Zootaxa 3702 (5): 459–472 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.3702.5.5 http://zoobank.org/urn:lsid:zoobank.org:pub:E7D7BEC6-5DD6-47D6-BE26-904A69403C76 A new Crossodactylodes Cochran, 1938 (Anura: : Paratelmatobiinae) from the highlands of the Atlantic Forests of southern Bahia, Brazil

MAURO TEIXEIRA JR.1,2, RENATO SOUSA RECODER1, RENATA CECÍLIA AMARO1, ROBERTA PACHECO DAMASCENO3, JOSÉ CASSIMIRO1& MIGUEL TREFAUT RODRIGUES1 1Departamento de Zoologia, Instituto de Biociências, Universidade de São Paulo, São Paulo, SP, Caixa Postal 11.461, CEP 05422– 970, Brazil. 2E-mail: [email protected] 3University of California, Berkeley, Museum of Vertebrate Zoology, 3101 Valley Life Sciences Building, Berkeley, CA 94720–3140, USA.

Abstract

A new Crossodactylodes is described from Serra das Lontras, in the highlands of the Atlantic Forests of southern Bahia. The new species can be distinguished from all other Crossodactylodes by having Finger I ending in an acute tip, a larger body size, by cranial features, and by molecular data. Like their congeners, the new species live in bromeliads but is widely geographically disjunct, being apparently restricted to the summit of a mountain range in Northeastern Brazil.

Key words: Crossodactylodes septentrionalis sp. nov., Serra das Lontras, Atlantic Forests, mountain endemism

Introduction

The genus Crossodactylodes Cochran, 1938, family Leptodactylidae (sensu Fouquet et al. 2013), is a poorly known bromeliad-dweller genus endemic to the Brazilian Atlantic Forests. Crossodactylodes pintoi Cochran, 1938 is the type species and “Macahé [= Serra de Macaé]”, Macaé mountains, in Nova Friburgo, state of Rio de Janeiro, is the type locality (Bokermann 1966). Given the presence of spines on the inner surface of the first finger, a similar tooth development, and the supposed presence of digital pad glands that appears when the digital tips are dried out, Cochran (1955) suggested Crossodactylodes to be closely related to Crossodactylus Duméril and Bibron, 1841. However, this last character was probably an artifact resulting from the presence of Y-shaped terminal phalanges (Lynch 1971). Lynch (1971) placed Crossodactylodes as part of his Grypiscini tribe of leptodactylids, along with Cycloramphus Tschudi, 1838 and Zachaenus Cope, 1866, and reported its occurrence from “Guanabara” (a former Brazilian state, presently the Rio de Janeiro municipality) to Espírito Santo, along the Brazilian coast. The first extensive molecular phylogenetic study on did not included Crossodactylodes (Frost et al. 2006), but based on the phylogenetic relationships of its putative relatives, from which samples were available, the genus was allocated in the resurrected family Cycloramphidae. Posterior molecular analyses, still not including Crossodactylodes, lead Pyron and Wiens (2011) to create the subfamily Paratelmatobinae to harbor Paratelmatobius and Scythophrys. Crossodactylodes and Rupirana were not included in the analysis and remained as incertae sedis. However, they did not present a diagnosis for their new subfamily, rendering Paratelmatobinae a nomen nudum. This was later emmended by Ohler and Dubois (2012) which diagnosed the Paratelmatobinae. In a recent molecular study, Fouquet et al. (2013) included Crossodactylodes in the analysis and recovered it closely related to Paratelmatobius, Scythrophrys and Rupirana. Unaware of Ohler and Dubois (2012) previous diagnosis of Paratelmatobinae (sensu Pyron & Wiens 2011), Fouquet et al. (2013) proposed the new subfamily Crossodactylodinae (Fouquet et al. 2013), now including Crossodactylodes and Rupirana, rendering it a junior synonym of Paratelmatobinae Ohler & Dubois, 2012 ( Dubois 2013).

Accepted by M. Vences: 20 Aug. 2013; published: 29 Aug. 2013 459 Tadpoles of Crossodactylodes pintoi were described from Santa Teresa municipality, in Espírito Santo (Peixoto 1981). However in the following year, two additional species were described from this locality: Crossodactylodes bokermanni and C. izecksohni by Peixoto (1982). The larvae formerly attributed to C. pintoi belonged to some of the newly described species. Currently, the genus comprises the three above referred species, still known only from the vicinities of their type localities, except for Crossodactylodes pintoi from which no additional specimens have ever been collected since the type series was obtained in 1909 (Peixoto & Carvalho-e-Silva 2004; Silvano & Peixoto 2004a; b). Herein based on recently collected specimens from the highlands of the Atlantic Forests in southern Bahia, we add a new species to the genus, providing its phylogenetic placement, along with first data on its osteology and natural history.

Material and methods

Field samplings

The new species was found during an herpetofaunistic survey at Parque Nacional Serra das Lontras, Bahia, Brazil, in which pitfall traps and active search were used as sampling techniques. The pitfall traps were installed in four sets, ranging from 400 to about 600 m a.s.l. Each set consisted of five traps, with four 20 L buckets buried at the ground level, arranged in “Y” shape, and connected by a 4 m long and 50 cm high plastic fences. Traps remained open from 2nd to 9th March 2009, during the rainy season, totalizing an effort equivalent to 1,000 buckets/day. Active searches were carried out from 200 to 900 m a.s.l. by four to six people, during four to six hours per night, totalizing an effort of about 200 h/person, exploring the whole range of available habitats. Collected specimens were fixed in 10% formalin, preserved in 70% ethanol, and housed at Museu de Zoologia da Universidade de São Paulo (MZUSP).

External Morphology and Morphometrics

External morphology nomenclature follows Duellman (1970). Examination of morphological characters and measurements were taken after fixation, using a micrometric ruler and a Mitutoyo digital caliper (to the nearest 0.01 mm) under a stereomicroscope Zeiss STEMI SV6. Morphometric measurements were: snout-vent length, from the tip of snout to the vent (SVL); thigh length, from vent to knee (THL); tibia length, from knee to heel (TL); foot length, from tip of the longest toe to heel (FL); arm length, from arm insertion to elbow (AL); forearm length, from elbow to tip of the longest finger (FAL); eye-eye distance, between the anterior margin of orbit (EE); eye- nostril distance, between anterior margin of orbit to nostril (EN); eye diameter, measured horizontally (ED); head length, from posterior end of jaw to snout (HL), and head width, on its widest point (HW). Comparisons were made with voucher specimens housed at MZUSP (Appendix I) and with data from the literature (Lynch 1971; Peixoto 1982; Gomes 1988).

Osteology

Individuals from each currently recognized species (Crossodactylodes pintoi: MZUSP 104; C. bokermanni: MZUSP 58079; C. izecksohni: MZUSP 109013) and the holotype of the new species were scanned with a SkyScan digital microtomograph (www.skyscan.be), with a resolution of 9µm and no filters added. The images were processed in CT Analyzer v. 1.11 software, setting an automatic threshold point at the skull level, for filtering low- density matter (soft tissue). The resulting 3D models were visualized on CT Volume v. 2.2. Osteological nomenclature follows Duellman and Trueb (1994).

Phylogenetics

We generated partial sequences of four mitochondrial (12S, 16S, cyt b and COI) and two nuclear (TYR and RHO) genes from one individual of the new species, six individuals of Crossodactylodes izecksohni and five of C.

460 · Zootaxa 3702 (5) © 2013 Magnolia Press TEIXEIRA JR. ET AL. bokermanni. As outgroups we used: Paratelmatobius sp. (aff. cardosoi), Pleurodema diplolister, Lithodytes lineatus, lutzi, Leptodactylus mystaceus, Rupirana cardosoi, and Odontophrynus americanus from Fouquet et al. (2013) (Appendix II). Total genomic DNA was extracted from liver and muscle according to Fetzner (1999) and amplification was carried using standard PCR protocols with primers 16SAR and 16SBR (Palumbi 1996) for amplification of 16S, primers LGL765 and V for cyt b (Bickham et al. 1995; Palumbi 1996), primers t-Phe- and t-Val-frog for 12S (Wiens et al. 2005), primers dgLCO1490 and dgHCO2198 for COI (Folmer et al 1994), primers TYR1E and TYR1G for tyrosinase (Bossuyt & Milinkovitch 2000), and primers Rhod1A and Rhod1C for rhodopsin (Bossuyt & Milinkovitch 2000). The PCR cycle protocol consisted of one initial cycle of 94 °C for 5 min followed by 35 cycles of 94 °C for 40 sec, 50 °C for 16S and cyt b, 55 °C for 12S, 60 °C for tyrosinase and 50-55 °C for rhodopsin for 40 sec, 72 °C for 40 sec. PCR products were directly purified with Exonuclease I and Shrimp Alkaline Phosphatase (USB or Fermentas). Automated sequencing was carried out using BigDye Terminator v3.1 Cycle Sequencing kit (Applied Biosystems), followed by analysis on ABI Prism 310, 3700 or 3170 Genetic Analyzer Sequencers (Applied Biosystems) according to the manufacturer’s instructions. Sequences were edited in CodonCode Aligner (CodonCode Corporation). The resulting sequences were aligned in ClustalX v. 2. (Larkin et al. 2007), and combined on BioEdit v. 7.2 (Hall 1999) in a matrix of 3,667 bp (16S: 891 bp; 12S: 554 bp; cyt b: 691 bp; COI: 658 bp; TYR: 532 bp; RHO: 341 bp). We selected the best fit model of nucleotide substitution per marker using jModelTest v. 2.1 (Darriba et al. 2012) and the Akaike Information Criterion (AIC): GTR+G for 16S and 12S; GTR+I+G for cyt b and COI; SYM+I+G for TYR and HKY +I for RHO. We inferred phylogenetic relationships using Bayesian and Maximum Likelihood methods on a concatenated dataset. Heterozygous sites for both nuclear markers were treated as ambiguities. The Bayesian analysis was implemented in MrBayes v. 3.2 (Ronquist et al. 2012), available on Cipres Science Gateway (Miller, Pfeiffer & Schwartz 2010) with four partitions, each including a set of genes with the same selected substitution model. Two independent runs were performed, with a random starting tree, four incrementally heated Markov chains each, and 20,000,000 generations, with trees sampled every 1,000. Stationarity for each run was assessed using Tracer v.1.5 (Rambaut & Drummond 2009), after discarding 25% of the samples as burn-in. Nodes with posterior probability ≥ 95% on a 50% majority rule consensus tree from both runs were considered significant support for a given clade. Maximum likelihood (ML) bootstrapping (1,000 replicates) was performed in RAxML 7.4.4. (Stamatakis 2006) also available on Cipres Science Gateway (Miller, Pfeiffer & Schwartz 2010). The majority 50% consensus trees saved with posterior probabilities and bootstrap values on the nodes were visualized using FigTree 1.3.1 (http://tree.bio.ed.ac.uk/). Uncorrected genetic distances (p distances) were calculated using PAUP* v 4.0b10 (Swofford 2001).

RESULTS Taxon description Crossodactylodes septentrionalis sp. nov. (Fig. 1–3)

Crossodactylodes sp. 1 – Fouquet et al. (2013: 452, 454)

Holotype: MZUSP 150209 (Fig. 1–3) an adult female from the summit of Peito de Moça (15°9'47.68"S, 39°20'34.03"W, 931m a.s.l.), Serra das Lontras mountain range, Arataca municipality, Bahia state, Brazil, collected by the authors on March 6th, 2009. Field number MTR 16370. Etymology: The specific epithet septentrionalis is a Latin word that means “of the north” in reference to the geographical position where the new species was found, as it is the northernmost known Crossodactylodes species. Diagnosis: A Crossodactylodes by the combination of columella absent, bromelicolous habits, and the close genetic similarity with the former recognized species of this genus. It is diagnosed by (1) large female body size (SVL of 17 mm); (2) snout rounded in dorsal view; (3) granules on upper eyelids absent; (4) hindlimbs without transverse bars; (5) dorsal skin granular; (6) Finger I ending in an acute tip; (7) disks on Finger III and IV rounded; (8) disks on Toes IV and V. Description of the holotype: Body small, 17 mm in SVL. Head almost as long as wide (HW = 97% of HL).

NEW SPECIES OF CROSSODACTYLODES Zootaxa 3702 (5) © 2013 Magnolia Press · 461 Snout rounded in dorsal view, obtuse to slightly oblique in profile. Upper eyelids slightly granular. Canthal ridge blunt, loreal region slightly concave. No cranial crests. No postrictal tubercles. Eyes small, eye diameter 27% of head length, and 1.2 times the eye to nostril distance. Tympanum hidden. Nares slightly prominent, nostrils elliptical. Tongue rounded, free posteriorly. Vomerine teeth present, two on the left side, in a short row posteromedial to choanae. Choanae large, rounded, separated by a distance of about 3.5 times its diameter. Arms long and slender. Finger length I

FIGURE 1. Holotype of Crossodactylodes septentrionalis sp. nov. (MZUSP 150209; SVL = 17.0 mm), in dorsal (left) and ventral (right) views. Color in life: Background color orange/brown tones; an irregular cream lateral line, extending from posterior edge of orbit to inguinal region, separates dorsal from ventrolateral color which is of a darker brown; a brown “X” mark on dorsum between scapulae, outlined by a whitish irregular line; whitish irregular spots on vertebral line from the middle of head to urostyle; irregular darker brown marks scattered on dorsum; dorsal surface of fore limbs orangish; ventral surface pale brown with brighten cream small irregular marks on gular region, chest, fore and hind limbs; Iris reddish. Color in preservative: The same pattern described above, but faded. Iris becomes brown. Comparison to other species (characters of the species in comparison are presented in parentheses): The new species can be distinguished from all other Crossodactylodes by having Finger I ending in an acute tip (disc of Finger I rounded), a larger body size, 17 mm SVL in the only adult female known (C. bokermanni females 12–15 mm SVL; C. izecksohni females 11–14.4 mm SVL) and iris in vivid red (brown), and zygomatic process of squamosal bone long, passing the anterior margin of the optic foramen (short, not reaching the anterior margin of optic foramen) (Fig. 4). Additionally, the new species can be distinguished from C. bokermanni and C. izecksohni by the absence of granules on upper eyelids and by the absence of well-defined bars pattern on hindlimb (granules present; bars present). From C. bokermanni in having dorsal skin granular (smooth), rounded digital disks on Finger III and IV (elliptical), rounded digital disks on Toes IV and V (elliptical) and ventral surface of body and limbs pale brown (dark brown). By a relatively slender body when compared with C. izecksohni and C. pintoi (relatively more robust). From C. pintoi by its digital disks rounded (elliptical); rounded snout in dorsal view (truncated). From C. izecksohni and C. pintoi it can be distinguished by the presence of vomerine teeth (absent). Comparative measurements for the genus are presented in Table 1.

462 · Zootaxa 3702 (5) © 2013 Magnolia Press TEIXEIRA JR. ET AL. Distribution and natural history: Crossodactylodes septentrionalis sp. nov. is currently known only from the Peito de Moça peak, at Serra das Lontras, a pre-Cambrian mountain range (Nacif et al. 2009), emerging from the low coastal plain, Arataca municipality (Fig. 5). Despite the wide altitudinal range covered by our pitfall traps, the new species was only found through active search. The holotype, the only adult obtained, was caught along with a minute juvenile at the summit of the mountain, above 930 m a.s.l., inside a 1 m diameter bromeliad. Local vegetation consists of lower and sparse trees with thick leaves; the ground is covered by a dense layer of bromeliads, herbaceous plants, lichens and mosses. Epiphytic bromeliads are also very common and some areas show a bare rock with lichens (Fig. 6). This vegetation is very distinct from that of the surrounding lower areas, which harbor much higher trees, with closed canopy producing a shaded environment. We did not detect any individuals calling.

FIGURE 2. Holotype of Crossodactylodes septentrionalis sp. nov. (MZUSP 150209): Dorsal (A) and lateral (B) views of the head; palmar (C) and plantar (D) views. Scale bars = 5 mm.

FIGURE 3. Holotype in life of Crossodactylodes septentrionalis sp. nov. (MZUSP 150209).

NEW SPECIES OF CROSSODACTYLODES Zootaxa 3702 (5) © 2013 Magnolia Press · 463 FIGURE 4. Skull of the four Crossodactylodes species. From left to right, dorsal, ventral and lateral views. (A) C. pintoi (MZUSP 104); (B) C. bokermanni (MZUSP 58079); (C) C. izecksohni (MZUSP 109013); and (D) C. septentrionalis sp. nov. (MZUSP 150209). al p pm = alary process of premaxilla; dent = dentary; exoc = exoccipital; fp = frontoparietal; max = maxilla; mmk = mentomeckelian bone; nas = nasal; occ c = occipital condyle; opt f = optic foramen; pa = palatine; pm = premaxilla; pro f = prootic foramen; pro = prootic; prsph = parasphenoid; pt = pterygoid; qj = quadradojulgal; sph = sphenethmoid; spmax = septomaxilla; sq = squamosal; vom = vomer; vt = vomerine teeth.

Phylogenetic relationships: All three species of Crossodactylodes included in the molecular phylogenetic analysis were recovered in a clade with high support (pp = 1; bootstrap = 96%), where the new species is sister to C. izecksohni (pp = 0.99; bootstrap = 86%) (Fig. 7). The genetic divergence between C. septentrionalis sp. nov. and C. izecksohni averaged on 5% for 16S, 6% for 12S, and ranged between 13–14% for cyt b and 5–6% for COI. Between C. septentrionalis sp. nov. and C. bokermanni differences averaged on 6–7% for 16S, 7–8% for 12S, 15% for cyt b and 5–6% for COI. Distances between C. izecksohni and C. bokermanni averaged on 5–7% for 16S, 7–8% for 12S, 14–15% for cyt b and 5–7% for COI (Table 2). Genetic distances are low and fairly homogeneous between specimens of C. izecksohni from Santa Teresa. Surprisingly, this is not the case for C. bokermanni where MNRJ 38412 obtained at a close locality (Reserva Biológica de Santa Lúcia, Santa Teresa) differs strikingly from their closer relatives. Nuclear markers are less variable than the mitochondrials (0–2% among TYR and 0–3% among RHO), but no haplotype sharing was observed for TYR among the three species of Crossodactylodes analyzed. RHO showed haplotype sharing among some individuals of C. izecksohni and C. bokermanni. Only a few heterozygous sites

464 · Zootaxa 3702 (5) © 2013 Magnolia Press TEIXEIRA JR. ET AL. were found for nuclear markers among the individuals of each Crossodactylodes species (six sites for RHO, but only 3 shared by more than two individuals and three sites for TYR, and only one shared by two individuals).

TABLE 1. Comparative measurements of Crossodactylodes septentrionalis sp. nov., C. izecksohni, C. bokermanni, and C. pintoi. Abbreviations are explained in the text. The values are combinations of literature data and data from examined specimens. C. septentrionalis sp. nov. C. izecksohni C. bokermanni C. pintoi ♀♀♂♀♂♂ SVL 17.0 11.4–14.4 12.0–15.0 13–14 14–17 15–17 THL 7.4 12.0–6.3 5.3–6.3 5.7–7.3 5.7 7.0 TL 7.0 5.2–5.8 5.0–6.2 5.7–7.4 5.6 6.1 FL 9.7 7.5–8.5 7.1–8.8 7.8–10.1 7.4 8.8 FAL 8.0 5.5–6.3 4.8–6.2 5.5–7.1 5.7 6.6 ED 1.8 1.2–1.4 1.2–1.4 1.4–1.5 1.4 1.6 HL 6.6 4.6–5.5 4.5–5.3 5.1–5.6 4.7 6.7 HW 6.5 5.0–5.9 5.3–6.0 5.0–5.6 4.8 6.1

Discussion

Crossodactylodes was described more than seventy years ago (Cochran 1938) and yet it is a poorly known genus endemic to the Atlantic Forests, referred only in a small fraction of papers from the growing literature on Neotropical amphibians (e.g. Cochran 1938; 1955; Lynch 1971; Peixoto 1981; 1982; Heyer 1983; Gomes 1988; Haddad & Prado 2005; Frost et al. 2006; Pertel et al. 2006; Gasparini et al. 2007; Rödder et al. 2007; Haddad et al. 2008; Langone et al. 2008; Noleto et al. 2011; Pyron & Wiens 2011; Fouquet et al. 2012). In the original description of Crossodactylodes izecksohni and C. bokermanni, Peixoto (1982) highlights as diagnostic characters: the skin texture (smooth in C. bokermanni and granular in C. izecksohni), body robustness (robust in C. bokermanni and slender in C. izecksohni) and the presence of an “X” mark on dorsum (absent in C. bokermanni and present in C. izecksohni). We have examined over 50 specimens assigned to C. izecksohni, and five of C. bokermanni, from their type locality, Santa Teresa. Although the number of examined individuals from the later species is low, they seemed fairly homogeneous in these features; however, among the specimens of C. izecksohni we found a large variation in all these characters. The dorsum varied from smooth to granular, dorsal coloration from X-marked to fairly homogenous marble-patterned and body habitus from robust to slender. This wide variation renders difficult further detection of diagnostic morphological features between C. septentrionalis sp. nov. and its congeners, especially because most diagnoses are based on males and information on females is not frequent in the literature. Nevertheless, the shape of Finger I, the digital discs, body size and molecular data can adequately separate it from all other Crossodactylodes for the moment. Our data support previous findings for C. bokermanni and C. izecksohni showing that males reach larger SVL than females (Gomes 1988). Accepting that the new species follows the same pattern, males of C. septentrionalis sp. nov. would have body sizes reaching up further than 17 mm of SVL, the size of the single adult female known. Characters from internal morphology may also help intrageneric diagnoses. Gomes (1988) reported diagnostic osteological features between Crossodactylodes bokermanni and C. izecksohni (e.g. longer zygomatic process of squamosal bones in C. izecksohni, skull more elongated in profile in C. bokermanni), which were confirmed with our osteological data. The zygomatic process of squamosal is also a diagnostic feature for C. septentrionalis sp. nov., in which it appears longer than those in both above mentioned species, and also in C. pintoi, extending beyond the anterior border of optic foramen (Fig. 4). The presence of two fenestrae just anteriorly to the optic foramen could be an additional diagnostic feature of the new species although this might be due to an incomplete closuring of sphenethmoid due to immaturity; an unlikely possibility because oviductal eggs were observed. A suggested synapomorphy of Crossodactylodes among the Paratelmatobinae was the absence of columella (Gomes 1988; Fouquet et al. 2013). Our data corroborates this suggestion, as it is not observed in any currently known Crossodactylodes species.

NEW SPECIES OF CROSSODACTYLODES Zootaxa 3702 (5) © 2013 Magnolia Press · 465 TABLE 2. Uncorrected p distances of mitochondrial genes among Crossodactylodes septentrionalis sp. nov., C. izecksohni, and C. bokermanni; Upper table: 16S (below) and 12S (above) and; Lower table: genes cyt b (below) and COI (above). 123456789101112 1) MTR16370 - C. septentrionalis sp. nov 0.05 0.05 0.05 0.06 0.05 0.05 0.06 0.07 0.07 0.07 0.06 2) RBF350 - C. izecksohni 0.05 0.00 0.00 0.00 0.00 0.00 0.06 0.06 0.06 0.06 0.05 3) RBF351 - C. izecksohni 0.05 0.00 0.00 0.00 0.00 0.00 0.06 0.06 0.06 0.06 0.05 4) RBF352 - C. izecksohni 0.05 0.00 0.00 0.00 0.00 0.00 0.06 0.06 0.06 0.06 0.05 5) RBF517 - C. izecksohni 0.06 0.00 0.00 0.00 0.01 0.01 0.07 0.07 0.07 0.07 0.06 6) RBF592 - C. izecksohni 0.05 0.00 0.00 0.00 0.01 0.00 0.06 0.06 0.06 0.06 0.05 7) RBF598 - C. izecksohni 0.05 0.00 0.00 0.00 0.01 0.00 0.06 0.06 0.06 0.06 0.05 8) RBF625 - C. bokermmanni 0.06 0.06 0.06 0.06 0.07 0.06 0.06 0.00 0.00 0.00 0.04 9) RBF648 - C. bokermmanni 0.07 0.06 0.06 0.06 0.07 0.06 0.06 0.00 0.00 0.00 0.04 10) RBF652 - C. bokermmanni 0.07 0.06 0.06 0.06 0.07 0.06 0.06 0.00 0.00 0.00 0.04 11) RBF653 - C. bokermmanni 0.07 0.06 0.06 0.06 0.07 0.06 0.06 0.00 0.00 0.00 0.04 12) MNRJ38412 - C. bokermmanni 0.06 0.05 0.05 0.05 0.06 0.05 0.05 0.04 0.04 0.04 0.04 123456789101112 1) MTR16370 - - C. septentrionalis sp. nov 0.13 0.14 0.13 0.13 0.13 0.15 0.15 0.15 0.15 0.15 2) RBF350 - - C. izecksohni 0.13 0.00 0.00 0.01 0.01 0.14 0.14 0.14 0.14 0.15 3) RBF351 - - C. izecksohni 0.14 0.00 0.00 0.01 0.01 0.14 0.14 0.14 0.14 0.16 4) RBF352 - - C. izecksohni ------5) RBF517 - - C. izecksohni 0.13 0.00 0.00 0.01 0.01 0.14 0.14 0.14 0.14 0.15 6) RBF592 - - C. izecksohni 0.13 0.01 0.01 0.01 0.00 0.14 0.14 0.14 0.14 0.15 7) RBF598 - - C. izecksohni 0.13 0.01 0.01 0.01 0.00 0.14 0.14 0.14 0.14 0.15 8) RBF625 - - C. bokermmanni 0.15 0.14 0.14 0.14 0.14 0.14 0.00 0.00 0.00 0.07 9) RBF648 - - C. bokermmanni 0.15 0.14 0.14 0.14 0.14 0.14 0.00 0.00 0.00 0.07 10) RBF652 - - C. bokermmanni 0.15 0.14 0.14 0.14 0.14 0.14 0.00 0.00 0.00 0.07 11) RBF653 - - C. bokermmanni 0.15 0.14 0.14 0.14 0.14 0.14 0.00 0.00 0.00 0.06 12) MNRJ38412 - - C. bokermmanni 0.15 0.15 0.16 0.15 0.15 0.15 0.07 0.07 0.07 0.06

The absence of vomerine teeth was proposed to be a synapomorphy of C. izecksohni + C. pintoi (Fouquet et al. 2013). The new species has a small protuberance in the corresponding area of its left vomer, and a large one on the right vomer (Fig. 4D). The protuberances emerge externally as a series of small vomerine teeth only at the right vomer, showing that some variation on this character may occur. In our phylogenetic hypothesis, although the monophyly of Paratelmatobinae was not recovered, probably due to the low representativeness of Leptodactylidae in our analysis, the relationships within Crossodactylodes are well resolved, recovering C. septentrionalis sp. nov.

466 · Zootaxa 3702 (5) © 2013 Magnolia Press TEIXEIRA JR. ET AL. as sister to C. izecksohni, and C. bokermanni as sister to both. The unavailability of molecular data for C. pintoi, and given we have not produced a morphological matrix, prevented us from testing if the vomerine teeth is indeed a synapomorphy in the group.

FIGURE 5. Distributional records of the genus Crossodactylodes. Yellow line represents Parque Nacional Serra das Lontras boundaries; green area in the more general map represents the Atlantic Forest.

The molecular differences between MNRJ 38412 and other sequenced specimens of C. bokermanni living nearby at Santa Teresa, together with the morphological variation observed in C. izecksohni, also raises some questions, indicating the possibility of more undetected sympatric species. Moreover, we refrain from extending the discussion on cryptic species, as well as on character evolution in this genus, given the general limited data availability, as our osteological data comes from a single individual of each species, allowing us only to suggest hypothesis (rather than draw conclusions), and also because it will be addressed in an ongoing taxonomic revision of the genus (P. Garcia pers. comm.). Crossodactylodes septentrionalis sp. nov. and its sister species, C. izecksohni, are currently separated from each other by more than 540 km of lowlands (Fig. 5), while C. bokermanni occurs in sympatry with C. izecksohni at Santa Teresa mountain region. This puzzling distributional pattern does not change even if the future phylogenetic analyses including C. pintoi recover it as sister to C. izecksohni. Based on the evidence available, it is difficult to suggest hypotheses on speciation scenarios that could have led to this (currently known) distributional pattern. Montane regions, however, have already been recognized as important elements driving the diversification for diverse groups (Rodrigues et al. 2002; McCain 2005; Cadena et al. 2012; Zou et al. 2012), and this could also be the case in Crossodactylodes. The new species was only found at about 930 m a.s.l., C. bokermanni and C. izecksohni at about 650 m a.s.l. (Peixoto 1982; Silvano & Peixoto 2004a; b), and C. pintoi, 1200 m a.s.l. (Peixoto & Carvalho-e-Silva 2004).

NEW SPECIES OF CROSSODACTYLODES Zootaxa 3702 (5) © 2013 Magnolia Press · 467 FIGURE 6. Habitats of Crossodactylodes septentrionalis sp. nov.: (A) General view of the Atlantic Forest over Peito de Moça peak; (B) detail of the vegetation, with low trees covered with mosses, at the top of Peito de Moça peak; (C) bromeliad were the individuals were found.

FIGURE 7. Phylogenetic relationship among Crossodactylodes, inferred through a Bayesian analysis, based on mitochondrial (16S, 12S, cyt b and COI) and nuclear (RHO and TYR) genes. Values above branches indicate posterior probabilities and bootstrap values from the Maximum Likelihood analysis, respectively. Scale bar represents number of substitutions per site.

These coastal mountains usually work as barriers to the moisture coming from the sea, which are pushed towards the continent by the southeasterly trade winds. This results in adiabatic cooling, leading to condensation and ultimately orographic rain (Ackerman & Knox 2012) creating a cooler and wetter environment at higher altitudes, than those found at the surrounding lowlands. Thus, the restricted geographic distribution and the recurrent association of Crossodactylodes species with these highland habitats indicate that climate and habitat structure may have played an important role on its dispersal and isolation.

468 · Zootaxa 3702 (5) © 2013 Magnolia Press TEIXEIRA JR. ET AL. As it is currently recognized the Atlantic Forest has suffered dramatic fragmentation and expansion due climatic fluctuations of the past (Carnaval & Moritz 2008; Carnaval et al. 2009). Thus, during these fluctuations the environmental conditions, currently found only at highland areas, may had been found also along the lowlands, allowing their ancestor to disperse; and that when the climate started to change, their optimum environment was pushed upwards, becoming restricted to the higher areas, isolating the populations and resulting in this narrow occurrence, geographically and environmentally found today. These narrow endemisms also raise concerns on the future of these species as these habitats are among those more directly threatened by climatic changes (Williams et al. 2007). As the warmer temperatures overtake the higher elevations, the optimum microclimates for summit- specialists species would shrink or disappear, placing all Crossodactylodes species in great danger. The new species discovered here, together with those described from this particular region in the last five years (Cruz et al. 2008; Cruz & Napoli 2010; Recoder et al. 2010; Napoli et al. 2011; Lourenço-de-Morais et al. 2012; Teixeira Jr et al. 2012) and with other yet undescribed (pers. obs.), show that despite the exploration of the Atlantic Forest has started five centuries ago, the discoveries are still far from done. In fact they are ascending, and this severely threatened biome still harbors a highly diversified fauna waiting to be uncovered.

Acknowledgements

We thank Fundação de Amparo à Pesquisa do Estado de São Paulo (FAPESP), Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq), National Science Foundation (NSF, DEB 0817035 to C. Moritz), and CAPES-Fulbright (PhD fellowship to R. Damasceno DEB-2740/06-0) for financial support. We are grateful to Gabriel Rodrigues dos Santos from the Instituto de Estudos Socioambientais do Sul da Bahia (IESB) and Bird Life for allowing access to the Parque Nacional da Serra das Lontras, and providing logistical support. We are also grateful to M. Vences and an anonymous reviewer that commented early versions of this manuscript that improved it considerably. We also thank Sabrina Baroni and Jéssica Gillung for help in laboratory, Hussam Zaher and C. Castro-Mello, for access to specimens at MZUSP, Rodrigo Barbosa Ferreira for access to tissue samples, Nerivaldo and “Zezito”, from Fazenda Dois Braços, for their help in the fieldwork and Phillip Lenktaitis and Enio Mattos for the 3D scanned images. Instituto Chico Mendes de Conservação da Biodiversidade (ICMBio) provided collection permits (14555–5).

References

Ackerman, S.A. & Knox, J.A. (2012) Meteorology: understanding the atmosphere. Jones & Bartlett Learning, USA, 580 pp. Bickham, J.W., Wood, C.C. & Patton, J.C. (1995) Biogeographic implications of cytochrome b sequences and allozymes in sockeye (Oncorhynchus nerka). Journal of Heredity, 86, 140–144. Bokermann, W.C.A. (1966) Lista Anotatada das Localidades Tipo de Anfíbios Brasileiros. R.U.S.P. - Serviço de Documentação, São Paulo, SP, 183 pp. Bossuyt, F. & Milinkovitch, M.C. (2000) Convergent adaptive radiations in Madagascan and Asian ranid reveal covariation between larval and adult traits. Proceedings of the National Academy of Sciences of the United States of America, 97, 6585–6590. http://dx.doi.org/10.1073/pnas.97.12.6585 Cadena, C.D., Kozak, K.H., Goméz, J.P., Parra, J.L., McCain, C.M., Bowie, R.C.K., Carnaval, A.C., Moritz, C., Rahbek, C., Roberts, T.E., Sanders, N.J., Schneider, C.J., VanDerWal, J., Zamudio, K.R. & Graham, C.H. (2012) Latitude, elevational climatic zonation and speciation in New World vertebrates. Procedings of the Royal Society B, 279, 194–201. http://dx.doi.org/10.1098/rspb.2011.0720 Carnaval, A.C. & Moritz, C. (2008) Historical climate modelling predicts patterns of current in the Brazilian Atlantic forest. Journal of Biogeography, 35, 1187–1201. http://dx.doi.org/10.1111/j.1365-2699.2007.01870.x Carnaval, A.C., Hickerson, M.J., Haddad, C.F.B., Rodrigues, M.T. & Moritz, C. (2009) Stability Predicts Genetic Diversity in the Brazilian Atlantic Forest Hotspot. Science, 323, 785–789. http://dx.doi.org/10.1126/science.1166955 Cochran, D.M. (1938) Diagnoses of new frogs from Brazil. Proceedings of the Biological Society of Washington, 51, 41–42. Cochran, D.M. (1955) Frogs of Southeastern Brazil. United States National Museum Bulletin, 206, 1–423. http://dx.doi.org/10.5479/si.03629236.206.1

NEW SPECIES OF CROSSODACTYLODES Zootaxa 3702 (5) © 2013 Magnolia Press · 469 Cruz, C.A.G., Feio, R.N. & Nascimento, L.B. (2008) A new species of Phasmahyla Cruz, 1990 (Anura : Hylidae) from the Atlantic Rain Forest of the States of Minas Gerais and Bahia, Brazil. Amphibia-Reptilia, 29, 311–318. http://dx.doi.org/10.1163/156853808785111995 Cruz, C.A.G. & Napoli, M.F. (2010) A new species of smooth horned frog, genus Proceratophrys Miranda-Ribeiro (Amphibia: Anura: Cycloramphidae), from the Atlantic Rainforest of eastern Bahia, Brazil. Zootaxa, 2660, 57–67. Darriba, D., Taboada, G.L., Doallo, R. & Posada, D. (2012) jModelTest 2: more models, new heuristics and parallel computing. Nature Methods, 9, 772. http://dx.doi.org/10.1038/nmeth.2109 Dubois, A. (2013) Crossodactylodinae Fouquet, Blotto, Maronna, Verdade, Juncá, de Sá & Trefaut Rodrigues, 2013 is an invalid junior synonym of Paratelmatobiinae Ohler & Dubois, 2012 (Amphibia, Anura). Zootaxa, 3700, 499–500. http://dx.doi.org/10.11646/zootaxa.3700.4.9 Duellman, W.E. (1970) The Hylid frogs of Middle America. Monograph of the Museum of Natural History University of Kansas, 1–753. http://dx.doi.org/10.5962/bhl.title.2835 Duellman, W.E. & Trueb, L. (1994) Biology of . The Johns Hopkins University Press, Baltimore, 670 pp. Fetzner, J. (1999) Extracting high–quality DNA from shed reptiles skins: a simplified method. BioTechiniques, 26, 1052–1054. Folmer, O., Black, M., Hoeh, W., Lutz, R. & Vrijenhoek, R. (1994) DNA primers for amplification of mitochondrial cytochrome c oxidase subunit I from diverse metozoan invertebrates. Molecular Marine Biology and Biotechnology, 3, 294–299. Fouquet, A., Loebmann, D., Castroviejo-Fisher, S., Padial, J.M., Orrico, V.G.D., Lyra, M.L., Roberto, I.J., Kok, P.J.R., Haddad, C.F.B. & Rodrigues, M.T. (2012) From Amazonia to the Atlantic forest: Molecular phylogeny of Phyzelaphryninae frogs reveals unexpected diversity and a striking biogeographic pattern emphasizing conservation challenges. Molecular Phylogenetics and Evolution, 65, 547–561. http://dx.doi.org/10.1016/j.ympev.2012.07.012 Fouquet, A., Blotto, B.L., Maronna, M.M., Verdade, V.K., Juncá, F.A., de Sá, R. & Rodrigues, M.T. (2013) Unexpected phylogenetic positions of the genera Rupirana and Crossodactylodes reveal insights into the biogeography and reproductive evolution of leptodactylid frogs. Molecular Phylogenetics and Evolution, 67, 445–457. http://dx.doi.org/10.1016/j.ympev.2013.02.009 Frost, D.R., Grant, T., Faivovich, J., Bain, R.H., Haas, A., Haddad, C.F.B., de Sá, R.O., Channing, A., Wilkinson, M., Donnellan, S.C., Raxworthy, C.J., Campbell, J.A., Blotto, B.L., Moler, P., Drewes, R.C., Nussbaum, R.A., Lynch, J.D., Green, D.M. & Wheeler, W.C. (2006) The amphibian tree of life. Bulletin of the American Museum of Natural History, 297, 1–370. http://dx.doi.org/10.1206/0003-0090(2006)297[0001:tatol]2.0.co;2 Gasparini, J.L., Almeida, A.P., Cruz, C.A.G. & Feio, R.N. (2007) Os Anfíbios Ameaçados de Extinção no Estado do Espírito Santo. In: Passamani, M. & Mendes, S.L. (Eds.), Espécies da Fauna Ameaçadas de Extinção no Estado do Espírito Santo. Instituto de Pesquisas da Mata Atlântica, Vitória, ES, pp. 75–86. Gomes, N. (1988) Osteologia comparada de duas especies do genero Crossodactylodes Cochran, 1938 (Amphibia, Anura, Leptodactylidae). Memorias do Instituto Butantan, 50, 51–61. Haddad, C.F.B. & Prado, C.P.A. (2005) Reproductive modes in frogs and their unexpected diversity in the Atlantic forest of Brazil. BioScience, 55, 207–217. http://dx.doi.org/10.1641/0006-3568(2005)055[0724:rmifc]2.0.co;2 Hall, T. (1999) BioEdit: a user-friendly biological sequence alignment editor and analysis program for Windows 95/98/NT. Nucleic Acids Symposium Series, 41, 95–98. Heyer, W.R. (1983) Variation and systematics of frogs of the genus Cycloramphus (Amphibia, Leptodactylidae). Arquivos de Zoologia, 30, 235–339. Langone, J.A., Segalla, M.V., Bornschein, M. & de Sá, R.O. (2008) A new reproductive mode in the genus Melanophryniscus Gallardo, 1961 (Anura: Bufonidae) with description of a new species from the State of Paraná, Brazil. South American Journal of Herpetology, 3, 1–9. http://dx.doi.org/10.2994/1808-9798(2008)3[1:anrmit]2.0.co;2 Larkin, M.A., Blackshields, G., Brown, N.P., Chenna, R., McGettigan, P.A., McWilliam, H., Valentin, F., Wallace, I.M., Wilm, A., Lopez, R., Thompson, J.D., Gibson, T.J. & Higgins, D.G. (2007) Clustal W and Clustal X version 2.0. Bioinformatics Applications Note, 23, 2947–2948. http://dx.doi.org/10.1093/bioinformatics/btm404 Lourenço-de-Moraes, R., Solé, M. & Toledo, L.F. (2012) A new species of Adelophryne Hoogmoed and Lescure 1984 (Amphibia: Anura: Eleutherodactylidae) from the Atlantic rainforest of southern Bahia, Brazil. Zootaxa, 3441, 59–68. Lynch, J.D. (1971) Evolutionary relationships, osteology, and zoogeography of leptodactyloid frogs. Miscellaneous Publications of the Museum of Natural History of the University of Kansas, 53, 1–238. McCain, C.M. (2005) Elevational gradients in diversity of small mammals. Ecology, 86, 366–372. http://dx.doi.org/10.1890/03-3147 Miller, M.A., Pfeiffer, W. & Schwartz, T. (2010) Creating the CIPRES Science Gateway for inference of large phylogenetic trees. Proceedings of the Gateway Computing Environments Workshop (GCE), 14 Nov. 2010, New Orleans, LA, 8 pp. http://dx.doi.org/10.1109/gce.2010.5676129

470 · Zootaxa 3702 (5) © 2013 Magnolia Press TEIXEIRA JR. ET AL. Nacif, P.G.S., Costa, O.V., Araújo, M. & Santos, P.S. (2009) Geomorfodinâmica da Região do Complexo de Serras das Lontras. In: Brasil, S., IESB & International, B. (Eds.) Complexo de Serras das Lontras e Una, Bahia: Elementos naturais e aspectos de sua conservação. SAVE Brasil, São Paulo, pp. 9–14. Napoli, M.F., Caramaschi, U., Cruz, C.A.G. & Dias, I.R. (2011) A new species of flea-toad, genus Brachycephalus Fitzinger (Amphibia: Anura: Brachycephalidae), from the Atlantic rainforest of southern Bahia, Brazil. Zootaxa, 2739, 33–40. Noleto, R.B., Amaro, R.C., Verdade, V.K., Campos, J.R.C., Gallego, L.F.K., Lima, A.M.X., Cestari, M.M., Kasahara, S., Yonenaga-Yassuda, Y., Rodrigues, M.T. & Toledo, L.F. (2011) Comparative cytogenetics of eight species of Cycloramphus (Anura, Cycloramphidae). Zoologischer Anzeiger, 250, 205–214. http://dx.doi.org/10.1016/j.jcz.2011.04.001 Ohler, A. & Dubois, A. (2012) Validation of two familial nomina nuda of Amphibia Anura. Alytes, 28, 162–167. Palumbi, S.R. (1996) Nucleic acids II: The Polymerase Chain Reaction. In: Hillis, D.M., Moritz, C. & Mable, B.K. (Eds.), Molecular Systematics. Sinauer, Sunderland, MA, 205–247. Peixoto, O.L. (1981) Notas sobre o girino de Crossodactylodes pintoi Cochran (Amphibia, Anura, Leptodactylidae). Revista Brasileira de Biologia, 41, 339–341. Peixoto, O.L. (1982) Duas novas espécies de Crossodactylodes de Santa Tereza, Estado do Espírito Santo (Amphibia, Anura, Leptodactylidae). Revista Brasileira de Biologia, 42, 619–626. Peixoto, O.L. & Carvalho-e-Silva, S.P. (2004) Crossodactylodes pintoi. In: IUCN 2012. IUCN Red List of Threatened Species. Version 2012.2. Available from: http://www.iucnredlist.org (Accessed 28 May 2013) Pertel, W., Teixeira, R.L. & Rödder, D. (2006) Anurans inhabiting soil bromeliads in Santa Teresa, southeastern Brazil. Amphibia, 5, 16–19. Pyron, R.A. & Wiens, J.J. (2011) A large-scale phylogeny of Amphibia including over 2800 species, and a revised classification of extant frogs, salamanders, and caecilians. Molecular Phylogenetics and Evolution, 61, 543–583. http://dx.doi.org/10.1016/j.ympev.2011.06.012 Rambaut, A. & Drummond, A.J. (2009) MCMC Trace Analysis Tool Version v1.5.0. Institute of Evolutionary Biology, University of Edinburgh and Department of Computer Science, University of Auckland. Available from: http:// beast.bio.ed.ac.uk/ (Accessed 20 Aug. 2013) Recoder, R.S., Teixeira Jr., M., Cassimiro, J., Camacho, A. & Rodrigues, M.T. (2010) A new species of Dendrophryniscus (Amphibia, Anura, Bufonidae) from the Atlantic Rainforest of southern Bahia, Brazil. Zootaxa, 2642, 36–44. Rödder, D., Teixeira, R.L., Ferreira, R.B., Dantas, R.B., Pertel, W. & Guarneire, G.J. (2007) Anuran hotspots: the municipality of Santa Teresa, Espírito Santo, southeastern Brazil. Salamandra, 43, 91–110. Rodrigues, M.T., Dixo, M., Pavan, D. & Verdade, V.K. (2002) A new species of Leposoma (Squamata, Gymnophthalmidae) from the remnant Atlantic forests of the state of Bahia, Brazil. Papéis Avulsos de Zoologia, 42, 335–350. Ronquist, F., Teslenko, M., van der Mark, P., Ayres, D.L., Darling, A., Höhna, S., Larget, B., Liu, L., Suchard, M.A. & Huelsenbeck, J.P. (2012) MrBayes 3.2: efficient Bayesian phylogenetic inference and model choice across a large model space. Systematic Biology, 61, 539–542. http://dx.doi.org/10.1093/sysbio/sys029 Silvano, D. & Peixoto, O.L. (2004a) Crossodactylodes bokermanni. In: IUCN 2012. IUCN Red List of Threatened Species. Version 2012.2. Available from: http://www.iucnredlist.org (Accessed 28 May 2013) Silvano, D. & Peixoto, O.L. (2004b) Crossodactylodes izecksohni. In: IUCN 2012. IUCN Red List of Threatened Species. Version 2012.2. Available from: http://www.iucnredlist.org (Accessed 28 May 2013) Stamatakis, A. (2006) RAxML-VI-HPC: maximum likelihood-based phylogenetic analyses with thousands of taxa and mixed models. Bioinformatics, 22, 2688–2690. http://dx.doi.org/10.1093/bioinformatics/btl446 Swofford, D.L. (2001) PAUP*. Phylogenetic Analysis Using Parsimony (*and Other Methods) version 4.0 beta 10 for 32-bit Microsoft Windows. Smithsonian Institution, Washington, DC. Teixeira Jr., M., Dal Vechio, F., Recoder, R.S., Carnaval, A.C., Strangas, M., Damasceno, R.P., Sena, M.A. & Rodrigues, M.T. (2012) Two new species of marsupial tree-frogs genus Gastrotheca Fitzinger, 1843 (Anura, Hemiphractidae) from the Brazilian Atlantic Forest. Zootaxa, 3437, 1–23. Wiens, J.J., Fetzner Jr., J.W., Parkinson, C.L. & Reeder, T.W. (2005) Hylid frog phylogeny and sampling strategies for speciose clades. Systematics Biology, 54, 719–748. http://dx.doi.org/10.1080/10635150500234625 Williams, J.W., Jackson, S.T. & Kutzbach, J.E. (2007) Projected distributions of novel and disappearing climates by 2100 AD. Proceedings of the National Academy of Sciences of the United States of America, 104, 5738–5742. http://dx.doi.org/10.1073/pnas.0606292104 Zou, F.-S., Chen, G.-Z., Yang, Q.-F. & Li, Y.-D. (2012) Bird species richness along an elevational gradient in a forest at Jianfengling, Hainan Island, China. Zoological Studies, 51, 362–371.

NEW SPECIES OF CROSSODACTYLODES Zootaxa 3702 (5) © 2013 Magnolia Press · 471 APPENDIX I

Examined specimens Crossodactylodes bokermanni: Brazil: Espírito Santo, Santa Teresa, MZUSP 58077–79, MZUSP 73759–60 (paratypes). Crossodactylodes izecksohni: Brazil: Espírito Santo, Santa Teresa, MZUSP 16370, MZUSP 59519, MZUSP 96221, MZUSP 96226–27, MZUSP 108983–86, MZUSP 108988, MZUSP 108990–93, MZUSP 108995, MZUSP 108997, MZUSP 108999, MZUSP 109001, MZUSP 109003–05, MZUSP 109008–09, MZUSP 109012–13, MZUSP 109015–18, MZUSP 109020–22, MZUSP 109025, MZUSP 109027–31, MZUSP 109033–35, MZUSP 109037–39, MZUSP 109041, MZUSP 109047, MZUSP 109058, MZUSP 109060, MZUSP 109062, MZUSP 109074–75. Crossodactylodes pintoi: Brazil: Rio de Janeiro: Serra de Macaé, MZUSP 104 (paratype).

APPENDIX II.

GenBank accession codes for the terminals used in our phylogenetic analysis. Newly generated sequences are from Rodrigo Barbosa Ferreira (RBF) collection, from Reserva Biológica Augusto Ruschi, Santa Teresa, Espírito Santo. Published sequences are from individuals on Miguel Trefaut Rodrigues (MTR) collection are from Serra das Lontras, Arataca, Bahia; and Museu Nacional do Rio de Janeiro (MNRJ) from Estação Biológica de Santa Lúcia, Santa Teresa, Espírito Santo.

Taxon ID 16S 12S cyt b COI TYR RHO C. septentrionalis sp. nov. MTR16370 KC603958 KC603957 KC603963 KC603985 KC604077 KC604105 C. izecksohni RBF350 KF534641 KF534632 KF534660 KF534651 KF534678 KF534669 RBF351 KF534642 KF534633 KF534661 KF534652 KF534679 KF534670 RBF352 KF534643 - - - - - RBF517 KF534644 KF534634 KF534662 KF534653 KF534680 KF534671 RBF592 KF534645 KF534635 KF534663 KF534654 KF534681 KF534672 RBF598 KF534646 KF534636 KF534664 KF534655 KF534682 KF534673 C. bokermanni RBF625 KF534647 KF534637 KF534665 KF534656 - KF534674 RBF648 KF534648 KF534638 KF534666 KF534657 KF534683 KF534675 RBF652 KF534649 KF534639 KF534667 KF534658 KF534684 KF534676 RBF653 KF534650 KF534640 KF534668 KF534659 KF534685 KF534677 MNRJ38412 KC593359 KC593359 KC593349 - KC593364 - A. lutzi ROM40167 KC603952 KC603951 KC603974 KC604002 KC604093 KC604103 Le. mystaceus SMNS12036 KC603959 - KC603979 KC603992 JN691817 KC604108 Ly. lineatus AMNHA166426 AY843690 AY843690 - - - AY844683 O. americanus JF1891 AY843704 AY843704 - - - AY844695 Pa. sp. (aff. cardosoi) CFBH240 EU224408 EU224408 - - - DQ283814 Pl. diplolister MTR15393 - - KC603981 KC603986 KC604080 - R. cardosoi JC1112 KC603956 KC603955 KC603964 KC603987 KC604078 KC604106

472 · Zootaxa 3702 (5) © 2013 Magnolia Press TEIXEIRA JR. ET AL.