Zootaxa 3616 (2): 173–189 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.3616.2.6 http://zoobank.org/urn:lsid:zoobank.org:pub:D6F85F14-2D4E-4B53-9B40-3B6E9BE723FF A new species of Gray, 1845 (: ) from the Eastern Brazilian Cerrado, and data on its ecology, physiology and behavior

MAURO TEIXEIRA JR1,3, RENATO SOUSA RECODER1, AGUSTÍN CAMACHO1, MARCO AURÉLIO DE SENA1, CARLOS ARTURO NAVAS2 & MIGUEL TREFAUT RODRIGUES1 1Laboratório de Herpetologia, Departamento de Zoologia, Instituto de Biociências, Universidade de São Paulo, CEP 05508-090, São Paulo, SP, Brazil 2Departamento de Fisiologia, Instituto de Biociências, Universidade de São Paulo, CEP 05508-090, São Paulo, SP, Brazil 3Corresponding author. E-mail: [email protected]

Abstract

A new species of Bachia of the bresslaui group, Bachia geralista sp. nov., is described from Planalto dos Gerais, an old and partially dissected plateau extending along the Cerrados of Bahia, Minas Gerais and Tocantins states, Brazil. The new species is morphologically similar to B. bresslaui, with which it has been confused; however head scalation resembles other species from sandy spots within the Cerrado (B. psamophila and B. oxyrhina). Like in B. psamophila and B. oxyrhi- na, the shovel-shaped snout of the new species is highly prominent, a typical trait of psammophilous habits in other gym- nophthalmids. The examination of specimens of B. bresslaui from several populations within the Cerrado revealed great variation among localities, leading to the reidentification of a specimen from Utiariti, Mato Grosso, previously referred to in the literature as the second record of B. bresslaui, as the recently described B. didactyla, suggesting that cryptic diversity might remain still undiscovered within this genus in the Cerrado. Despite occurring in a relatively open Cerrado, thermal physiology of Bachia geralista sp. nov. restricts its occurrence to shaded microhabitats within this habitat.

Key words: Bachia geralista sp. nov., psammophilous habits, Peruaçu, limb reduction, fossoriality.

Introduction

At the end of the XVIII century Bonnaterre (1789) described Chalcides flavescens, a small worm-like from South America, with vestigial limbs. Few decades later Duméril and Bibron (1839) described C. dorbignyi, a second South American species from Bolivia. Later, Gray (1845) recognized Duméril and Bibron’ species as belonging to a distinct genus, and described Bachia to accommodate it. More than 150 years has passed and the genus Bachia, including Bonnaterre’ species, currently comprises 22 species distributed mostly over the tropical South America, and southern Central America (Dixon 1973; Castrillon & Strussmann 1998; Kizirian & McDiarmid 1998; Rodrigues et al. 2007; Rodrigues et al. 2008; Freitas et al. 2011). Forest dwelling species of Bachia frequently show a rounded snout, and fingers and toes relatively developed (Dixon 1973). However, in the last few decades, several species have been described from the open areas within the Cerrado of Central Brazil (Castrillon & Strussmann 1998; Rodrigues et al. 2007; Rodrigues et al. 2008; Freitas et al. 2011), all showing reduced limbs, and those from sandy habitats (e.g. B. micromela Rodrigues, Pavan & Curcio 2007, B. psamophila Rodrigues, Pavan & Curcio 2007 and B. oxyrhina Rodrigues, Camacho, Nunes, Recoder, Teixeira, Valdujo, Ghellere, Mott & Nogueira 2008) also showing reduction on head scalation and a shovel-like snout (Rodrigues et al. 2007, Rodrigues et al. 2008). The relationship between morphology, ecology, physiology and behavior of species are basic to recognize potential limitations to their ecological or geographical distribution, and support further conservation actions or evolutionary studies. However, as for most gymnophthalmids, these traits are very poorly known in Bachia (Colli 1998; Wiens et al. 2006; Henderson & Powell 2009; Bentz et al. 2011).

Accepted by S. Carranza: 4 Jan. 2013; published: 19 Feb. 2013 173 Based on a series of specimens obtained at the southernmost end of Planalto dos Gerais, in the eastern part of Brazilian Cerrado, as well as on exemplars from nearby localities, we recognize those populations, which have been previously included in the widely distributed B. bresslaui (Amaral, 1935) (Rodrigues et al. 2008), as a distinct new species which we describe, providing data on its ecology, physiology and behavior, and discussing the implications of the morphological variation found within B. bresslaui.

Material and methods

Field sampling. Specimen collection was made through pitfall traps with drift fences and active search. The pitfall traps were installed throughout the main habitats at Parque Nacional das Cavernas do Peruaçu: 15 traps on dry forest habitats; 15 on savanna; 17 on an arborescent scrubland, locally known as “carrascos”, and two on riparian evergreen forest. Each trap was composed of four 30 L buckets connected by 4 m long vertical plastic fences, which remained opened for 20 consecutive days each sampling period: January 2008 and 2009 (rainy seasons), and July 2008 and 2009 (dry seasons); totalizing an effort of 14.560 buckets/day. Active searches were performed during July 2008 and January 2009, and only at the savanna habitat. Searches consisted in racking all the leaf litter, trunks or shallow soil of haphazardly selected microhabitat patches, divided in three categories: (i) open areas, (ii) bushes and (iii) trees. At each patch, the individuals found were collected, and the size of the patch estimated by measuring its two major perpendicular axis and using the ellipse’s formulae to calculate the area. Concomitantly to that, environmental temperatures were measured hourly by HOBO data loggers (resolution 1ºC, with white probe weighting 1g), laid at two different microhabitats, (i) ground’s surface or the leaf litter and (ii) the deepest limit of the loose soil (usually 2cm), in both open areas and under vegetation, totalizing four microhabitats. In the latter case, the logger probe was covered by leaf litter plus loose soil layer. To show potential thermal restrictions imposed by the different microhabitats over Bachia’s ecological distribution at this site, maximum and minimum temperatures were compared among microhabitats. When not designated to physiological experimentation, or after that, the individuals were sacrificed, fixed in 10% formalin, preserved in 70% ethanol, and housed at Museu de Zoologia da Universidade de São Paulo, São Paulo, Brazil. Behavioral and physiological observations. Escape and defensive behavior of some individuals was observed and annotated during data collection and within set ups for focal trials (10 individuals). These set ups consisted of a recipient filled with sandy soil 4 cm’s deep, having half of its area covered by leaf litter. In each trial, a single specimen was placed in one of both substrates (sand or sand covered by leaf litter), and its first reaction was categorized as: (i) slide, (ii) hide (meaning partial burrowing or hiding under leaf litter), or (iii) burrow. All ten specimens were subjected to trials in both substrates, in random order. Ten individuals, maintained for a year, where observed from 5:00 to 23:00. were kept in plastic terraria, keeping together the groups found in single patches during fieldwork. All terraria received UV (L12:D12) and heating light (L10:D14), with diurnal thermal gradients ranging between 20º–45ºC, falling to 22–20ºC at nights. All were fed with termites, and immature cockroaches and crickets. Water was provided every two days spraying it directly on leaf litter and terrarium walls. Thermal preferences were measured over the day and beginning of the night (8–19:30h, lights were switched off at 18:30) in thermal gradients ranging between 14–55ºC. A T-type thermocouple was fixed to the back of lizards using surgical tape, allowing body temperatures to be registered in a computer using a multiple thermocouple register (model 8018, Superlogics®). The critical thermal maximum was assessed through incidental (Cowles & Bogert 1944) miss function of the thermal chamber, set to 39ºC, that suddenly raised to 41ºC and cause the death of individuals. Morphology. Length measurements were taken after fixation to the nearest millimeter with a digital caliper; scale counts and observations of other morphological characters were performed with a stereomicroscope Zeiss STEMI SV6. Scale nomenclature follows Dixon (1973). Comparisons were made primarily through examined specimens housed in the herpetological collection of the Museu de Zoologia, Universidade de São Paulo (MZUSP) (Appendix I), and were supplemented with data from the literature (Vanzolini 1961; 1966; Dixon 1973; MacLean 1973; Avila-Pires 1995; Castrillon & Strussmann 1998; Colli et al. 1998; Rodrigues et al. 2007; Rodrigues et al. 2008; Freitas et al. 2011).

174 · Zootaxa 3616 (2) © 2013 Magnolia Press TEIXEIRA JR ET AL. Meristic characters were: (SO) supraoculars; (SC) superciliaries; (SL) supralabials; (TS) temporal, scales along each three oblique series of temporals; (FP) femoral pores; (PP) preanal pores; (PS) preanal shields; (SAB) scales around midbody; (DO) dorsals, between parietals to the row over the insertion of hindlimbs; (VE) ventrals, between interbrachial shields and preanal shields; (GU) gulars, between interbrachial and mental plates; (CA) caudals. The measurements were: (SVL) snout-vent length, from border of cloaca to the tip of snout; (IBNL) interbrachial-nasal length, from the anterior border of the median interbrachial scale to the tip of snout; (LBM) length between members, from the anterior margin of hind limb to the posterior margin of forelimb; (HH) head high at highest point in the longitudinal axe; (HW) head width at widest point; (HL) head length from the anterior margin of tympanic aperture to the tip of snout; (FLL) fore-limb length; (HLL) hind-limb length; (END) eye- nostril distance; (NND) nostril-nostril distance; (TL) tail length, measured only in intact tails. Distribution. Distributional records of species from B. bresslaui group were gathered from the literature (Vanzolini 1961; 1966; Dixon 1973; MacLean 1973; McDiarmid & Foster 1987; Avila-Pires 1995; Castrillon & Strussmann 1998; Colli et al. 1998; Ferreira et al. 2005; Rodrigues et al. 2007; Rodrigues et al. 2008; Freitas et al. 2011), from online databases (GBIF 2012; SpeciesLink 2012), and from examined specimens (Appendix I).

Results

Taxon description

Bachia geralista sp. nov. (Figs. 1–3)

Bachia bresslaui Rodrigues et al. (2008:45)

Holotype: MZUSP 99408, an adult from Parque Nacional das Cavernas do Peruaçu, (15°09'14.04"S, 44°18'4.32" W, 740 m a.s.l., WGS84), Januária municipality, state of Minas Gerais, Brazil, collected by the authors on 25th January, 2008. Field number MTJ 212. Paratopotypes: MZUSP 99406, 99407, 99409, 103211–10321, all collected by the authors, in January and July 2008, and January and July 2009. Paratypes: MZUSP 100021, an adult from Roda Velha (12°47'1.86"S, 45°56'12.77"W, 750 m a.s.l.), São Desidério municipality, state of Bahia, Brazil, collected by M.A. Freitas on 5th January, 2010. MZUSP 94473–94474, Parque Nacional Grande Sertão Veredas (15°13'0.12"S, 45°47'60.00"W, 670 m a.s.l.), Formosa municipality, Minas Gerais, Brazil, collected by C. Nogueira on 23–29th October, 2001. Etymology: The specific name is a noun in apposition; “geralista” is an old Brazilian term applied to the inhabitants of the “Planalto dos Gerais” from which the new species seems to be endemic. Diagnosis: (1) A large-sized species of Bachia (101 mm of maximum known SVL); (2) prefrontals absent; (3) six supralabials; (3) 2-2 supraoculars and 2-2 supraciliars; (4) 2-2 femoral and 1-1 preanal pores; (5) 33–35 scales around midbody; (6) 46–51 dorsal rows of scales; (7) 37–40 ventral rows of scales; (8) 8–9 gulars; (9) 5 large preanal scales, and two small, on each side of the preanal plate; (10) sixth supralabial in contact with parietal, in most of the individuals; (11) ventral scales quadrangular, smooth; (12) fore and hindlimbs ending in an single apical scale; (13) first temporal absent. Description of the holotype: Body elongate, with a slight cervical constriction on head, snout highly prominent and wedge shaped, tail longer than body. Rostral broad, prominent, contacting first supralabial, nasal and frontonasal. Viewed from above the rostral is about twice as wide as high; on lateral view it projects broadly anteriorly toward, forming a horizontal surface ventrally, at the level of ventral surface of the upper lips. Frontonasal trapezoidal, as wide as long, wider posteriorly, contacting rostral, nasal, first supraocular and frontal. Prefrontals absent. Frontal pentagonal, longer than wide, with anterior margin slightly convex, narrow than, and in broad contact, with frontonasal; lateral margins straight to slightly concave, in contact with first supraocular; posteriorly angulose, broadly contacting parietals and in short contact with interparietal. Frontal about three times wider than anterior supraocular. Frontoparietals absent. Interparietal narrow, longer than wide, subrectangular, slightly wider posteriorly, shorter than parietals, as long as frontal. Parietals very large, longer than wide, longer

NEW SPECIES OF BACHIA Zootaxa 3616 (2) © 2013 Magnolia Press · 175 and wider than frontal, roughly hexagonal; their anterior margin deeply indented and in broad contact with frontal, contacting narrowly the first and broadly the second supraoculars, the postocular, punctually the sixth supralabial, two temporal and the dorsals; internally it contacts frontal and interparietal. Posterior borders of interparietal and parietals and dorsals coincides with a very slight transverse cervical constriction in the occipital region. Two supraoculars, first largest, about three times longer than wide, contacting frontal, frontonasal, nasal, loreal, first superciliary, second supraocular and parietal. Second supraocular smaller, above second superciliary, contacting postocular, longer than wide, separated from frontal by the slight contact between parietal and first supraocular. Two superciliaries, the first longer, their suture coincides with that between supraoculars. Nasal large, longer than high. Nostril in the middle of lower margin of nasal. Loreal roughly squared, in contact with nasal, first supraocular, first superciliary, preocular and second and third supralabials. Preocular small, curved. Frenocular present in contact with subocular. Six supralabials, third, fourth and fifth under the orbital region, sixth the highest and largest, contacting punctually parietal. One long subocular. Eyelid present with an undivided semitransparent disc. A large and elongate postocular between fifth and sixth supralabials and parietal. First temporal absent. Second temporal enlarged, longer than wide, temporal scale between parietal and sixth labial, in broad contact with both. Ear opening absent. All head scales smooth and juxtaposed. Mental roughly trapezoidal, wider than long, wider than the ventral surface of rostral. Postmental heptagonal, as wide as long. Two pairs of chin shields, both contacting infralabials; the anterior pair smaller, in broad contact at midline; second pair in narrower contact at the midline; a pair of symmetric flat and diagonally disposed elongate pregulars. Five infralabials, first the smallest, second, third and fourth with about the same size. Gulars smooth, imbricate, rounded posteriorly, in nine transversal rows; scales of gular rows increasing gradually in size toward interbrachial region. Interbrachial region with four scales, the central ones largest, twice longer than wide. 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, strongly keeled, with lateral sides almost juxtaposed. Fifty transverse rows between interparietal and the level of hind limbs. Lateral scales about the same size as dorsals but smooth and less acuminate; those closer to ventrals slightly wider. A distinctive area with granular scales surrounds the area of arm insertion and the posterior part of leg insertion. Thirty two scales around 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; 38 transverse rows between interbrachials and preanals. Seven scales (five larger, central scales, and one small placed in each side of the preanal plate) in the posterior part of preanal plate, central one largest. One preanal and two femoral pores on each side. Scales of tail similar to midbody dorsals, keeled, lanceolate, strongly imbricate. Fore limbs rudimentary, stiliform, covered by smooth and imbricate scales, ending in a single apical scale. Its length corresponds to one row of lateral scales width. Hind limb also stiliform but larger than fore limb, covered by smooth, large and imbricate scales ending by one apical scale; its length equivalent to one and half scale rows. Two femoral pores present at each side, proximal one the largest. Background dorsal and lateral surfaces of body and tail cream to light brown, with a dorsolateral light stripe. Ventral parts of body and tail cream immaculate. Measurements of the holotype (mm): SVL=87.86; IBNL=15.91; LBM=67.90; HH=4.08; HW=5.31; HL=8.90, FLL=2.44; HLL=3.25; END=3.96, NND=2.60; TL=140 (regenerated) Variation: Most individuals show the same color pattern as described for the holotype, however larger individuals tends to have a more homogeneous coloration, with faded stripes (Fig. 2A,B). Although none individual possess a first temporal (Figs. 1,3), in some specimens none supralabial is in contact with parietal, because the postocular is in contact with the second temporal row. In some individuals in one side of the head there is contact between supralabial and parietal and in the other side there is not. There is also variation in the number of scales on the second temporal row: in some individuals the two scales on the second temporal row are merged. Meristics and morphometric values of B. geralista sp. nov. and B. bresslaui are given on Table 1.

176 · Zootaxa 3616 (2) © 2013 Magnolia Press TEIXEIRA JR ET AL. FIGURE 1. Lateral (A), dorsal (B) and ventral (C) views of the head of the holotype of Bachia geralista sp. nov. (MZUSP 99408). Bar represents 5 mm.

NEW SPECIES OF BACHIA Zootaxa 3616 (2) © 2013 Magnolia Press · 177 FIGURE 2. Living paratopotypes of Bachia geralista sp. nov. from Parque Nacional Cavernas do Peruaçu, Minas Gerais state, Brazil, showing color pattern with a distinct dorsolateral yellowish stripe (A) and with a dorsolateral stripe faded (B).

178 · Zootaxa 3616 (2) © 2013 Magnolia Press TEIXEIRA JR ET AL. TABLE 1. Meristic and morphometric (mm) values of Bachia geralista sp. nov. and B. bresslaui. Abbreviations are explained in the text. Meristics Morphometrics B. geralista sp. nov B. bresslaui B. geralista sp. nov B. bresslaui SAB 30–33 33–35 SVL 71.9–101.3 64.8–104.2 DO 46–51 47–53 IBNL 14.6–18 14.2–19.2 VE 36–40 36–41 TL 141–162 122.5–148.5* GU 8–9 9–10 LBM 54.6–79.5 48.1–80.8 SL 6 6–7 HH 3.2–4.6 3.5–4.5 CA 108–115 117–123 HW 4.5–5.8 4.6–5.8 PS 7 5–7 HL 7.6–10.1 7.6–10.5 SO 2 2 FLL 1.8–2.7 2–2.8 SC 2 2 HLL 2–3.9 1.9–3.5 FP 2 1–2 END 3.2–4.2 3–4.2 PP 1 1 NND 2.1–2.8 2.1–2.9

*Only smaller adult individuals had their tails intact, thus this range may not be complete.

FIGURE 3. Paratypes of Bachia geralista sp. nov. (A) from Parque Nacional Grande Sertão Veredas, MG (MZUSP 99473) and (B) from São Desidério, BA (MZUSP 100021).

NEW SPECIES OF BACHIA Zootaxa 3616 (2) © 2013 Magnolia Press · 179 Comparison with other species (data from species in comparison are given in parentheses): the new species is a member of the B. bresslaui group by having lanceolated, keeled, imbricate dorsal scales, quadrangular, juxtaposed ventrals, 2-2 femoral pores, 1-1 preanal pores, limbs ending in one apical scale. These characters promptly distinguish B. geralista sp. nov. from all species of the B. dorbignyi group (B. dorbignyi (Dumeril & Bibron, 1839) B. barbouri Burt & Burt, 1931, B. bicolor (Cope, 1896), B. huallagana Dixon, 1973, B. intermedia Noble, 1921, B. peruana Werner, 1901, B. talpa Rutven, 1925, and B. trisanale (Cope, 1868)), from the B. heteropa group (B. heteropa (Lichtenstein, 1956) and B. pallidiceps (Cope, 1862)) (both having hexagonal, smooth dorsal scales), and from B. heteropa and B. flavescens (B. flavescens (Bonnaterre, 1789) and B. monodactyla (Daudin, 1802)) groups (hindlimbs with more than one digit). Among the species of B. bresslaui group (diagnostic features for species of B. bresslaui group are summarized in Table 2), B. geralista sp. nov. can be promptly distinguished from all species by having 30–33 scales around midbody (29–30 in B. oxyrhina and 33–47 in the combination of the remaining species); from all species, with exception of B. oxyrhina and B. bresslaui, it differs in having both fore and hindlimbs ending in a apical single scale (forelimbs ending in four apical scales in B. cacerensis Castrillon & Strüssmann, 1998, hindlimbs with two apical scales in B. micromela, hindlimbs with four clawed toes in B. psamophila, both limbs with four reduced toes and fingers in B. panoplia Thomas, 1965, and B. scolecoides Vanzolini, 1961, both limbs with free fingers and toes in B. pyburni Kizirian & McDiarmid, 1998, and forelimbs with two apical scales in B. didactyla Freitas, Strüssmann, Carvalho, Kawashita-Ribeiro & Mott, 2011); from all other species except B. oxyrhina and B. psamophila by the acute, shovel-shaped snout in profile, (snout rhomboid, rounded in profile in B. scolecoides, B. didactyla, B. pyburni, B. cacerensis, B. micromela and rhomboid, slightly protruding upwards in B. bresslaui and B. panoplia); from all species except B. cacerensis, B. oxyrhina and B. psammophila by the absence of the first temporal, allowing a narrower contact between supralabial and parietal (first temporal present, preventing contact between supralabial and parietal); from all species except B. micromela by a contact between parietal and supralabial (parietal and supralabial without contact), although the contact is with the 6th supralabial in B. geralista sp. nov. and with the 5th supralabial in B. micromela; from B. panoplia, B. pyburni and B. scolecoides by the absence of prefrontals (present and in contact in P. panoplia and B. pyburni, and widely separated and reduced in size in B. scolecoides); from B. oxyrhina and B. pyburni by having six supralabials (five and seven, respectively); from B. oxyrhina and B. didactlyla by having the first supralabial and nasal separated (merged); from B. pyburni, B. panoplia and B. micromela by having all ventral scales smooth, quadrangular (lanceolated, keeled in B. pyburni and B. panoplia, and anteriorly quadrangular, smooth, and posteriorly, close to vent, lanceolated, keeled in B. micromela). Distribution: Bachia geralista sp. nov. is currently known to occur only in the Planalto dos Gerais, at the eastern Brazilian Cerrado, a large eroded sandstone plateau that covers all western Bahia state, and some parts of Minas Gerais and Tocantins; emerging from the surrounding lowlands, it reaches more than 900 m a.s.l. in some points (Fig. 4). Although we are aware of only a few localities where B. geralista sp. nov occurs, literature records indicates the presence of Bachia at other points of Planalto dos Gerais: Correntina (Colli 1998) and Cocos (Rodrigues et al. 2008), from which specimens not examined. Further examination are need to attest if they belong to this new species. Natural history: Habitat—Although we sampled all available environments in the area (e.g. dry forest, riparian forest, carrasco and savannas), pitfall trapping at the Peruaçu region, found only three specimens of B. geralista sp. nov., which were found only at the savanna. Active searches within this habitat showed that B. geralista sp. nov. inhabits primarily the leaf litter under larger trees (9 out of 11, found using standardized active search, of which 6 were found under ‘Pequi’ trees (Caryocar brasiliense) (Fig. 5). Only one specimen was found under a dead trunk and two under the leaf litter of bushes. Mean abundance was 5 individuals/ha, among all microhabitats, and 6.9 individuals/ha, considering only where lizards were found. Typically, we found single specimens under each patch. Only in two occasions we found more. In one of these situations, we found two large adults and a smaller specimen under an unidentified tree, in the other, four hatchlings within a square meter under a Pequi tree. Thermal ecology—Temperatures above 40ºC (lethal for B. geralista sp. nov.) were attained at all microhabitats (Fig. 6A), in both seasons. Lethal temperatures occurred from 7:00h–16:00h, both in the surface and the exposed subsoil and, only between 13:00–15:00h, under the loose soil covered by vegetation. Temperatures below the fifth percentile (11.38º C) of the frequency distribution of preferred temperatures occurred at all the

180 · Zootaxa 3616 (2) © 2013 Magnolia Press TEIXEIRA JR ET AL. microhabitats measured. Absolute minimum (7.43ºC) was reached in July (dry season), at 6:00h in the surface of the leaf litter, at the same time, the temperature registered under the loose soil layer was between 14–16ºC. Preferred temperatures of B. geralista sp. nov. were 31.22ºC (50% central range 33.84–28.18ºC) (Fig. 6A).

TABLE 2. Summary of diagnostic features among Bachia species of the B. bresslaui group. B. geralista sp. nov. B. bresslaui B. cacerensis B. didactyla B. psamophila Prefrontals absent absent absent absent absent Supralabials six six* six six six First supralabial separated separated separated merged separated and nasal Scales around 30–33 33–35 35 34–37 35–38 midbody Contact 6th supralabial no contact** 6th supralabial no contact 6th supralabial supralabial/parietal First temporal absent present** absent present absent Contact loreal/ no no no yes no internasal Ventral scales smooth quadrangular smooth quadrangular smooth smooth smooth quadrangular quadrangular quadrangular Forelimb one apical scale one apical scale*** four apical scales two apical scales one apical scale Hindlimb one apical scale one apical scale one apical scale one apical scale four clawed toes continued. B. oxyrhina B. panoplia B. psamophila B. pyburni B. scolecoides Prefrontals absent present in contact absent present in contact present separated Supralabials five six six seven six First supralabial merged separated separated separated separated and nasal Scales around 29–30 43–47 35–38 41 36–40 midbody Contact 5th supralabial no contact 6th supralabial no contact no contact supralabial/parietal First temporal absent present absent present present Contact loreal/ no no no no no internasal Ventral scales smooth lanceolate keeled smooth lanceolate keeled smooth quadrangular quadrangular quadrangular Forelimb one apical scale four reduced fingers one apical scale four free fingers four reduced fingers Hindlimb one apical scale four reduced toes four clawed toes four free fingers four reduced toes

*Some individuals present seven supralabials. **Some individuals lack the first temporal, then the 6th supralabial is in contact with parietal. ***Some individuals present two apical scales.

Activity —Although not standardized observations were made, captive specimens, were only seen to be active at hours ranging between 15:00–22:30h. Behavior—All observed animals, but two, tried to escape using the same tactic in the wild and in both soil types during trials. They first slide quickly over the surface and then burrowed in the loose soil. Another, when found in the wild, jumped a distance of nearly half a meter and entered the leaf litter. A last specimen tried to directly burrow during trials, in both soil types. When grabbed by hand, all the individuals exhibited tail’s prehensile ability.

NEW SPECIES OF BACHIA Zootaxa 3616 (2) © 2013 Magnolia Press · 181 In the laboratory, one specimen was recorded whipping its tail in an apparently defensive display (http:// www.youtube.com/watch?v=yUi7luTTxlI). Another specimen used its tail as a long support to climb one of its terrarium’s walls (Fig. 6B). The four hatchlings found together, were further observed together in three occasions in the laboratory, along the following year, suggesting possible grouping behavior of this species among juveniles (Fig. 6C).

FIGURE 4. Distributional records of Bachia species of B. bresslaui group at the Central Brazil. Examined specimens are represented by symbols outlined in white; Type localities represented by central black dot. Two records of B. bresslaui at Planalto dos Gerais presented with an question mark were not examined and may represent B. geralista sp. nov. São Paulo state record presented with a question mark represents the unknown type locality for Bachia bresslaui.

182 · Zootaxa 3616 (2) © 2013 Magnolia Press TEIXEIRA JR ET AL. FIGURE 5. Habitat and microhabitat of Bachia geralista sp. nov., at Peruaçu valley region: (A) an individual and its track imprinted in the sandy soil; (B) typical cerrado vegetation found at the area; (C) a Pequi tree (Caryocar brasiliense); (D) the large cover of leaves under a Pequi tree; (E) detail of soil profile covered by a large layer of leaves; (F) general view of the regenerated cerrado habitat, dominated by Porcada bushes (Copaifera martii).

Discussion

We found great variation in head scalation among specimens of Bachia bresslaui examined (Fig. 7): while some individuals (Brasilia, DF) show the same head scale pattern of B. bresslaui’s holotype (Fig. 7A,B), in others the first temporal was lacking (Bataguassu, MS, and from an unknown locality) (Fig. 7 C,D), and some individuals have seven supralabials and two apical scales on forelimbs (from Mato Grosso and Mato Grosso do Sul border) (Fig. 7E,F). One individual has the first supralabial and nasal merged, and two apical scales in the forelimbs (MZUSP 10300, from Utiariti, MT) (Fig. 8). This last one was the second reported specimen of Bachia bresslaui in the literature (Vanzolini 1966).

NEW SPECIES OF BACHIA Zootaxa 3616 (2) © 2013 Magnolia Press · 183 FIGURE 6. Physiological and behavioral results: A) Box-plots represent maximum daily temperatures reached in the different microhabitats available to B. geralista sp. nov. Horizontal grey bars show temperatures voluntarily experienced by 10 individuals of B. geralista sp. nov. within laboratory thermal gradients. Dashed red line indicates highest temperature experienced by B. geralista sp. nov. in the lab, grey indicates voluntary maximum. B) Use of tail in one captive B. geralista sp. nov., arrow indicates cloacal region. C) Juveniles B. geralista sp. nov. grouped together within the terrarium.

Vanzolini noted some differences between this specimen and the holotype of B. bresslaui, like the contact between the loreal and internasal scales, preventing the contact between first supraocular and nasal (Fig. 7A, 8). This feature is absent in all other specimens of B. bresslaui (except one from Bataguassu, MT, MZUSP 78211) and B. geralista sp. nov., that we have examined (Fig. 1–3,6). However, Vanzolini (1966) did not mention the merging between first supralabial and nasal, and the two apical scales on forelimbs. These are exactly the diagnostic characters defining Bachia didactyla, a species recently described from the western Brazilian Cerrado, close to Utiariti (Freitas et al. 2011), in which there is no contact between the nasal and the first supraocular, the first supralabial and nasal are merged, and forelimbs end by two apical scales. Therefore, here we recognize this specimen (MZUSP 10300) as an individual of B. didactyla. The great variation found here, together with the recent descriptions of new species (Castrillon & Strüssmann 1998; Rodrigues et al. 2007; Rodrigues et al. 2008; Freitas et al. 2011) strongly suggest that a revision of Bachia bresslaui might uncover a cryptic diversity in the genus. By now, the paucity of specimens at each locality (often one, less often two) imposes difficulty in addressing this matter; the description of both B. didactyla and B. geralista sp. nov. were only possible due to the relatively large number of individuals known. The acquiring of additional molecular data and good morphological series through more directed surveys to find specimens seem necessary to solve this issue. This scarcity of representative series of Bachia bresslaui led it to be historically considered rare (MacLean 1973), despite Coli (1998) argued that it might just be inadequately sampled. However given our directed and standardized collection method, our data suggest that B. geralista sp. nov. is relatively rare locally, when compared

184 · Zootaxa 3616 (2) © 2013 Magnolia Press TEIXEIRA JR ET AL. to other open habitat fossorial limb-reduced gymnophthalmids, such as Calyptommatus spp., from the sand dunes of middle course of São Francisco River (Rodrigues 1991).

FIGURE 7. Variation on head scalation on Bachia bresslaui: A) the holotype from an unknown locality at São Paulo state (MZUSP 4737); B) from Brasilia, DF (MZUSP 91658), with a similar head scalation to the holotype; C) from an unknown locality (MZUSP 91599), showing contact between parietal and supralabial; D) from Bataguassu, MS (MZUSP 78211) also with parietal and supralabial in contact, and with no contact between frontal and nasal; E) from UHE Ponte de Pedra, MT (MZUSP 98760), showing seven supralabials, and also; F) from Itiquira, MT (MZUSP 99345).

Despite previous ecological observations of B. bresslaui were very limited (Colli 1998), B. bresslaui, B. dorbigny and B. flavescens, have been considered a fossorial, tail-reduced morph in the context of a global survey on the evolution of body elongation and limb-reduction in lizards (Wiens et al. 2006). Tail reduction in fossorial lizards has been attributed to a more efficient locomotion within the soil (Pianka & Vitt 2003). However, B. geralista sp. nov., and other Bachia species from the Cerrado, although having long tails, show morphological (e.g. reduced head scalation) and behavioral (e.g. burrowing when disturbed) traits indicating fossorial habits, being considered shallow burrowers, as other Bachia species (Pianka & Vitt 2003). These long tails are thought to serve as defense for predators, which would be effective via autotomy (Pianka & Vitt 2003). We observed the long tail of B. geralista sp. nov. being actively used in defensive displays (e.g. vibrating it, grabbing collectors’ finger) and seven out of 16 specimens had their tails autotomized and regenerated, which supports this explanation, although in all but one specimen, only the very tip of the tail was autotomized (apparently, less than a fifth of the complete tail).

NEW SPECIES OF BACHIA Zootaxa 3616 (2) © 2013 Magnolia Press · 185 However, they may also use the tail for overcoming obstacles and even jumping (behavior which has also been observed in other Bachia species; pers. obs.). Moreover, the observed specimens were not prone to autotomize their tails despite considerable handling. In sum, contrasting to what has been observed in other strictly fossorial gymnophthalmids like Calyptommatus, in which the tail is much reduced, we suggest that Bachia’s long tail is actually serving for a variety of functions, apart from being only an expendable target.

FIGURE 8. Specimen housed at Museu de Zoologia da Universidade de São Paulo (MZUSP 10300), from Utiariti, MT, the second specimen of Bachia bresslaui to be known, here recognized as the recently described B. didactyla, showing the first supralabial merged with the nasal, and supraocular not touching the nasal.

Temperatures reached in most of the potential microenvironments, seem to obligate B. geralista sp. nov. to seek refuge under the leaf litter of large trees (its actual microhabitat choice) during most of the day, however, even under the leaf litter there is still risk of overheating, and this risk may extend along the year, especially during the early evening. That might reduce the area of suitable microhabitats to a few thermally stable spots under the vegetation. Although we do not have critical minima for B. geralista sp. nov., the fifth percentile of environmental temperatures (11.34ºC) are below the critical minimum temperature observed for any other fossorial gymnopthalmid lizard from open areas (e.g. Calyptommatus spp, Camacho, unpub. data) and was experienced at all the microenvironments measured in July. Nonetheless, the loose sub soil remained at temperatures 7–9ºC above the absolute minimum registered during this study. That suggests that loose soil gives some degree of thermal protection against minimum extremes within the study region. The evening-nocturnal activity time displayed by B. geralista sp. nov. during laboratory observations, could be related to avoidance of critical maximum temperature reached during the warmest hours. Bachia are known to lay only two eggs (Colli 1998, and studies therein) but nothing is known about nest site selection or grouping behavior in Bachia. We did not find any pregnant individual but we found four hatchlings grouped together in July, and three well grown adults in January under the same microhabitat patch. That suggests that egg laying may occur in April–May, and that either B. geralista sp. nov. may reutilize or share their nest sites or that their hatchlings are able to detect co-specifics, possibly chemically. Posterior observations of the four hatchlings grouped in their terrarium during nearly a year, suggest they find some advantage in that behavior. However this is still speculative as we are still very far from understanding most of B. geralista sp. nov.’s biology. The endemicity and ecological traits of B. geralista sp. nov. make this species especially vulnerable to landscape changes, such as the wide agricultural expansion undergoing within the Cerrado (Alho & Martins 1995; Pinto 1994). This expansion is evident at Planalto dos Gerais, where most of the original vegetation was replaced by soy plantations. This region is a Mesozoic sandstone plateau that has been eroded for several million years producing large Cenozoic sandy deposits around and above it. The Jalapão area (northwestern end of Planalto dos Gerais) is a good example of a large sandy deposit, harboring a rich herpetofauna (Recoder et al. 2011; Valdujo et

186 · Zootaxa 3616 (2) © 2013 Magnolia Press TEIXEIRA JR ET AL. al. 2011) including several squamate endemic to that region somehow associated with sandy soils (Colli et al. 2003; Rodrigues et al. 2008; Colli et al. 2009; Ribeiro et al. 2009; Pinto & Curcio 2011). Nonetheless, most of Planalto dos Gerais is still far from being well inventoried. That has prevented until now the use of biogeographical or ecological information to direct conservationist policies. However several lines of evidence suggest that this region might be an important diversity center. These are: the discovery of the new species we describe herein, other fossorial squamates recently described (Ribeiro et al. 2011), our yet unpublished data and the recognition of this region as a Cerrado refuge (Werneck et al. 2012). All that testify in favor of its potential for harboring a diverse endemic fauna (Nogueira et al. 2011); and also that we are probably loosing most of its diversity while the agricultural frontiers are still spreading throughout its extent.

Acknowledgements

We are grateful to ICMBio officers at Parque Nacional Cavernas do Peruaçu, Evandro Silva, Flávio Túlio Gomes, Lilian Bonjorne de Almeida and Maria Carolina A. Camargos; and FIAT employees, Mr. Norinho, Ms. Nita and their family for help and support during field work; ICMBio for the authorization to work at the National Park (permit numbers 10129-2 and 17086-1); FAPESP, CNPq and CAPES for financial support; Hussam Zaher and Carolina Castro-Mello for access to specimens. We thank Adriana Yamanouchi, for help with laboratory experiments. We also thank Salvador Carranza for comments and suggestions on an early version of this manuscript. The senior author is especially grateful to Renato Kipnis for presenting Peruaçu valley region to him, rendering all this possible.

References

Alho, C.J.R., & Martins, E.S. (1995) De Grão em Grão, o Cerrado Perde Espaço. WWF-Fundo Mundial para a Natureza, Brasilia, 66 pp. Amaral, A. (1935) Estudos Sobre Lacertilios Neotropicos - II. Novo Genero e Espécie de Lagarto do Brasil. Memórias do Instituto Butantan, 9, 49–50. Avila-Pires, T.C.S. (1995) Lizards of Brazilian Amazonia (Reptilia: Squamata). Zoologische Mededelingen, 299, 1–706. http:// dx.doi.org/10.5962/bhl.title.59326 Bentz, E.J., Rodriguez, M.J.R., John, R.R., Henderson, R.W. & Powell, R. (2011) Population densities, activity, microhabitats, and thermal biology of a unique crevice- and litter-dwelling assemblage of reptiles on Union Island, St. Vincent and the Grenadines. Herpetological Conservation and Biology, 6, 40–50. Bonnaterre, P.J. (1789) Tableau Encyclopédique et Méthodique des Trois Règnes de la Nature. Erpétologie. Panckouke, Paris, 70 pp. Burt, C.E. & Burt, M.D. (1931) South American lizards in the collection of the American Museum of Natural History. Bulletin of the American Museum of Natural History, 61, 315–323. Cope, E.D. (1862) Catalogues of the reptiles obtained during the explorations of the Parana, Paraguay, Vermejo and Uruguay rivers, by Capt. Thos. J. Page, U.S.N., and those procured by Lieut. N. Michler, U.S. Trop. Eng., Commander of the expedition conducting survey of the Atrato River. Proceedings of the Academy of Sciences of Philadelphia, 14, 346–359. Cope, E.D. (1868) An examination of the Reptilia and Batrachia obtained by the Orton Expedition to Ecuador and the Upper Amazon, with notes on other species. Proceedings of the Academy of Sciences of Philadelphia, 20, 96–140. Cope, E.D. (1896) On the hemipenes of the Sauria. Proceedings of the Academy of Sciences of Philadelphia, 48, 461–467. Castrillon, M.I. & Strüssmann, C. (1998) A new species of Bachia and the occurrence of B. dorbignyi (Duméril and Bibron) in southwestern Mato Grosso, Brazil (Sauria, Gymnophthalmidae). Revista Brasileira de Zoologia, 15, 567–581. http:// dx.doi.org/10.1590/S0101-81751998000300001 Colli, G.R., Caldwell, J.P., Costa, G.C., Gainsbury, A.M., Garda, A.A., Mesquita, D.O., Filho, C.M., Soares, A.H., Silva, V.N., Valdujo, P.H., Vieira, G.H., Vitt, L.J., Werneck, F.P., Wiederhecker, H.C. & Zatz, M.G. (2003) A new species of Cnemidophorus (Squamata, Teiidae) from the Cerrado biome in Central Brazil. Occasional Papers Sam Noble Oklahoma Museum of Natural History, 14, 1–14. Colli, G.R., Giugliano, L.G., Mesquita, D.O. & França, F.G.R. (2009) New species of Cnemidophorus from Jalapão region, in the Central Brazilian Cerrado. Herpetologica, 65, 311–327. http://dx.doi.org/10.1655/08-049R1.1 Colli, G.R., Zatz, M.G. & da Cunha, H.J. (1998) Notes on the ecology and geographical distribution of the rare gymnophthalmid lizard, Bachia bresslaui. Herpetologica, 54, 169–174. Cowles, R.B. & Bogert, C.M. (1944) Preliminary study of the thermal requirements of desert reptiles. Bulletin of the American Museum of Natural History, 83, 261–296.

NEW SPECIES OF BACHIA Zootaxa 3616 (2) © 2013 Magnolia Press · 187 Daudin, F.M. (1802) Histoire Naturelle, génerale et particulière des reptiles, ouvrage faisant suite, a l'histoire naturelle, générale et particulière composée par Leclerc de Buffon, et redigée par C. S. Sonnini, vol. 4. F. Dufart, Paris, 397 pp. http:/ /dx.doi.org/10.5962/bhl.title.60678 Dixon, J. (1973) A systematic review of the teiid lizards, genus Bachia with remarks on Heterodactylus and Anotosaura. Miscellaneous Publications of the Museum of Natural History of the University of Kansas, 57, 1–47. Duméril, A.M.C. & Bibron, G. (1839) Erpétologie Générale on Histoire Naturelle Complète des Reptiles. Vol.5. Roret/Fain et Thunot, Paris, 871 pp. Ferreira, V.L., Petroni, D.M. & Caceres, N.C. (2005) Bachia bresslaui. Herpetological Review, 36, 78. Freitas, J.L., Strüssmann, C., Carvalho, M.A., Kawashita-Ribeiro, R.A. & Mott, T. (2011) A new species of Bachia Gray, 1845 (Squamata: Gymnophthalmidae) from the Cerrado of Midwestern Brazil. Zootaxa, 2737, 61–68. GBIF (2012) Global Biodiversity Information Facility - Data Portal, Global Biodiversity Information Facility. Available from: http://data.gbif.org/welcome.htm Gray, J.E. (1845) Catalogue of the specimens of lizards in the collection of the British Museum. Edward Newman, London, 289 pp. Henderson, R.W. & Powell, R. (2009) The state of natural history: a perspective from the literature on West Indian herpetology. Herpetological Review, 40, 273–275. Kizirian, D.A. & McDiarmid, R.W. (1998) A new species of Bachia (Squamata: Gymnophthalmidae) with plesiomorphic limb morphology. Herpetologica, 54, 245–253. Lichtenstein, H. & von Martens, E. (1856) Nomenclator reptilium et amphibiorum Musei Zoologici Berolinensis. Namenverzeichniss der in der zoologischen Sammlung der Königlichen Universität zu Berlin aufgestellten Arten von Reptilien und Amphibien nach ihren Ordnungen, Familien und Gattungen. Königliche Akademie der Wissenschaften, Berlin, 48 pp. http://dx.doi.org/10.5962/bhl.title.45245 MacLean, W.P. (1973) On the third specimen of Bachia bresslaui (Sauria, Teiidae). Papéis Avulsos de Zoologia, 27, 81–82. McDiarmid, R.W. & Foster, M.S. (1987) Additions to the Fauna of Paraguay with Notes on a Small Herpetological Collection from Amambay. Studies on Neotropical Fauna and Environment, 22, 1–9. Noble, G.K. (1921) Two new lizards from nothwestem Peru. New York Academy of Sciences, 29, 141–143. Nogueira, C., Ribeiro, S., Costa, G.C. & Colli, G.R. (2011) Vicariance and endemism in a Neotropical savanna hotspot: distribution patterns of Cerrado squamate reptiles. Journal of Biogeography, 38, 1907–1922. http://dx.doi.org/10.1111/ j.1365-2699.2011.02538.x Pianka, E.R. & Vitt, L.J. (2003) Lizards: windows to the evolution of diversity. University of California Press, Berkley, 333 pp. Pinto, M.N. (1994) Cerrado: Caracterização, Ocupação e Perspectivas - 2nd ed. Editora Universidade de Brasilia, Brasilia, 681 pp. Pinto, R.R. & Curcio, F.F. (2011) On the generic identity of Siagonodon brasiliensis, with the description of a new leptotyphlopid from Central Brazil (Serpentes: Leptotyphlopidae). Copeia, 53–63. http://dx.doi.org/10.1643/CH-09-119 Recoder, R.S., Teixeira Junior, M., Camacho, A., Sales Nunes, P.M., Mott, T., Valdujo, P.H., Ghellere, J.M., Nogueira, C. & Rodrigues, M.T. (2011) Répteis da Estação Ecológica Serra Geral do Tocantins, Brasil Centra. Biota Neotropica, 11, 263–281. http://dx.doi.org/10.1590/S1676-06032011000100026 Ribeiro, S., Castro-Mello, C. & Nogueira, C. (2009) New Species of Anops Bell, 1833 (Squamata, Amphisbaenia) from Jalapão Region in the Brazilian Cerrado. Journal of Herpetology, 43, 21–28. http://dx.doi.org/10.1670/07-299R1.1 Ribeiro, S., Nogueira, C., Cintra, C.E.D., Silva Jr, N.J. & Zaher, H. (2011) Description of a New Pored Leposternon (Squamata, Amphisbaenidae) from the Brazilian Cerrado. South American Journal of Herpetology, 6, 177–188. http://dx.doi.org/ 10.2994/057.006.0303 Rodrigues, M.T. (1991) Herpetofauna das dunas interiores do rio São Francisco: Bahia: Brasil. 1. Introdução à área e descrição de um novo gênero de microteiideos (Calyptommatus) com notas sobre sua ecologia, distribuição e especiação (Sauria, Teiidae). Papéis Avulsos de Zoologia, 37, 285–320. Rodrigues, M.T., Camacho, A., Nunes, P.M.S., Recoder, R.S., Teixeira Jr, M., Valdujo, P.H., Ghellere, J.M.B., Mott, T. & Nogueira, C. (2008) A new species of the lizard genus Bachia (Squamata: Gymnophthalmidae) from the Cerrados of Central Brazil. Zootaxa, 1875, 39–50. Rodrigues, M.T., Pavan, D. & Curcio, F.F. (2007) Two new species of lizards of the genus Bachia (Squamata, Gymnophthalmidae) from Central Brazil. Journal of Herpetology, 41, 545–553. http://dx.doi.org/10.1670/06-103.1 Rutven, A.G. (1925) Lizards of the genus Bachia. Proceedings of Boston Society of Natural History, 28, 101–109. Thomas, R. (1965) A new species of Bachia (Sauria: Teiidae) from Brasil. Herpetologica, 21, 18–22. Valdujo, P.H., Camacho, A., Recoder, R.S., Teixeira Junior, M., Beloti Ghellere, J.M., Mott, T., Sales Nunes, P.M., Nogueira, C. & Rodrigues, M.T. (2011) Anfíbios da Estação Ecológica Serra Geral do Tocantins, região do Jalapão, Estados do Tocantins e Bahia. Biota Neotropica, 11, 251–261. http://dx.doi.org/10.1590/S1676-06032011000100025 Vanzolini, P.E. (1961) Bachia: Especies Brasileiras e conceito generico (Sauria: Teiidae). Papéis Avulsos de Zoologia, 14, 193–209. Vanzolini, P.E. (1966) Sobre o segundo exemplar de Bachia bresslaui (Sauria, Teiidae). Papéis Avulsos de Zoologia, 19, 189–192. Werneck, F.P., Nogueira, C., Colli, G.R., Sites Jr., J. & Costa, G.C. (2012) Climatic stability in the Brazilian Cerrado: implications for biogeographical connections of South American savannas, species richness and conservation in a

188 · Zootaxa 3616 (2) © 2013 Magnolia Press TEIXEIRA JR ET AL. biodiversity hotspot. Journal of Biogeography, 3, 1–12. Werner, F. (1901) Reptilien und Batrachier aus Peru und Bolivien. Abhandlungen und Berichte des Zoologischen und Anthropologisch-Ethnographischen Museums zu Dresden, 9, 1–14. Wiens, J.J., Brandley, M.C. & Reeder, T.W. (2006) Why does a trait evolve multiple times within a clade? Repeated evolution of snakelike body form in squamate reptiles. Evolution, 60, 123–141. http://dx.doi.org/10.1111/j.0014- 3820.2006.tb01088.x

APPENDIX I. Specimens examined.

Bachia bresslaui: MZUSP 4737 (holotype) São Paulo, Brazil; MZUSP 78211 Bataguaçu, Faz. Água Branca, Mato Grosso do Sul, Brazil; MZUSP 98760 UHE Ponte de Pedra, Rio Corrente, Mato Grosso-Mato Grosso do Sul, Brazil; MZUSP 99345 Itiquira, Mato Grosso, Brazil; MZUSP 91658–91659 Brasília, Distrito Federal, Brazil; MZUSP 91599 no data. Bachia didactyla: MZUSP 10300 Utiarití, Mato Grosso, Brazil. Bachia micromela: MZUSP 91317 (holotype), MZUSP 91315–91316, 91318, 91030–91031 (paratypes) Guaraí, Tocantins, Brazil; Bachia oxyrhina: MZUSP 98080–98084 (paratypes) EESGT, Almas, Tocantins, Brazil; MZUSP 98086 (holotype) Estação Ecológica Serra Geral do Tocantins, Mateiros, Tocantins, Brazil. Bachia panoplia: MZUSP 57329, 57561–57562, 57638–57639, 57852, 58814 Manaus, Amazonas, Brazil. Bachia psamophila: CHUNB 24209, MZUSP 95079 (paratypes), MZUSP 95080 (holotype) UHE Luis Eduardo Magalhães, Tocantins, Brazil. Bachia scolecoides: MZUSP 3293 (holotype), MZUSP 3289–3295, 3337–3344 (paratypes) Rio Teles Pires, Mato Grosso, Brazil; MZUSP 38374 Sinop, Mato Grosso, Brazil; MZUSP 82525–82526 Fazenda Iracema, Cláudia, Mato Grosso, Brazil; MZUSP 103762–103766 Sapezal, Mato Gross, Brazil.

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