Herpetologica, 64(4), 2008, 452–457 E 2008 by The Herpetologists’ League, Inc.

ON THE IDENTITY OF JORDANENSIS HEYER, 1983 (ANURA: )

1,2 1 VANESSA K. VERDADE AND MIGUEL T. RODRIGUES 1Departamento de Zoologia, Instituto de Biocieˆncias, Universidade de Sa˜o Paulo, Caixa Postal 11.461 CEP: 05422-970, Sa˜o Paulo, Brazil

ABSTRACT: Cycloramphus jordanensis was described based on a single preserved specimen from Campos do Jorda˜o (22u 449 S, 45u 359 W), State of Sa˜o Paulo, Brazil. While examining the holotype; we noticed the presence of toe and tarsal fringes. Because these characters are absent in Cycloramphus, we suspected that the species was mistakenly placed in the genus. X-ray images of the holotype revealed T-shaped terminal phalanges and fang-like teeth. Together with the presence of toe and tarsal fringes, these characters squarely place the specimen in the genus . Considering the striking niche differences between Cycloramphus and Megaelosia, we expect the new combination will facilitate location of new individuals of this rare . Key words: Anura; Atlantic forest; Cycloramphus; Megaelosia; Megaelosia jordanensis (Heyer) comb. nov.;

THE GENUS Cycloramphus Tschudi, 1838 is characters, the presence of dermal scutes on presently allocated to Cycloramphidae (Frost the top of digital discs, and T-shaped terminal et al., 2006; Grant et al., 2006) and comprises phalanges (Frost et al., 2006). Like Cyclor- 27 nocturnal species (Brasileiro et al., 2007; amphus, hylodids are stream dwellers, living SBH, 2007) restricted to the Atlantic forest along the Atlantic forest and in open areas Domain (Ab’Saber, 1970). Most forms are beyond the tree line in eastern and southeast- associated with the fast flowing streams in ern Brazil (Frost, 2007; IUCN, 2006). How- montane areas of southern and southeastern ever members of the are primarily Brazil (Frost, 2007; Heyer, 1983a,b; Heyer diurnal, whereas Cycloramphus are nocturnal and Maxson, 1983). Relationships among . Cycloramphus species were recently assessed We took X-ray images of the holotype of in a phylogenetic analysis based on morpho- Cycloramphus jordanensis (MZUSP 4522) logical characters in Verdade’s (2005) unpub- using a Faxitron MX-20 machine with a lished Ph.D. thesis. According to this analysis, specimen radiography system formatted to Cycloramphus can be considered a monophy- take digital images captured under a 10 3 letic assemblage only if species of 5 cm area. Museum acronyms are Museu de Cope 1866 are included in the genus. We Zoologia da Universidade de Sa˜o Paulo, Sa˜o follow such an arrangement. Paulo, Brazil (MZUSP) and Museu Nacional, Cycloramphus jordanensis Heyer, 1983 was Rio de Janeiro, Brazil (MNRJ). described from a single preserved specimen The external morphology of the holotype of collected in Campos do Jorda˜o (22u 449 S, Cycloramphus jordanensis was described and 45u 359 W), State of Sa˜o Paulo, Brazil. No illustrated by Heyer (1983a: 308–309). Al- other specimen has been collected since the though we fully agree with most of Heyer’s original description. While examining the description, we did see the tympanum exter- holotype, we were surprised to find well- nally, and we clearly recognized a tarsal fringe developed toe and tarsal fringes. These on the specimen (Fig. 1). Heyer (1983a), characters are absent in Cycloramphus but however, described the existence of a hidden are considered to be a synapomorphy of tympanum and the presence of a tarsal fold. We believe that this difference may be due to Hylodidae (Frost et al., 2006; Grant et al., dehydration of the preserved specimen. Ad- 2006). Hylodids also share, among other ditionally, we noticed the existence of expand- ed toe tips bearing a barely visible pair of 2 CORRESPONDENCE: e-mail, [email protected] dorsal scutes (Fig. 2).

452 December 2008] HERPETOLOGICA 453

FIG. 1.—Tarsal fringe (arrow) on ventral left tarsus of Cycloramphus jordanensis holotype (MZUSP 4522); inner and outer metatarsal tubercles on left side of the picture.

Despite the lateral elements of the skull being obliterated by the hands of the speci- men, the X-ray images of the holotype (Fig. 3– 5) allow the following osteological character- ization. Premaxilla with an alary process projecting posterodorsally (Fig. 4); large fang-like teeth on premaxilla and maxilla (Figs. 4–5); vomers separated, with one vo- merine tooth (Fig. 4); palatines present, FIG. 2.—A pair of poorly developed dorsal scutes (arrow) on dorsal third toe of Cycloramphus jordanensis projecting parallel to the base of the skull holotype (MZUSP 4522). and widely separated (Fig. 3–4); pterygoids without a well developed ventral flange (Fig. 3); prootic and frontoparietal not fused parallel to the base of the skull. As pointed out (Fig. 3); condyles widely separated (Fig. 3); earlier, the external morphology of the spec- coracoid short, with anterior and posterior imen suggests that it is a member of family margins deeply concave (Fig. 3); clavicle Hylodidae, and the osteology strongly sup- slender and almost straight (Fig. 3); transverse ports this conclusion. Our osteological obser- process of presacral vertebra II the longest; vations agree with the detailed description of transverse processes of last presacral verte- Hylodidae osteology presented and illustrated brae shortened and projecting perpendicular- by Lynch (1971, under subfamily Elosiinae); ly to the vertebral axis; terminal phalanges T- in particular the T-shaped terminal phalanges shaped (Fig. 3); annulus tympanicus present and the alary processes of premaxillae with a (Fig. 5); maxilla massive with a distinctive lateral projection clearly place the species postorbital process (Fig. 5); condition of within the family. zygomatic process of the squamosal touching Hylodinae genera have been externally the maxilla not clear (Fig. 5); mandibles diagnosed on the basis of snout–vent length bearing an odontoid ridge (Fig. 5). The (SVL) ( and reaching images also show some of the damage inflicted SVL 30–45 mm; Megaelosia SVL 70–120 to the specimen: the pelvic girdle is dislocat- mm), condition of nuptial asperities (present ed, the left ilium shaft is broken near the base, in Crossodactylus and Hylodes; absent in the sacral vertebra is lacking, and the left Megaelosia), and vocal sac morphology (a humerus is broken, as are the maxilla and single median vocal sac in Crossodactylus; mandible (Figs. 3–5). paired lateral or absent vocal sac in Hylodes The following characters clearly exclude the and Megaelosia) (Lynch, 1971). In terms of specimen from Cycloramphus: annulus tym- osteological characters, Crossodactylus lacks a panicus visible externally, toe tips expanded quadratojugal. Megaelosia, in turn, is easily into small disks bearing dorsal scutes, pres- differentiated from the remaining Hylodidae ence of inner and outer metatarsal fold, a well by the presence of fang-like teeth (Cochran, developed tarsal fold, terminal phalanges T- 1955; Lynch; 1971), a massive construction of shaped, vomers widely separated bearing one maxillae, the presence of an enlarged zygo- large tooth, and palatines short and projecting matic ramus of the squamosal, which touches 454 HERPETOLOGICA [Vol. 64, No. 4

FIG. 3.—X-ray images of Cycloramphus jordanensis (MZUSP 4522). (A) Dorsal view of head and trunk. (B) Detail of terminal phalanx on left foot. Alary process of premaxilla (a), vomer (b), palatine (c), pterygoid (d), frontoparietal not fused to prootic (e), condyle (f), coracoid (g), clavicle (h), broken humerus (i), transversal process of vertebrae V (j), and broken ilium shaft (k). the maxilla, and the presence of a distinc- that C. jordanensis is a Megaelosia; accord- tive odontoid ridge in the mandibles (Lynch, ingly, we formally transfer this species to that 1971: 41, 169). Characters of external anat- genus: Megaelosia jordanensis (Heyer) comb. omy and our X-ray images clearly indicate nov. December 2008] HERPETOLOGICA 455

FIG. 4.—X-ray images of Cycloramphus jordanensis holotype (MZUSP 4522). Dorsal view of snout: alary process of premaxilla (a), fang-like tooth (b), vomerine tooth (c), and palatine (d). FIG. 5.—X-ray images of Cycloramphus jordanensis holotype (MZUSP 4522). Lateral view: phalanx (a), zygomatic process of squamosal (b), annulus tympanicus Currently, Megaelosia Miranda-Ribeiro (c), postorbital process of maxilla (d), broken mandible 1923 is comprised of six endemic species (e), T-shaped terminal phalanx (f), fang-like teeth (g), and living in the mountain streams of the Atlantic odontoid ridge on mandible (h). forest of southeastern Brazil: M. apuana Pombal, Prado and Canedo 2003, M. bocai- their semi-terrestrial tadpoles live on rocks in nensis Giaretta, Bokermann and Haddad the splash zone (Heyer, 1983a,b; Lutz, 1928, 1993, M. boticariana Giaretta and Aguiar 1929). Our field experience suggests that 1998, M. goeldi Baumann 1912, M. lutzae searches for Megaelosia should be more Izecksohn and Gouveˆa 1987 ‘‘1985’’, and M. successful during the dry season. massarti (Witte, 1930). Campos do Jorda˜ois The paucity of data on natural history of located in the Serra da Mantiqueira, between Megaelosia is striking. Except for M. goeldi, the ranges of M. boticariana and M. lutzae classified as Least Concern (LC) in the Global (Fig. 6). Giaretta and Aguiar (1998) referred Assessment database (IUCN, to a Megaelosia sp2 for the same mountain 2006), all other species are categorized as range, without specifying the locality. This Data Deficient (DD). It is therefore important specimen was never again mentioned in the to increase efforts searching for both new literature, but because its locality coincides populations of these frogs and additional with the Campos do Jorda˜o general area details concerning their natural history. These (Fig. 6) it may be possible that the specimen are top predators associated with corresponds to M. jordanensis. For now, we mountain streams at altitudes from 800– leave comparisons of M. jordanensis with its 2000 m a.s.l. in the Atlantic Forest areas, a congeners until new specimens have been well-known hotspot (Giaretta and collected. The main characters differentiating Aguiar, 1998; Giaretta et al., 1993; Myers et the species within Megaelosia are skin gran- al., 2000). Furthermore, the distributional ulation and development of finger and toe area of Megaelosia is entirely within the range discs and dorsal scutes (Giaretta et al., 1993). of occurrence of Batrachochitrium dendroba- These characters are barely assessable due to tidis in Brazil (Carnaval et al., 2006; Toledo et the poor condition of the holotype. al., 2006) suggesting a dangerous combination The new generic allocation proposed here of factors that may lead to population crashes may help in locating new specimens of M. in the near future (e.g., Lips et al., 2003, 2006; jordanensis. Despite occupying similar envi- Skerratt et al., 2007). ronments, the biology of Cycloramphus and Acknowledgments.—We thank H. Zaher and C. Castro- Megaelosia differ strikingly. Megaelosia are Mello (MZUSP), and J. P. Pombal Jr. and C. Canedo diurnal frogs (but see Pombal et al., 2003) (MNRJ) for access to specimens; M. C. Pinna and E. G. living on rocks near torrents; and Megaelosia Baena for the X-ray images; A. B. Carvalho and J. Cassimiro for photographs; IBAMA (process 2001. tadpoles live in slow-flowing pools of forest 004024/01-42) and IF/COTEC (process 42.495/2003) for streams. The stream-associated Cycloramphus collection permits; the staff of the Parque Estadual are nocturnal and live on rock crevices, and Campos do Jorda˜o for logistics and J. Cassimiro, F. F. 456 HERPETOLOGICA [Vol. 64, No. 4

FIG. 6.—Distribution of the Megaelosia in southeastern Brazil (adapted from Giaretta and Aguiar, 1998; including also localities listed at Appendix 1). Areas above 800 m are shaded in gray.

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