Zootaxa 1761: 59–68 (2008) ISSN 1175-5326 (print edition) www.mapress.com/zootaxa/ ZOOTAXA Copyright © 2008 · Magnolia Press ISSN 1175-5334 (online edition)

A large and enigmatic new eleutherodactyline (Anura, ) from Serra do Sincorá, Espinhaço range, Northeastern Brazil

JOSÉ CASSIMIRO1, VANESSA K. VERDADE2 & MIGUEL T. RODRIGUES3 Departamento de Zoologia, Instituto de Biociências, Universidade de São Paulo, Caixa Postal 11461, CEP 05422–970, São Paulo, Brazil. E-mail: [email protected]; [email protected]; [email protected]

Abstract

We describe a new of a large eleutherodactyline frog from the mountain rocky meadows (“campos rupestres”) of the Serra do Sincorá, Espinhaço mountain range, Mucugê municipality, State of Bahia, Brazil. The new species is promptly diagnosed from all the other Brazilian eleutherodactylines by its large size (males SVL 40.3–41.1; females SVL 75.2–79.7mm), broad head (head width 43–49% of SVL), presence of frontoparietal crests, pars fascialis of the maxilla deepened, discs absent on fingers, toes with poorly developed discs, first and second toes ridged, and tarsal fold absent. On the basis of these characters the new species is attributed to the up to now restricted to southern part of Central America and northwest part of South America.

Key words: Strabomantis aramunha sp. nov., , Strabomantidae, Amphibia, Atlantic forest, Campos rupestres,

Resumo

Descrevemos uma nova espécie de eleuterodactilíneo endêmico dos campos rupestres da Serra do Sincorá, Cadeia do Espinhaço, município de Mucugê, estado da Bahia, Brasil. A espécie é facilmente diagnosticada de todos os outros mem- bros brasileiros do grupo pelo seu grande tamanho (CRC dos machos 40.3–41.1; CRC das fêmeas 75.2–79.7 mm); cabeça larga (largura da cabeça 43–49% do CRC), presença de cristas frontoparietais, pars fascialis da maxilla profunda, discos ausentes nos dedos, artelhos com discos pouco desenvolvidos, primeiro e segundo artelhos com dobras dérmicas, e ausência de dobra tarsal. Com base nesses caracteres, a nova espécie é atribuída ao gênero Strabomantis, até o presente momento restrito ao sul da América Central e ao noroeste da América do Sul.

Palavras-chave: Strabomantis aramunha sp. nov., Eleutherodactylus, Strabomantidae, Amphibia, Mata Atlântica, Cam- pos rupestres, Taxonomia

Introduction

The genus Strabomantis is currently allocated in Strabomantidae (Hedges et al. 2008) and comprises 16 large species of forest-litter dweller , occurring from Costa Rica through the cloud forest of Colombia, Ecua- dor, Peru, and Amazon basin in western Brazil (Lynch 1975, 1997; Heinicke et al. 2007; Frost 2007; Hedges et al. 2008). The genus was resurrected by Hedges et al. (2008) that recently split the large assemblage of eleutherodactyline frogs (sensu Lynch 1976) in monophyletic units based on molecular data. Strabomantis is currently applied to all frogs previously belonging to the bufoniformis species series and Strabomantis bipor- catus (Hedges et al. 2008). Despite the many historical attempts to address relationships within “Eleuthero-

Accepted by M. Vences: 5 April 2008; published: 2 May 2008 59 dactylus” in smaller and coherent groups (e.g. Lynch 1975, 1976, 1979, 1980; Savage 1987; Hedges 1989; Lynch & Duellman 1997; Crawford & Smith 2005; Frost et al. 2006; Heinicke et al. 2007; Hedges et al. 2008), some of the proposed taxa lack unambiguous phenotypic diagnoses, and well-supported evidence of monophyly. The situation is particularly undefined for most species from Eastern Brazil, where the taxon sam- pling used to address relationships is inadequate (Frost et al. 2006; Heinicke et al. 2007; Hedges et al. 2008). While collecting and reptiles at Mucugê municipality (13° 00’ S and 41° 22’ W; Fig. 1), State of Bahia, Brazil, we found a new large broad-headed species of “Eleutherodactylus” bearing well-developed frontoparietal crests, living in the rocky meadows of Serra do Sincorá. Herein we describe this new species and, aware of the taxonomic complexity involving the Brazilian spe- cies of “Eleutherodactylus”, we discuss the resemblance of this highly distinctive and enigmatic species to other South American eleutherodactylines. We follow Lynch’s (1997) diagnosis of Eleutherodactylus sulcatus group, presently allocated in the genus Strabomantis to allocate the new species –– large size, broad head, posterior pars fascialis of maxillae deepened, V3 ramus of trigeminus nerve superficial to the m. levator pos- terior mandibulae subexternus (“S” condition of Lynch 1986).

Material and methods

The Serra do Sincorá is a high elevated area (reaching up 1600 m a.s.l.) located at the northern portion of the Espinhaço range. The collections in the area were made from 2 to 27 March, 2005, at the municipality of Mucugê (13°00’S, 41°22’W), State of Bahia, Brazil, using active search. Specimens were euthanized using alcohol 15%, fixed in 10% formalin, preserved in 70% ethanol, and deposited at the Museu de Zoologia da Universidade de São Paulo (MZUSP). Other specimens examined are listed in the Appendix 1. Geographical coordinates and elevations were obtained at collecting sites with a Magellan MAP 330 (map datum WGS 84). We studied the external morphology of preserved specimens, and verified the condition of V3 ramus of trigeminus nerve (Lynch 1986) by dissection. The sex was determined by the examination of gonads, the sex- ual maturity was inferred by the SVL of specimens (see Table 1 for minimum values of adult SVL) and gonads state of development. The following measurements were taken with calipers, to the nearest 0.1 mm (Table 1), according to Lynch and Duellman (1997): snout-vent length (SVL), head length (HL), head width (HW), eye-diameter (ED), eye-nostril distance (EN), interorbital distance (ID), tympanum diameter (TYD), tympanum height (TYH), tympanum-eye distance (TED), hand length (HAL), femur length (FL), tibia length (TL), foot length (FOL), length of the inner metatarsal tubercle (IMT).

Strabomantis aramunha sp. nov. (Figs. 1–3)

Holotype: MZUSP 138693 (field number JC-1212), an adult female, from Serra do Sincorá, Espinhaço range, 13° 04' 07" S and 41° 20' 09" W, 998 m elevation, municipality of Mucugê, State of Bahia, Brazil, collected by J. Cassimiro and F. S. F. Leite on 11 March 2005. Paratypes (N=6): MZUSP 138687 (JC-1198), an adult female, 13° 00' 33" S and 41° 22' 46" W, elevation 1191 m, on 8 March 2005; MZUSP 138688 (JC-1206), an adult male, 13° 00' 32" S and 41° 22' 48" W, eleva- tion 1207 m, same date; MZUSP 138690 (JC-1214), an adult male, 13° 03' 43" S and 41° 20' 26" W, elevation 947 m, on 11 March 2005; MZUSP 138689 (JC-1213), juvenile female, 13° 04' 07" S and 41° 20' 09" W, ele- vation 998 m, same date; MZUSP 138691 (JC-1251), juvenile male, on 20 March 2005; MZUSP 138692 (JC- 1277), juvenile female, 13° 01' 20" S and 41° 21' 24" W, elevation 1068 m, on 26 March 2005. All collected by J. Cassimiro and F. S. F. Leite at the surroundings of the type locality.

60 · Zootaxa 1761 © 2008 Magnolia Press CASSIMIRO ET AL. TABLE 1. Morphometrics (in millimeters) of the type material of Strabomantis aramunha sp. nov. For abbreviations of variables, see Material and Methods. Measurements of holotype in bold.

Variable MZUSP MZUSP MZUSP MZUSP MZUSP MZUSP MZUSP 138693 138687 138688 138690 138691 138689 138692 Adult female Adult female Adult male Adult male Young male Young female Young female SVL 75.2 79.7 40.3 41.1 32.6 40.2 46.0 HL 32.6 35.1 17.4 18.1 14.2 17.5 19.6 HW 36.4 39.5 17.7 19.0 14.4 17.4 19.9 ED 9.9 9.8 6.6 6.7 4.9 6.7 6.9 EN 10.0 10.2 5.5 6.1 4.7 5.7 5.9 TYD 4.5 4.7 2.5 2.6 2.1 2.9 2.9 TYH 5.5 6.3 3.3 3.2 2.7 3.3 3.6 TED 3.6 4.4 1.4 1.8 1.2 1.7 2.0 ID 7.9 9.1 3.9 3.9 3.1 3.7 4.9 HAL 19.3 20.6 10.9 11.2 9.1 10.7 12.2 FL 40.0 41.6 21.1 21.5 16.5 21.4 23.3 TL 38.3 40.8 21.3 22.5 17.0 21.5 24.3 FOL 34.9 38.3 19.1 20.1 15.6 19.9 21.5 IMT 3.7 3.5 1.7 1.9 1.3 2.1 2.2

Etymology: The specific name is from the Tupi language aramunha, meaning giant, in allusion to the large size of the species, and is used as noun in apposition. Diagnosis: A large broad-headed species (adult male 40.3–41.1 mm SVL, adult female 75.2–79.7 mm; HW respectively 44–46% and 48–49% of SVL), adult females bearing well-developed frontoparietal crests, posterior part of pars fascialis of maxilla deepened, mandibular ramus of the trigeminal nerve passing lateral to the m. levator mandibulae posterior subexternus, eyelid tubercles present posteriorly, not elongated, skin of venter coarsely areolate, leg relatively short (FL 51–53% of SVL) finger discs absent, first finger longer than second, poorly developed discs on toes, first and second toes ridged, fifth toe smaller than third, tarsal fold absent, testes white. Comparison to other species: The adult Strabomantis aramunha is promptly diagnosed from all other eleutherodactylines by the combination of large size, broad-head, flared maxilla, frontoparietal bearing well- developed crests in females, mandibular ramus of the trigeminal nerve passing lateral to the m. levator man- dibulae posterior subexternus, posterior eyelid tubercles present and not elongated, skin of venter coarsely areolate, absence of finger discs, first finger longer than second, poorly developed discs on toes, first and sec- ond toes ridged, fifth toe smaller than third, and, tarsal fold absent. These characters place squarely the new species in the Eleutherodactylus sulcatus group (Lynch 1997; Lynch & Duellman 1997), currently in the genus Strabomantis (Hedges et al. 2008). The only other species of Strabomantis occurring in Brazil is S. sul- catus (Lynch 1975, 1997; Frost 2007; Hedges et al. 2008). differ from S. aramunha by the smaller size (SVL 28–42 mm; Lynch 1975, 1980), concealed surfaces of thigh and shank presenting light blotches, and by the presence of tarsal fold. From the other species of Strabomantis, S. aramunha differ by the larger size (adult female S. cadenai, SVL 40.6 mm; Lynch 1997), skin of venter areolate (smooth in S. cade- nai, S. laticorpus; Lynch 1997), by not presenting elongate conical tubercles on eyelid (present in S. cadenai, S. cerastes, S. ingeri, S. laticorpus, S. necopinus, S. ruizi; Lynch 1997), large flattened warts absent (present in S. helonotus; Lynch 1997), tarsal fold absent (present in S. anomalus, S. cornutus, S. ingeri, S. necopinus, and S. ruizi; Lynch 1997), cranial crests present (absent in S. anomalus, and S. cheiroplethus; Lynch & Myers

NEW ELEUTHERODACTYLINE FROG FROM BRAZIL Zootaxa 1761 © 2008 Magnolia Press · 61 1983; Lynch 1990), reduced basal web only between first and third toes (extensive toe webbing in S. anoma- lus, S. anatipes and S. zygodactylus; Lynch & Myers 1983). Males and the young specimens, by being smaller and presenting narrower heads most resemble the adult (formerly “Eleutherodactylus” binotatus, Hedges et al. 2008). However, under close examination H. binotatus is promptly differentiated from S. aramunha by the smoother dorsal skin and the presence of tubercles not bearing keratinized tips (present in S. aramunha). Additionally, the head is narrower and longer than wide (wider than long in S. aramunha), the body is slender, the legs are longer, and the super- numerary tubercles are more numerous in H. binotatus than in S. aramunha.

FIGURE 1. Strabomantis aramunha sp. nov. (paratype, MZUSP 138687, adult female); from Serra do Sincorá, munici- pality of Mucugê, State of Bahia, Brazil. Photo by Felipe S. F. Leite.

Description of holotype: Head wider than long (HW = 36.4 mm; HL = 32.6 mm); a pair of frontoparietal crests externally visible; eyes dorsolateral, median sized (ED = 9.9 mm), diameter similar to the eye-nostril distance (EN = 10.0 mm); snout sub-elliptical in dorsal view, rounded in lateral view, overhanging lower jaw; nostrils small, slightly protuberant, directed laterally and positioned distally at the canthus rostralis; canthus rostralis sharp, accentuated by bony keel; loreal region deeply concave, tuberculate; flared maxilla, with pos- terior pars fascialis deepened; upper eyelid bearing small tubercles posteriorly, not elongated; interorbital space highly depressed, bearing small tubercles; tympanic membrane smooth; tympanic annulus prominent, higher than long (TYD = 4.5 mm; TYH = 5.5 mm), oblique; tympanum-eye distance less than tympanum diameter (TED = 3.6 mm); supratympanic fold evident, extending from posterior corner of eye to axils. Skin of dorsum spiculate under magnification, with tubercles more concentrated on posterior half; dorso- lateral ridges present; skin of venter coarsely areolate; anal opening plicate, surrounded by small tubercles; ulnar ridge slightly evident, evidenced by small and white tubercles; relative finger lengths I > III > II ˜ IV;

62 · Zootaxa 1761 © 2008 Magnolia Press CASSIMIRO ET AL. subarticular tubercles present; finger tips slightly expanded, not bearing discs; fingers free, not fringed or ridged; dorsal surface of thigh and shank tuberculate, tubercles with keratinized tips; hidden parts of thigh and shank smooth; no calcar ornamentation; sole of foot smooth; toe tips slightly expanded, with poorly devel- oped discs; toe relative lengths I < II < V < III < IV; reduced basal web between first and third toes; internal margins of first and second toes ridged.

FIGURE 2. Ventral view of left hand (A) and foot (B) of the holotype of Strabomantis aramunha sp. nov. Scale: 5 mm.

Color in life: Ground color of dorsum light brown, with two dark dorsolateral stripes extending from pos- terior corner of the eyelids to groin, medial dorsum with undefinite darker mottling, canthus rostralis dark brown, flanks barred from the dorsolateral stripes to venter, venter irregularly mottled with brown; dorsal sur- face of forearms, thighs and shanks barred; anterior and posterior surface of thigh uniformly faintly pigmented with brown; iris bronze. Color in preservative: Similar in life, although slightly faded. Measurements of holotype: SVL 75.2 mm, HL 32.6 mm, HW 36.4 mm, ED 9.9 mm, EN 10.0 mm, TYD 4.5 mm, TYH 5.5 mm, ID 7.9 mm, HAL 19.3 mm, FL 40.0 mm, TL 38.3 mm, FOL 34.9 mm, IMT 3.7 mm. Variation (N=6): Adult male and females of S. aramunha differ in size (male SVL 40.3–41.1 mm; female SVL 75.2–79.7 mm), HW/SVL ratio (males 0.44–0.46, females 0.48–0.49), development of flared maxilla,

NEW ELEUTHERODACTYLINE FROG FROM BRAZIL Zootaxa 1761 © 2008 Magnolia Press · 63 and presence of frontoparietal crests. Females are larger, present more developed flared maxilla, and are unique in presenting cranial crests. Males present the skin of dorsum spiculate with tubercles densely and evenly distributed, while in females, the density of tubercles is more variable, usually more concentrated on the posterior half. There is ontogenetic variation in the development of frontoparietal crests and labial flaring in females. In the smaller female (SVL 40.2 mm) of the type series cranial crests are absent; they are already seen in a slightly enlarged specimen (SVL 46 mm) becoming highly conspicuous in large specimens. Sexual dimorphism and ontogenetic variation observed is congruent with those reported for broad-headed eleuthero- dactylines (Lynch 1975, 1997; Lynch et al. 1994; Savage & Myers 2002).

FIGURE 3. Preserved adult male (left, MZUSP 138688, paratype) and female (right, holotype) of Strabomantis aramu- nha sp. nov.

Distribution and ecology: Strabomantis aramunha is presently known only and supposed to be endemic to the “campos rupestres” of the neighborhood of Mucugê municipality, Serra do Sincorá, Espinhaço Range (Fig. 4). The Espinhaço range is well known by the high level of species endemism (e.g. Bokermann 1956, 1964; Cunha 1966; Bokermann & Sazima 1973, 1978; Sazima & Bokermann 1978, 1983; Caramaschi & Saz- ima 1984, 1985; Vanzolini & Heyer 1988; Pinna 1992; Wege & Long 1995; Stattersfield et al. 1998; Heyer 1999; Lugli & Haddad 2006a,b; Napoli & Juncá 2006; Rodrigues et al. 2006). This mountain complex lies over Eastern Brazil at the transitional limits of the Atlantic forest and the Cerrado biomes with elevations from 700–2000 m a.s.l. (Stattersfield et al. 1998). The region is characterized by open and rocky areas domi- nated by shrubs and grasses referred to as “campos rupestres” or “campos de altitude” (for detailed character- ization of Brazilian “campos de altitude” see Safford 1999a,b). All specimens of S. aramunha were found at night, motionless on exposed rock outcrops in the “campos rupestres”. This is also the way broad-headed frogs have been frequently found as reported by Lynch (1997). Despite the rain, we did not hear the frogs calling. Nevertheless, the apparent absence of advertisement call is not infrequent among eleutherodactylines (Savage 1987; Savage & Myers 2002).

64 · Zootaxa 1761 © 2008 Magnolia Press CASSIMIRO ET AL. Remarks: This is the first record of an endemic eleutherodactyline from the “campos rupestres” of the Espinhaço range. These frogs are generally leaf-litter inhabitants that deposit their eggs in terrestrial humid habitats (Duellman 1978; Duellman & Trueb 1994). Presently the only recorded eleutherodactyline frog from these high elevation areas is the widespread ramagii (MTR unpublished data). In Brazil, eleuth- erodactyline frogs generally are associated with forested habitats, and, even those associated to non-forested biomes, like the Brazilian Cerrado, occur inside riparian forests. The single exception is Ischnocnema juipoca, known to occur in open areas (Haddad & Sazima, 1992). In this sense, the discovery of the new species Stra- bomantis aramunha from “campos rupestres” of the Espinhaço range is highly surprising and may represent a relict lineage evolved along with retraction of forests in this mountain complex (Giulietti & Pirani 1988).

FIGURE 4. Type-locality of Strabomantis aramunha sp. nov. (star), Mucugê municipality, State of Bahia, Brazil, and distribution of the genus Strabomantis (circles; except for S. biporcatus) in South America (Lynch, 1975, 1997; Frost, 2007).

The discovery of a large broad-headed eleutherodactyline in the high altitude mountains of Espinhaço range in eastern Brazil is enigmatic and reveals how complex is the understanding of the relationships and biogeography of these mountains. In the absence of an exhaustive phylogenetic study including east and southeastern South American Terrarana (Hedges et al. 2008), we can only rely on comparative examples to study the origin of Strabomantis aramunha. Phylogenetic and biogeographical relationships of endemic spe- cies from the Espinhaço range are virtually unexplored but some patterns are evident. Some frogs, like Thoropa megatympanum, Bokermannohyla alvarengai, B. nanuzae, B. oxente, B. saxicola, Rupirana car- dosoi, jandaia, otavioi, seem to have their closest relatives in species occurring in the eastern Atlantic forest (Bokermann 1956, 1964; Bokermann & Sazima 1973, 1978; Sazima & Bokermann 1983; Caramaschi & Sazima 1984; Heyer 1999; Lugli & Haddad 2006a,b). This is also the case of some liz-

NEW ELEUTHERODACTYLINE FROG FROM BRAZIL Zootaxa 1761 © 2008 Magnolia Press · 65 ards like the genera Placosoma and Enyalius (Cunha 1966; Rodrigues et al. 2006). A very distinctive and pos- sibly much older pattern is shown by species revealing relationships with species present in high elevation areas of northern and/or western South America. For example, species of Microlophus, the closest outgroup known for the lizard genus Eurolophosaurus, are Andean or transandean (Frost et al. 2001). Similarly, among the species of the cisandean Tropidurus of the torquatus group, Tropidurus mucujensis most resemble Tropi- durus bogerti, a species from the Ayuantepui in the Tepui region of Venezuela (Rodrigues 1987). Old Atlantic forest relationships are also disclosed by the relictual presence of the Atlantic forest anguid lizard Diploglos- sus fasciatus in western Amazon Basin (Ávila-Pires 1995). Again among lizards, the Rhachisaurinae, an endemic subfamily of gymnophthalmids restricted to these mountains was recently recognized (Pellegrino et al. 2001). Such examples are indicative of how far we are to understand the faunal relationships of this old, Precambrian ridge of eastern South America. The new species is placed in the genus Strabomantis based on morphological characters. However, males of S. aramunha are similar to Haddadus binotatus. This could indicate the generic misplacement of the spe- cies. Phylogenetic studies including species from a wide range of Eleutherodactylines (e.g. Craugastor, Had- dadus, , Ischnocnema, , , Strabomantis, etc.) would be needed to asses this and other hypothesis of relationships within this group of frogs.

Acknowledgments

We thank IBAMA, for collection permits (process number 02001.003933/01); F.S.F. Leite and L.E. Lopes for his valuable help in the field; H. Zaher and C. Castro-Mello for the access to specimens at MZUSP. We thank M. Vences and an anonymous reviewer who provided important and useful remarks that improved its quality. This work was supported by the Fundação de Amparo à Pesquisa do Estado de São Paulo (FAPESP) and CNPq (Conselho Nacional de Desenvolvimento Científico e Tecnológico).

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Appendix 1. Specimens examined

Haddadus binotatus: MZUSP 63557–63568 Fazenda Unacau, São José, Bahia, Brazil; MZUSP 53539–53555 Santa Ter- esa, Espírito Santo. : MZUSP 8084 Rancho Grande, Aragua, Venezuela. Strabomantis bufoniformis: MZUSP 120544–120545 Barro Colorado, Panamá. Strabomantis sulcatus: MZUSP 121859 Rio Vill- ano, Pastaza, Ecuador; MZUSP 121860 Loreto, Napo, Ecuador; MZUSP 121858 Cabeceras, Arajuno, Ecuador.

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