.06 ± 1.09 1.09 ± 8.06 (6–10) 2.97 ± 0.17 0.17 ± 2.97 (2–3) 29.14 ± 1.29 1.29 ± 29.14 (26–31) 4.86 ± 0.64 0.64 ± 4.86 (3–7) 53.94 ± 8.04 8.04 ± 53.94 (28–66) 125.6 ± 41.79 41.79 ± 125.6 (68–170) 61) 61) – (24–26) — (3–3) (15–20) (6–7) (6–7) (6–8) (11–15) (16–20) (7–9) (3–3) (27–32) (26 (4–5) Rondonops biscutatus (24) Females (36) Males (47–140) (60) Total Rondonops xanthomystax Summary of morphometric and meristic variables of of variables meristic and of morphometric Summary Scales around midbody midbody around Scales 0.81 ± 25.29 Pores — Pores Ventrals Ventrals 1.65 ± 18.12 Gulars 6.71 ± 0.46 0.46 0.48 ± ± 6.71 Gulars Infralabials 6.33 Supraoculars 3.00 ± 0.00 0.00 ± 3.00 Supraoculars Fourth toe lamellae lamellae toe Fourth 1.01 ± 18.13 Supralabials 6.96 ± 0.46 0.46 ± 6.96 Supralabials Fourth finger lamellae lamellae finger Fourth 1.00 ± 13.00 Superciliaries 3.00 ± 0.00 0.00 ± 3.00 Superciliaries Nuchals in double row row double in Nuchals 0.75 ± 8.29 Dorsals (incl. nuchals) nuchals) (incl. Dorsals 1.27 ± 30.29 Snout-vent length (mm) (mm) length Snout-vent 8.06 ± 50.46 Suboculars 4.79 ± 0.41 ± Suboculars 4.79 TABLE 1. only. tails to intact refer statistics length Tail parentheses). (in range and deviation Variables (mm) length Tail 28.67 ± 101.56

418 · Zootaxa 4000 (4) © 2015 Magnolia Press COLLI ET AL. Distribution, habitat and natural history. Rondonops xanthomystax is known from the forests of the middle Rio Abacaxis, Amazonas, and of the upper Rio Tapajós, in southwestern Pará, Brazil (Fig. 6). Seven specimens were obtained in two adjacent municipalities: Borba and Nova Olinda do Norte, on opposite sides of the Rio Abacaxis, Amazonas. All specimens were collected in primary terra firme forest, characterized by open understory, high abundance of palm trees, dense leaf litter and frequent large trees with more than 80 cm diameter and up to 40 m, like the Brazil nut Bertholletia excelsa (Fig. 8). One adult and two juveniles were collected by hand around 11:00 AM when foraging in sunny spots among the leaf litter. Three were taken from pitfall traps (Fig. 8) consisting of a set of four 30-liter buckets, arranged in a Y, separated by 4 m long drift fences, for a total effort of 175 traps x day. One specimen was collected on a small glue trap about 1.7 m high on the trunk of a diagonally- oriented fallen tree, approximately 50 cm in diameter. Six specimens from Rio Tapajós were collected in pitfall traps in terra firme forest, but MPEG 31107 and 31110 (both from Jacareacanga) were collected in riparian areas. Of the 10 females collected two were gravid, the holotype (MZUSP 98085) and MPEG 31106, each with one egg.

Phylogenetic analyses

The parsimony analysis of morphological data produced four equally parsimonious trees in which only about one- half of the nodes had bootstrap values >50 (not shown). In the Bayesian analysis of morphology, similar levels of support were found for the same nodes as in the parsimony tree. Thus we show only the Bayesian morphology tree (Fig. 9, right) with parsimony bootstrap values added to the figure. We collapse those nodes in which the Bayesian posterior probability (BPP) is < 0.50. Hereafter, we refer to this as the morphology tree. This tree weakly supported the sister-group relationship of Iphisa and Rondonops (PBS = 69; and BPP = 0.79; Fig. 9, right).

FIGURE 9. Bayesian phylogenies of the combined DNA and morphology dataset (left) and the morphological dataset only (right). The parsimony bootstrap values are shown above the branches or horizontal line; these values apply to this particular tree and do not indicate that the parsimony analysis supported these nodes most strongly. The Bayesian posterior probabilities are shown below the branches or horizontal line. In the tree on the right, the polytomies were formed by collapsing those nodes with <50% posterior probability. The dots refer to the names of the higher taxa.

NEW LIZARDS FROM BRAZILIAN AMAZONIA Zootaxa 4000 (4) © 2015 Magnolia Press · 419 FIGURE 10. Maximum likelihood topology for the DNA-only dataset. The parsimony bootstrap values are shown above the branches or horizontal line; these values apply to this particular tree and do not indicate that the parsimony analysis supported these nodes most strongly. The Bayesian posterior probabilities are shown below branches. Asterisks represent bootstrap values of 100% or Bayesian posterior probabilities of 1. The dots refer to the names of the higher taxa.

420 · Zootaxa 4000 (4) © 2015 Magnolia Press COLLI ET AL. The DNA-only tree presented in Fig. 10 is the optimum RAxML topology with the parsimony bootstrap support values above the branch and the Bayesian posterior probabilities below the branch. Our analyses recovered several strongly supported (BPP >95 and PP = 1) clades; we rank these as subfamilies: Alopoglossinae, Ecpleopodinae, Bachiinae (subfam. nov.), Rhachisaurinae, Gymnophthalminae, and Cercosaurinae. Alopoglossinae is the most basal clade, followed by Ecpleopodinae. Cercosaurinae is the sister-group of a clade containing Bachiinae + (Rhachisaurinae, Gymnophthalminae), although this relationship is poorly supported. We also recovered as monophyletic three groups, ranked as tribes, within Gymnophthalminae, all with PP = 1: Chirocolini, Iphisini, and Gymnophthalmini. The clade of Rondonops and Iphisa is strongly supported (BPP = 82 and PP = 1). Elsewhere within Gymnophthalminae some nodes are weakly supported; seven have posterior probabilities <0.90. The combined DNA and morphology analysis (Fig. 9, left) yielded a tree that is generally similar to the morphology tree, also supporting a sister-group relationship between Iphisa and Rondonops (PBS = 54 and BPP = 1). In summary, there are no strongly supported conflicts in topology between the two trees. If the complete DNA tree (Fig. 10) is pruned to retain only those taxa in common with those in Fig. 9, the two trees are almost identical, the only minor differences being in two weakly supported nodes (not shown). Overall, phylogenetic analyses indicate that Rondonops is clearly a member of Iphisini and the closest relative of Iphisa.

Revised Classification of Gymnophthalmidae

Gymnophthalmidae Merrem, 1820

Alopoglossinae Pellegrino, Rodrigues, Yonenaga-Yassuda, and Sites Jr., 2001. Content: Alopoglossus Boulenger, 1885; Ptychoglossus Boulenger, 1890.

Ecpleopodinae Fitzinger, 1843. Content: Adercosaurus Myers and Donnelly, 2001; Amapasaurus Cunha, 1970; Anotosaura Amaral, 1933; Boulenger, 1885; Cunha and Lima Verde, 1991; Rodrigues, Freire, Pellegrino, Sites Jr. 2005; Ecpleopus Duméril and Bibron, 1839; Kaieteurosaurus Kok, 2005; Leposoma Spix, 1825; Marinussaurus Peloso, Pellegrino, Rodrigues and Ávila- Pires, 2011; Pantepuisaurus Kok, 2009.

Bachiinae, subfam. nov. Diagnosis: Bauplan associated with fossorial habits. Body serpentiform, elongate, cylindrical. Four reduced limbs, especially hind limbs, with variable number of digits, some lacking claws. External ear absent; lower eyelid with unsegmented semi-transparent disc. Head scales reduced in number; frontoparietals absent. Ventrals smooth. Distribution: , from Costa Rica into South America east of the Andes, south to Paraguay, and on some islands. Content: Bachia Gray, 1845.

Rhachisaurinae Pellegrino, Rodrigues, Yonenaga-Yassuda, and Sites Jr., 2001. Content: Rhachisaurus Pellegrino, Rodrigues, Yonenaga-Yassuda, and Sites Jr., 2001.

Gymnophthalminae Merrem, 1820 Chirocolini Gray, 1838. Content: Caparaonia Rodrigues, Cassimiro, Pavan, Curcio, Verdade and Pellegrino, 2009; Colobodactylus Amaral, 1933; Heterodactylus Spix, 1825.

Iphisini Gray 1851. Content: Acratosaura Rodrigues, Pellegrino, Dixo, Verdade, Pavan, Argolo and Sites Jr., 2007; Alexandresaurus Rodrigues, Pellegrino, Dixo, Verdade, Pavan, Argolo and Sites Jr., 2007; Colobosaura Boulenger, 1887; Iphisa Gray, 1851; Rondonops gen. nov.; Stenolepis Boulenger, 1888.

NEW LIZARDS FROM BRAZILIAN AMAZONIA Zootaxa 4000 (4) © 2015 Magnolia Press · 421 Gymnophthalmini Merrem, 1820. Content: Rodrigues, 1991; Merrem, 1820; Micrablepharus Dunn, 1932; Rodrigues, 1984; Procellosaurinus Rodrigues, 1991; Psilophthalmus Rodrigues, 1991; Scriptosaura Rodrigues and Santos, 2008; Tretioscincus Cope, 1862; Rodrigues, 1991.

Cercosaurinae Gray, 1838. Content: Gray, 1845; Wagler, 1830; Boulenger, 1890; Tschudi, 1845; Noble, 1921; Duméril and Bibron, 1839; Doan and Castoe, 2005; Pholidobolus Peters, 1862; Tschudi, 1847; Doan and Castoe, 2005; Tschudi, 1845; Gray, 1858; Uzzell, 1973; Teuchocercus Fritts and Smith, 1969.

Discussion

We conducted maximum parsimony and Bayesian analyses of morphological and molecular datasets to determine the phylogenetic placement of Rondonops among the major clades of Gymnophthalmidae. Overall, our results are very similar to those of Pyron et al. (2013) regarding the relationships among the major groups of Gymnophthalmidae, except for the placement of Cercosaurinae, which those authors recovered as the sister-group of Ecpleopodinae and which we recovered as the sister-group of Bachiinae + (Rhachisaurinae + Gymnophthalminae). In neither arrangement are the relationships strongly supported. Our results consistently support a sister-group relationship between Rondonops and Iphisa, both being members of the Iphisini radiation of Gymnophthalminae. Members of this clade were previously considered as part of Heterodactylini (sensu Pellegrino et al. 2001), but Rodrigues et al. (2009b) found Heterodactylini to be paraphyletic on the basis of Bayesian analyses of combined molecular and morphological data. Therefore, they restricted Heterodactylini (presently Chirocolini) to Caparaonia, Colobodactylus, and Heterodactylus, and proposed a new tribe, Iphisiini (=Iphisini of this paper), to accommodate Acratosaura, Alexandresaurus, Colobosaura, Iphisa, and Stenolepis (Rodrigues et al. 2009b). Our study further corroborates this arrangement. Rodrigues et al. (2007b) suggested that Atlantic Forest members of Iphisini occupy a more basal position and have more restricted ranges than those from Amazon Forest or Cerrado. Occupying a more crown position in Iphisini and having a broad geographic distribution in Amazonia, Rondonops corroborates this hypothesis. Nevertheless, this pattern may be artificial and the wide-ranging species from Amazonia and Cerrado may in fact consist of a number of cryptic species with much smaller ranges, as observed in other wide-ranging lizard species (e.g., D'Angiolella et al. 2011; Gamble et al. 2012; Gamble et al. 2011; Werneck et al. 2012). Indeed, recent analyses of hemipenial morphology and nucleotide sequences revealed that the widespread Amazonian Iphisa elegans is a complex of cryptic species (Nunes et al. 2012). A phylogeographic analysis of , widespread in Cerrado and also occurring in parts of eastern Amazonia, may reveal a similar pattern. Both new species were collected mainly in pitfall traps in the forest or near the forest edge, but R. biscutatus also occurs in open vegetation enclaves. A similar pattern is found in Iphisa (Vitt et al. 2008; SMS, personal observation). As far as we know, R. xanthomystax is restricted to forest . They also differ in habitat use: Rondonops biscutatus is predominantly a leaf litter species, like most gymnophthalmids, whereas R. xanthomystax apparently uses the forest structure more broadly. Rondonops xanthomystax and R. biscutatus are known from several localities in Amazonia, from Rondônia to Pará. In the Rio Tapajós area of southern Pará (Itaituba and Jacareacanga) the species are sympatric, but have not yet been found syntopically. No clear differences were found between habitats where they occurred. Most localities are in the "arc of deforestation", a crescent-shaped area where deforestation is concentrated, including southwestern Maranhão, northern Tocantins, southern Pará, northern Mato Grosso, Rondônia, southern Amazonas and southeastern Acre (Brooks et al. 2002; Fearnside & Graça 2006; Ferreira et al. 2005). An analysis based on high-spatial-resolution Landsat imagery from 2000 and 2005 indicated that the "arc of deforestation" accounts for ca. 50% of clearing in humid tropical forests globally, nearly four times the level of Indonesia, which has the next highest rate (13%) (Hansen et al. 2008). In addition to habitat loss and fragmentation, deforestation in the "arc of deforestation" is also promoting changes in cloud cover that may lead to increased seasonality (Costa & Pires 2010; Durieux et al. 2003) and increased frequency of fires (Morton et al. 2013; Righi et al. 2009). The combined effects of these changes are promoting severe biodiversity erosion (Coe et al. 2013; Korfanta et al. 2012; Lees & Peres 2006; Mahood et al. 2012).

422 · Zootaxa 4000 (4) © 2015 Magnolia Press COLLI ET AL. Human occupation in Brazilian Amazonia steadily moves northward (e.g., along the BR-163 and BR-319 highways), radiating from the "arc of deforestation," and the resulting loss of pristine habitat may cause loss of biodiversity, especially in small cryptozoic species such as most herpetofauna. It is therefore important to stem this mostly illegal activity as soon as possible, but apparently in Brazil this is a moot point, as impunity for illegal deforestation is the rule rather than the exception. After some years of decreasing deforestation rates in Brazilian Amazonia (especially in the state of Pará), the most recent data indicate an increase in deforestation between 2012 and 2013 of 28.9%. One of the reasons of this increase is the fact that, because of technical limits, the satellite monitor system cannot detect invaded areas of less than 25 ha, and small farmers used this knowledge to deforest small plots (Beer 2014). Data from 2013–2014 might even be more alarming, but apparently have been kept back because of Brazilian presidential elections. The increasing demand for mineral resources and hydropower is another factor that severely threatens the Amazonian biodiversity (Ferreira et al. 2014). Given the rate and extent of habitat loss in this region, it is likely that many species will go extinct before being discovered.

Acknowledgements

We thank Coordenação de Aperfeiçoamento de Pessoal de Nível Superior—CAPES, Conselho Nacional de Desenvolvimento Científico e Tecnológico—CNPq, Fundação de Amparo à Pesquisa do Estado de São Paulo— FAPESP, Fundação de Amparo à Ciência e Tecnologia do Estado de Pernambuco—FACEPE and Fundação de Apoio à Pesquisa do Distrito Federal—FAPDF for financial support. DCC and GRC also thank the Fulbright Scholar Program for financial support.

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