A new species of Coleodactylus Parker, 1926 (Squamata Sphaerodactylidae) from the Atlantic Forest of...

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ZOOTAXAISSN 1175-5326 (print edition)

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Zootaxa 3204: 20–30 (2012) www.mapress.com/zootaxa/ Article

A new species of Coleodactylus Parker, 1926 (Squamata: Sphaerodactylidae) from the Atlantic Forest of northeast Brazil

UBIRATAN GONÇALVES1, SELMA TORQUATO2,4, GABRIEL SKUK & GEORGE DE ARAÚJO SENA3

Museu de História Natural, Setor de Zoologia, Seção de Herpetologia, Universidade Federal de Alagoas. Avenida Aristeu de Andrade, 452, 57021-090, Farol, Maceió, Alagoas, Brazil. E-mail: [email protected]., [email protected], [email protected] author. E-mail: [email protected]

Abstract

A new species, Coleodactylus elizae sp. nov., is described for the Atlantic Forest of Northeastern Brazil. It is distinguished from other Coleodactylus species by the following character combinations: one nearly completely divided or two rostral scales, dorsal scales smooth and four scales forming the ungual sheath. The color pattern consists of white spots distributed over a pinkish background. Coleodactylus elizae sp. nov. is a forest species inhabiting bromeliads.

Key words: lizard, bromeliad fauna, rainforest, Brazil

Introduction

The genus Coleodactylus Parker (Squamata: Sphaerodactylidae) is distributed in South America east of the Andes range. Up to now, it was composed of five species of small leaf-litter diurnal geckos: C. amazonicus (Andersson), distributed in Central and Eastern Amazonia, including Southern Venezuela, Southern Guyana, Suriname, and French Guyana; C. brachystoma (Amaral) in the Cerrado in the Brazilian states of Goiás, Mato Grosso, Mato Grosso do Sul and Piauí; C. meridionalis (Boulenger), in the Atlantic Forest, from the coastal Brazilian states to more mesic open formations from Ceará to Bahia; C. natalensis Freire restricted to the forest in the dune fields of Natal and Parnamirim, in Rio Grande do Norte State, and C. septentrionalis Vanzolini, which occurs in the north-ern state of Roraima (Vanzolini 1957, 1980; Ávila-Pires 1995; Freire 1999; Vitt et al. 2005; Geurgas et al. 2008; Geurgas & Rodrigues 2010).

Until recently the characteristic considered most informative in recognizing genera of Sphaerodactylidae, due to its conservative nature, was digital scutellation, particularly of the ungual sheath. Kluge (1995) conducted a cla-distic analysis using 25 internal and external morphological synapomorphies. However, he explored the ungual sheath arrangement, based on the homonomy and heteronomy of individual digital scales, as the element of great-est weight to sustain the hypothesis regarding the historical evolution of the group. The most parsimonius hypothe-sis of his study established the following relationship (Gonatodes (Lepidoblepharis (Sphaerodactylus(Coleodactylus, Pseudogonatodes)))), corroborating the monophyly of each genus described in other studies (Parker 1926; Vanzolini 1968; Huey & Dixon 1970; Hoogmoed 1985).

More recently, molecular genetic studies have been carried out to help interpret relationships based on mor-phological aspects. The phylogenetic study of Coleodactylus conducted by Geurgas et al. (2008) based on mito-chondrial 16S and two nuclear genes, identified the monophyly of the genus, albeit weakly supported, as well as two main clades. The first composed of populations attributed to C. amazonicus and the second of C. meridionalis, C. brachystoma, C. natalensis and C. septentrionalis. The last was named as the C. meridionalis group.

However, in a phylogenetic study by Gamble et al. (2011), using fragments of five nuclear genes from New World geckos, established that Coleodactylus was polyphyletic with regard to other genera. Coleodactylus ama-zonicus was reported as the sister taxon to the remaining Sphaerodactylini, a clade consisting of Gonatodes, Lepi-

20 Accepted by A. Bauer: 6 Jan. 2012; published: 23 Feb. 2012


doblepharis, Pseudogonatodes, Sphaerodactylus and the other Coleodactylus species sampled: C. brachystoma, C. cf. brachystoma and C. septentrionalis. This relationship was strong support in oposition to the hypothesis of Coleodactylus monophyly.

An extensive morphological study using traditional characters did not reveal significant differences between groups of geographically distant populations within C. amazonicus (Ávila-Pires 1995), despite the variability in color pattern and meristic characters.

In this work we describe a new species of Coleodactylus found during a project aimed at assessing the effects of forest fragmentation on the structure of squamate reptile populations in two forest remnants of the Atlantic For-est in the Brazilian State of Alagoas.

Material and methods

Weekly two-day excursions related to the project “Effects of forest fragmentation on the structure of squamate rep-tile populations in two forest remnants of the Atlantic Forest in the Brazilian State of Alagoas” occurred between November 2005 and November 2006. Capture effort was 23,544 container days = total number of sampling days (218) x total number of containers (108) and 1104 man hours of active daily and nighttime searches calculated as total number of sampling days (46) x total number of persons (3) x hours of search per day (8) investigating litter, trunks and tree branches, as well as bromeliad and epiphytes, with the help of a rake and machete. The three speci-mens of Coleodactylus collected were photographed, euthanized by anesthesia, fixed in 10% formalin and pre-served in 70% ethanol. The material used to describe the new species is deposited in the collection of amphibians and reptiles of the Seção de Herpetologia do Museu de História Natural (MUFAL) at the Universidade Federal de Alagoas (UFAL). These specimens were compared with information in the literature and with specimens from sci-entific collections of other species of Coleodactylus (Appendix).

Morphometric analysis. All measurements and scale nomenclature follows Ávila-Pires (1995), and Kluge (1995), especially for the ungual sheath. The specimens obtained were compared with specimens of C. amazoni-cus, C. brachystoma, C. meridionalis, C. natalensis and C. septentrionalis. Measurements were taken to the nearest mm with a caliper ruler. The following characters were recorded for each fixed specimen: snout-vent length (SVL) —from tip of snout to cloaca and tail length (TL)—from cloaca to tip of tail; shape and number of rostral scale (RS); postrostral (PR); supranasal (SN); postnasal (PN); supralabial (SL); infralabial (IL); number of dorsal and ventral scales around the midbody (MBS); scales around the ungual sheath (USS); number and arrangement of lamellae under the fourth toe and the presence or absence of keeled scales.

Other characteristics were observed and used to describe the new species of Coleodactylus. Head length, width and height; shape of snout; mental scale width; number of postmental scales; height of the supraocular and supra-ciliary region; shape, arrangement and ratios between head scales; number of scales in a longitudinal line between the postnasal and the orbit; shape and length of neck, head, body, limbs and tail; shape of pupil; number of rows, size of subocular scales and relationship with other scales; shape and arrangement of scales on temporal and poste-rior region of the head; shape of ear opening; shape and arrangement of supralabials; shape and relationship between the mental and postmental and infralabial scales; number, shape and arrangement of postmentals up to the gular region;size and arrangement of infralabials; shape and arrangement of neck and dorsal scales; number of ven-tral scales along the medial ventral line, between the anterior level of the forelimbs and the anterior level of the hindlimbs, size and arrangement of ventral scales; shape of preanal scales; shape and arrangement of tail, limb, arm, and thigh scales; number of lamellae on the fourth finger, number, size and arrangement of ungual sheath scales; color pattern of head, body and tail.

Results

Distribution area. The specimens of the new taxon were obtained in an Atlantic Forest remnant of Serra da Sau-dinha (Saudinha range), Usina Cachoeira (Figure 1). This area is characterized as rainforest. The property is owned by an alcohol and sugar-producing company and is surrounded by extensive sugarcane plantations. The forest frag-ment covers about 1,210 hectares and is situated in the northern part of the municipality of Maceió at coordinates 09º 23’31” S and 35º 43’38” W. The area, a mountain range with an elevation of 304 meters, is composed mainly of

Zootaxa 3204 © 2012 Magnolia Press · 21COLEODACTYLUS FROM ATLANTIC FOREST OF BRAZIL


granite rocks (Assis 2000). There are a large number of rocky outcrops covered by ground bromeliads, which are also common as epiphytes. Two specimens of the new species were collected in the axils of epiphytic bromeliads of the species Aechmea muricata, at two meters above ground. The third specimen was collected from the rupicolous bromeliad Aechmea gustavoi.

FIGURE 1. Location of Serra da Saudinha, type locality of Coleodactylus elizae sp. nov. (star).

Holotype. MUFAL 3253, adult male; Serra da Saudinha, Municipality of Maceió, Alagoas, Brazil. 09°23’36.6”S, 35°43’30.6”W; collected by Ubiratan Gonçalves on December 12, 2005.

Paratypes. MUFAL 3252 and MUFAL 3755, adult females; same data as the holotype (Figure 2).Etymology. The species is named after Eliza Maria Xavier Freire, a researcher who has contributed signifi-

cantly to the knowledge of the herpetofauna of northeastern Brazil.Diagnosis. A forest species inhabiting bromeliads. SVL 26.2 mm in males, maximum SVL of 27.4 mm. Cylin-

drical body, slightly depressed with short limbs and tail. Claw on the fourth toe protected by ungual sheath com-posed of four asymmetrical imbricate scales. Among these is a dorsal scale that is half as long as the others, not reaching the apex of the sheath. Phylloid dorsal scales smooth and imbricated. Ventral scales larger than dorsal scales, smooth, in 48 rows (n=3). Scales around midbody 50 (n=3). Rostral divided or nearly completely divided. Diamond-shaped mental scale with a depression in the posteromedial portion that is continuous with groove between lower edge of infralabial and gular scales. Snout narrow with a ratio of mental width to head length of 0.28. Four postmentals. First infralabial long, with a ratio of mental width to length of 0.29 (Figure 3, Table 1).

Description. (Based on holotype - MUFAL 3253). Adult male, SVL 26.2 mm (Figure 1). Head 0.22 times SVL; 1.47 times longer than broad; 1.41 times broader than high. Snout narrow and depressed, rising slightly near the posterior margin of the head, ratio between mental scale width to head width 0.28. Four postmental scales.

GONÇALVES ET AL.22 · Zootaxa 3204 © 2012 Magnolia Press


Neck slightly narrower than head and body. Supraocular and supraciliary region above head level. Body nearly cylindrical, slightly depressed. Limbs relatively short, forelimbs 0.23 times SVL, hindlimbs 0.29 times SVL. Tail cylindrical in cross section, tapered towards the tip, 0.95 times SVL. Rostral divided, longer than broad, distinctly visible from above; posterior portion between the nostrils slightly depressed with a median fissure; bordered poste-riorly by four scales, with the lateral (supranasal) much larger than the others. Nostril bordered by the rostral (around 1/3 of the perimeter), first supralabial, two postnasals and one postrostral (supranasal); postnasal scales as large as adjacent loreals. Scales on the anterior area of the snout irregularly polygonal, smooth, flat, juxtaposed; decreasing in size, subimbricate, then granular towards the top of the head. Loreal region with scales similar to those on the anterior portion of the snout, decreasing in size posteriorly; 12 scales in a longitudinal line between the postnasal and the orbit. Scales of the supraorbital region flat, smooth, imbricate, smaller posteriorly. A supraciliary ridge is present with enlarged scales three times as long and wider than the supraciliary scales. They continue in the anteroposterior direction; posterior supraorbital scale followed by three rows of 6, 7 and 10 scales, respectively. Pupils round. Three rows of subocular scales; middle row with smaller scales; row next to supralabials with six scales, the first at mid-eye level, four times as long as the others. Scales on temporal region smooth, imbricate, irregularly polygonal. Scales on the posterior region of the head granular. Ear opening small and subtriangular. 6/6 supralabials, four extending to approximately below the posterior margin of the eye. First and second supralabial scales three times as long as high, third and fourth (below the eye) twice as long as high, fifth and sixth small, fol-lowed by three very narrow scalesup to the labial commissure. Sixth is about size of a granular scale. Mental scale rounded anteriorly, posterior margin edentate medially; posteromedial depression from the center of the scale con-tinuing in the postmentals; anterior and lateral margins of the mental scale boomerang-shaped, continuing along the margins of the first pair of infralabials, forming a convex edge; posterior median fissure absent. Four postmentals, irregularly polygonal, smooth, juxtaposed, subequal in size. Scales on the anterior portion of the jaw similar to the postmentals, decreasing gradually and changing to rounded, imbricate and granular in the gular region. Four infralabials decrease in size in the anteroposterior direction. First infralabial four times as long as high, second scale twice as long as high, third 1.5 times as long as high, fourth scale approximately equal in length and height and located below the central region of the eye. Five small scales continue to the labial commissure. Two rows of scales augmented between the second and third infralabials and small scales on the jaw. Scales on the neck forming a narrow transition zone between granular head scales and imbricate body scales. Dorsal scales smooth, imbricate, with posterior margin rounded. Ventral scales similar, larger; 48 scales along the midventral line between the ante-rior level of the forelimbs and cloaca; 52 up to the anterior margin of the hindlimbs. Dorsal and ventral scales in longitudinal and oblique rows, respectively. Scales around midbody 50, dorsal scales enlarged towards the ventral region. Scales on the preanal plate similar to the ventrals, except for scales on the cloacal edge, which are smaller. Anal plates absent. Scales on tail smooth, similar to dorsal and ventral scales. Scales on limbs smooth, imbricate, with posterior margin rounded; much smaller scales, nearly granular, on posterior surface of arms, forearms and thighs. Six lamellae on the fourth finger, nine on the fourth toe. Claws protected by an ungual sheath composed of four asymmetrical scales on the fourth finger, imbricate, which nearly cover the entire claw: In this species we dis-tinguished a single dorsal, a superolateral and two inferolaterals. The enlarged, more ventral, bulbous, pilose and dark scale of the sheath is presumably a composite, including two scales, whose original positions would be one postaxial ventral + one inferolateral. As observed by Kluge (1995), the asymmetry of the digit appears to be a sim-ple function of the larger size of postaxial ventral in relation to the other member of the pair. Fusion of scales in the infradigital region, increasing pad area, may have evolved along with the likely habit of climbing, irrespective of C. amazonicus. The dorsal scale is proximal, half as long as the others, not reaching the apex of the sheath.

Paratype variation. (Female): SVL (n=2) 26.1–27.4mm; tail length 22.6mm (n=1); hindlimb 7.8–7.9mm (n=2); two rostral scales or one nearly completely divided; six postnasal scales; 5–6 lamellae on the fourth toe (Table 1).



FIGURE 2A. Coleodactylus sp. nov. paratype, MUFAL 3252. Photo by Gabriel Skuk. B. Dorsal, left lateral, and ventral views of the head of Coleodactylus elizae sp. nov. holotype (MUFAL 3253).

Live coloration of holotype. Rostral dark brown; postrostral with blotches extending across the entire nasal region and preocular edge. Area from the nasal depression to the ocular region white with dark brown reticulations, posterior area of head pink with dark brown reticulations, mainly on the median line, where there is a concentration of white blotches that reach the edge of the nuchal marking. Nuchal marking U-shaped, white with dark brown margin, starting in the postocular area and joining its mate at the back of the neck; nuchal marking becomes gradu-ally wider in the anteroposterior direction, starting at two granules wide and reaching six granules at the median line. White band starting after the labial commissure, following the level of the ear, ending at the base of the fore-limb, where it is less distinct. Dark wide bands alternating with narrow white bands from the lips to the lateral neck area, up to the edge of the ear opening; the last of these white bands runs behind the mouth, to the forelimb. Dorsal region pink and lateral regions light colored; four white dorsolateral blotches, tail dark orange with five white dor-



solateral blotches, larger than those on the body; white dorsolateral band extending from the anterior margin of the hind feet to the sacral region, where it is more distinct and symmetrical; this band is bordered in the sacral region by a black and dark brown line; the rest of its length is light brown sprinkled with dark brown spots. There are three to four semi-spherical white blotches between the dorsolateral bands: three at the level of the anterior margin of the hindlimbs that are more distinct and one near the posterior margin of the hindlimbs that is less distinct; two white spherical blotches on the dorsal face of the forelimbs. Dorsal surfaces of the limbs, tail, feet and hands with fewer dark brown spots.

Coloration variation in live paratypes. Tail dark orange with two white dorsolateral blotches without black border (Figure 2).

FIGURE 3A–C. Dorsal, left lateral, and ventral view of the head of Coleodactylus elizae sp. nov. holotype. D. Palmar view of right hand (first finger to the right). E. Plantar view of right foot (first toe to the left). F. Palmar view of fourth toe from left foot (from nearly 45º angle). H. Apical view of fourth toe. G. Apical view of third toe tip (left foot). A–C are to the same scale. Sim-ilarly D–E and F–H are to the same scale, respectively. Scales of the finger and toe: a—ventral, b—lateral, c—ventral + lateral, d—dorsal, 1—superolateral, 2—inferolateral.

Coloration in preservative. Orange tail color becomes grey; white dorsal blotches less distinct.Paratype variation in preservative MUFAL 3252. White dorsolateral and dorsoventral bands and white

blotches at the tip of the body less distinct; white blotches on hindlimbs less distinct; hindlimbs and tail lighter in color. Abdomen white, sprinkled with dark brown spots; margins of the head, neck, the rear two-thirds of the body, tail and limbs with greater pigment concentration; sparse black spots on the rest of the body.



Discussion

Comparison with other Coleodactylus species. Despite the limitations in characters traditionally used in taxon-omy of the group to interpret intra and interspecific relationships, Coleodactylus elizae sp. nov. exhibited a combi-nation of clearly distinct traits of the remaining species of its genus: number and shape of rostrals (one to two, longer than wide; against one, wider than long in all remaining species), shape and length of snout (long and depressed; in remaining species it is short and higher); size (can reach slightly over 27 mm; among the others C. brachystoma is the largest, reaching around 26 mm, while C. natalensis and C. amazonicus are the smallest at slightly over 22 mm); a larger number of post-nasals (six to seven, against two to five for C. brachystoma and C. amazonicus); ungual sheath (four scales, five in remaining species, except C. amazonicus with four too); dorsal scales (smooth, the same of other species, except C. amazonicus with keeled scales) and pholidosis larger for post-nasals and infralabials (Table1). Furthermore, for the remaining pholidosis characters considered, higher counts were shown by the other two species from eastern Brazil, C. meridionalis (post-rostrals, supra nasals and supra labials), distributed from Ceará to Bahia and sympatric with C. elizae sp. nov., endemic to Serra da Saudinha and C. natalensis (ventrals), also sympatric with C. meridionalis, but more restricted to the dunes of Rio Grande do Norte (Table 1).

Coleodactylus elizae sp. nov. exhibited apparent sexual dimorphism with respect to body size, the females being larger than the males. This characteristic was also observed by Freire (1999) in three species of the genus (C. natalensis, C. meridionalis and C. septentrionalis). Freire (1999) also describes sexual dichromatism in C. natalen-sis, a characteristic observed in C. elizae sp. nov., where the male exhibited more detailed coloration, mainly on the tail.

In relation to color pattern, C. elizae sp. nov. resembles individuals of C. septentrionalis from the state of Roraima, in that they exhibit a transverse white band on the posterior part of the head, extending to the posterior edge of the eye; body and tail with a series of white blotches and a white U-shaped band in the sacral region. How-ever, the background color may be brown or reddish brown and the white band on the head is less evident near the eyes.

Complementary morphological studies such as osteological investigations can increase the number and quality of characters currently used in the taxonomy of Sphaerodactylidae, helping clarify the relationship between Coleo-dactylus elizae sp. nov. and the C. amazonicus group, with which it shares the presence of four ungual sheath scales and with the Coleodactylus meridionalis group, whose scales are smooth and sympatrically distributed. Coleodactylus elizae sp. nov. exhibits sympatry with C. meridionalis, although it is not syntopic, given that the lat-ter is found in undergrowth and all the individuals of C. elizae sp nov. were obtained in epiphytic or rupicolous bro-meliads, isolated from litter. The individuals are skillful climbers, in both their natural habitat and the laboratory. They are even able to climb smooth surfaces, suggesting that their habitat is arboreal. Such a conclusion is rein-forced by the absence of records of this species in terrestrial bromeliads, which were consistantly examined during collection excursions. However, despite the significant effort, the small number of lizards obtained in this species precludes definitive conclusions about its ecology.

The genetic study C. elizae sp. nov. and of individuals of C. meridionalis from Serra da Saudinha should pro-vide important information on the evolutionary history of the genus.

The study by Geurgas et al. (2008), based on the mitochondrial 16S gene and two nuclear genes of the genus Coleodactylus found the existence of two clades, C. amazonicus and the C. meridionalis group, is supported by morphological differentiation: C. amazonicus is the only species that has keeled dorsal scales, whereas in the C.meridionalis group they are smooth. The keeled state of the dorsal scales seems to be apomorphic, given that it is not present in the genera related to Coleodactylus. Another distinctive morphological characteristic is the number of scales that form the ungual sheath: four in C. amazonicus and five in the C. meridionalis group. The latter char-acteristic is considered one of the few and most phylogenetically informative autopomorphies of C. amazonicus(Vanzolini 1980; Ávila-Pires 1995; Freire 1999). Meristic and other color pattern variations (Vanzolini 1980; Ávi-la-Pires 1995; Freire 1999) do not show geographical trends within species of genus Coleodactylus, unlike molec-ular studies, which recognized five haplotypes in populations of C. amazonicus (Geurgas et al. 2008; Geurgas & Rodrigues 2010).

The discovery of a new species of Coleodactylus in the Eastern Atlantic Forest supports the idea that the area exhibits unique historical and biological characteristics. A recent phylogenetic study of Phyllomedusinae identified genetic differences in the population of Agalychnis granulosa Cruz from Serra da Saudinha (Faivovich et al. 2010).



Moreover, remaining forest fragments of Usina Cachoeira shelter 39 species of reptiles inventoried during the proj-ect entitled “Effects of forest fragmentation on the structure of squamate reptile populations of two Atlantic Forest remnants in the State of Alagoas, Brazil” among these is a population of Tropidurus aff. hygomi (data unpublished). The known distribution of T. hygomi Reinhardt & Luetken encompasses coastal sand dunes and sandy soils near the caatinga of the states of Sergipe and Bahia (Rodrigues 1987; Rodrigues 1988, Rodrigues 2003). In other studies on the herpetofauna of forest remnants in the state of Alagoas, including areas the type locality of C. elizae sp. nov., four new species of snakes, Bothrops muriciensis Ferrarezzi & Freire, 2001; Liotyphlops trefauti Freire, Caramaschi & Argôlo, 2007; Atractus caete Passos, Fernandes, Bérnils & Moura-Leite, 2010; Dendrophidion atlantica Freire, Caramaschi & Gonçalves, 2010 and six of amphibians, Chiasmocleis alagoanus Cruz, Caramaschi & Freire, 1999; Phyllodytes edelmoi Peixoto, Caramaschi, & Freire, 2003; Phyllodytes gyrinaethesPeixoto, Caramaschi, & Freire, 2003; Hyla studerae Carvalho-e-Silva, Carvalho-e-Silva & Izecksohn, 2003;Scinax skuki De Lima, Cruz & Azevedo Júnior, 2011 and Scinax muriciensis Cruz, Nunes & De Lima, 2011 were described. These facts indicate the importance of conserving the rainforests of the region, regardless of the size of the remnant or other aspects related to fragmentation, given that their biodiversity reflects the history of environ-mental evolution. Serra da Saudinha, as well as most other Atlantic Forest remnants in the state of Alagoas are not legally protected in conservation units (Menezes et al. 2004). This is an extremely worrisome situation, especially in light of the significant ongoing destruction of Atlantic Forest remnants in the aforementioned state.

Acknowledgements

Thanks to Filipe Augusto C. do Nascimento and Katyuscia Vieira, who took part in field collections and to Teresa Ávila-Pires from the Museu Emílio Goeldi and Fernanda Werneck and Guarino Colli from Universidade de Brasilia (UNB) for the use of the material. We thank the Fundação de Amparo à Pesquisa do Estado de Alagoas, for financing the project: “Effects of Forest Fragmentation on the Structure of Squamate Reptile Populations of Two Atlantic Forest Remnants in the State of Alagoas, Brazil. Protocol no. 2004 0930160–1. The Usina Caeté S. A., Cachoeira unit provided logistical support. The Instituto Brasileiro do Meio Ambiente e dos Recursos Renováveis–IBAMA issued collection licence no. 184/05–IBAMA/RAN.

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APPENDIX. Material examined.

Coleodactylus septentrionalis:Farm Bom Intento, Boa Vista, Roraima, Brazil: MPEG 4182, 4375, 4445.

Coleodactylus meridionalis:Serra da Saudinha Forest, Maceió, Alagoas, Brazil: MUFAL 3337, 3339, 3491, 3492, 3565, 3638, 3899, 3699, 3733, 4042.Catolé Forest, Maceió, Alagoas, Brazil: MUFAL 523, 531, 532, 533, 534, 537, 538, 539, 543, 545, 549, 550, 552, 553, 556,

557, 559, 561, 566, 567, 634, 1629, 1633, 1635, 6653, 6656, 6658Ecological Station of Xingó, Alagoas and Sergipe, Brazil: MUFAL 6501, 6502, 6504, 6505, 6506, 6508, 6509, 6511, 6512,

6514, 6548, 6661, 6662, 6663

Coleodactylus natalensis:Coastal sand dunes of Natal, Rio Grande do Norte, Brazil: MUFAL 0471, 0474, 0475, 2278, 2279, 2280, 2281, 2282.

Coleodactylus brachystoma:Minaçu, Goiás, Brazil: CHUNB 13140, 13180, 13261, 13263, 13266, 13267, 13282, 13283, 13288, 13289, 13292, 13293,

13295, 13297, 13300, 13301, 13303, 13304, 13306, 24278.

Coleodactylus amazonicus:Oriximiná Pará, Brazil: MPEG: 15369, 15372, 15368.Melgaço, Pará, Brazil: MPEG: 16466,16596, 16568, 16459, 16634, 16640, 17820.Santarém, Pará, Brazil: MPEG: 17429, 17526, 17559, 17582.Ouro Preto do Oeste, Rondônia, Brazil: MPEG: 14063.

List of acronyms of collectionsCHUNB—Universidade de BrasíliaMPEG—Museu Paraense Emílio GoeldiMUFAL—Museu de História Natural da Universidade Federal de Alagoas


A new species of Coleodactylus Parker, 1926 (Squamata Sphaerodactylidae) from the Atlantic Forest of...

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