Description of a New Species of Worm Salamander (Caudata

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Description of a new species of worm salamander (Caudata, Plethodontidae, Oedipina) in the subgenus Oedopinola from the central portion of the Cordillera Nombre de Dios, Honduras

Article in Zootaxa · August 2011 DOI: 10.11646/zootaxa.2990.1.4

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Description of a new species of worm salamander (Caudata, Plethodontidae, Oedipina) in the subgenus Oedopinola from the central portion of the Cordillera Nombre de Dios, Honduras

JAMES R. MCCRANIE1,3 & JOSIAH H. TOWNSEND2 110770 SW 164th Street, Miami, FL 33157–2933, USA 2School of Natural Resources and Environment, University of Florida, Gainesville, Florida 32611, USA, and Centro Zamorano de Bio- diversidad, Escuela Agrícola Panamericana Zamorano, Departamento de Francisco Morazán, Honduras 3Corresponding author. E-mail: [email protected]

Abstract

We describe a new species of Oedipina (subgenus Oedopinola) from Parque Nacional Pico Bonito in the central portion of the Cordillera Nombre de Dios in north-central Honduras. The new species was previously assigned to O. gephyra, but a combination of molecular analyses and its differences in foot morphology diagnose it from O. gephyra.

Key words: Oedipina, subgenus Oedopinola, Oedipina petiola sp. nov., Honduras, Parque Nacional Pico Bonito, morphology, mtDNA, 16S, cytochrome b

Resumen

Describimos una nueva especie de Oedipina (subgénero Oedopinola) del Parque Nacional Pico Bonito en la porción central de la Cordillera Nombre de Dios en el nor-central de Honduras. La nueva especie fue previamente asignada a O. gephyra, pero una combinación de análisis moleculares y diferencias en la morfología de los pies la diagnostican aparte de O. gephyra.

Introduction

Plethodontid worm salamanders of the genus Oedipina Keferstein are among the most challenging neotropical salamanders to study, due to their secretive fossorial habits and the resulting infrequency with which they are encountered (Brame 1968, García-París & Wake 2000, McCranie 2006a, Sunyer et al. 2010). The application of molecular phylogenetic analyses to systematic study of Oedipina has shown that several populations of these salamanders represent cryptic lineages warranting species-level recognition, and in some cases entire divergent clades were concealed by the relative lack of morphological distinctiveness that is typical of the genus (García-París & Wake 2000; McCranie et al. 2008; Sunyer et al. 2010, 2011). The new species described herein can now be added to that grow- ing list of cryptic species. García-París & Wake (2000) provided a phylogenetic analysis of the genus Oedipina using two mitochondrial genes (cytochrome b and 16S DNA). García-París & Wake (2000) resurrected Oedopinola Hilton as a subgenus for a clade containing, among others, O. gephyra McCranie, Wilson & Williams, a species previously considered to inhabit two isolated localities in north-central Honduras (Figure 1). Oedipina gephyra was described from a locality on the leeward side of Reserva de Vida Silvestre Texíguat in the western portion of the Cordillera Nombre de Dios, Honduras (McCranie et al. 1993), and subsequently reported from Parque Nacional Pico Bonito in the central portion of the Cordillera Nombre de Dios (McCranie 1996). A single representative from each of the two populations assigned to O. gephyra was included in the phylogenetic analyses of García-París & Wake (2000). Those

Accepted by M. Vences: 8 Jul. 2011; published: 9 Aug. 2011 59 analyses indicated that two species might be concealed under that nominal form based on interspecific-level sequence divergence between the two samples (16S= 5.25%; cyt b= 6.61%; García-París & Wake 2000, 70). How- ever, with only one sample each from the western and central localities, adequate characterization of intraspecific and interspecific variation would await collection of additional specimens of these rarely encountered animals. Despite frequent visits by herpetologists to the vicinity of Parque Nacional Pico Bonito, no additional specimens have been uncovered subsequent to the discovery of a single sub-adult male in 1995 (McCranie 1996). However, several recently collected samples of O. gephyra from the species’ type locality in the western portion of the Cor- dillera Nombre de Dios were collected in 2008 and analyzed for this study, and support the recognition of the western and central populations as distinct species. Comparison of the single specimen from Parque Nacional Pico Bonito with a series of O. gephyra from its type locality revealed differences in foot morphology between the two populations, providing diagnosable morphological evidence further supporting recognition of two taxa. The phylogenetic analysis included herein also recovered a monophyletic group for Oedopinola, itself containing a monophyletic group endemic to northern Honduras and made up of O. gephyra, O. tomasi McCranie from the Sierra de Omoa, and the heretofore un-named sister species to O. gephyra, which we describe below.

FIGURE 1. Map of Honduras showing the known localities for Oedipina petiola sp. nov. (square) and its closest relatives, O. gephyra (circle) and O. tomasi (star).

Methods

Morphology and measurements. The description of the holotype of the new species follows the format used in McCranie et al. (1993), except for some additional characters. All measurements were made in millimeters to the nearest tenth with dial calipers and the aid of a dissecting microscope. The abbreviation SVL stands for snout-vent- length (= standard length [SL] of some workers). Maxillary and vomerine tooth row counts are both sides summed. Comparative data for O. gephyra were taken by McCranie and published in McCranie & Wilson (2002) and McCranie et al. (1993), and that for O. tomasi is from McCranie (2006a). The color names and numeric codes used are those of Smithe (1975–1981). Comparative specimens examined are listed in Appendix I. DNA extraction, PCR amplification, and sequencing. Template DNA was extracted from muscle tissue using the QIAGEN PureGene DNA Isolation Kit following manufacturer’s instructions. Two mitochondrial gene

60 · Zootaxa 2990 © 2011 Magnolia Press MCCRANIE & TOWNSEND fragments were amplified using the primers MVZ15L and MVZ18H (Moritz et al. 1992) for cytochrome b (cyt b) and 16Sar-L and 16Sbr-H (Palumbi et al. 1991) for 16S large subunit RNA (16S). Unincorporated nucleotides were removed from PCR product using 1 uL of ExoSAP-IT (USB Corporation) per 10uL of product, which was then cycle sequenced using the BigDye Terminator 3.1 Cycle Sequencing kit for both forward and reverse strands, further purified using spin column filtration through Sephadex, and electrophoresed on an ABI 3130xl (Applied Biosystems, Inc.) at the University of Florida WEC/SFRC Molecular Ecology Lab.

TABLE 1. Samples utilized in the phylogenetic analysis, voucher numbers, locality information, and GenBank accession numbers. Taxon Locality Voucher 16S cyt b Oedipina petiola sp. nov. Honduras: Atlántida USNM 343462 AF199217 AF199161 Oedipina alleni Costa Rica: Puntarenas MVZ 190857 AF199207 AF199149 Oedipina carablanca Costa Rica: Limón No voucher FJ196862 FJ196869 Oedipina collaris Panamá: Coclé SIUC H 8896 FJ196863 FJ196870 Oedipina complex Panamá: Colón MVZ 236255 AF199213 AF199157 Oedipina cyclocauda Costa Rica: Heredia MVZ 138916 AF199214 AF199158 Oedipina elongata Honduras: Copán UTA A-56809 JN190932 - Honduras: Cortés UF 144649 - JN190938 Guatemala: Izabal UTA A-51906 AF199216 AF199160 Oedipina gephyra Honduras: Yoro USNM 530582 AF199218 AF199162 UF JHT2443 JN190930 JN190936 UF JHT2451 JN190931 JN190937 Oedipina gracilis Costa Rica: Heredia MVZ 203753 - AF199163 Oedipina grandis Costa Rica: Puntarenas MVZ 219593 - AF199165 Oedipina ignea Honduras: Ocotepeque USNM 530586 AF199231 AF199192 Oedipina kasios Honduras: Francisco Morazán UF 156500 HM113477 HM113484 Oedipina sp. Nicaragua: Atlántico Norte UF 156456 JN190926 JN190933 Oedipina leptopoda Honduras: Yoro MVZ 167772 - AF199193 Oedipina maritima Panamá: Bocas del Toro MVZ 219997 AF199221 AF199166 Oedipina nica Nicaragua: Jinotega UF 156445 HM113474 HM113482 Oedipina pacificensis Costa Rica: Puntarenas UCR 12063 AF199222 AF199169 Oedipina parvipes Panamá: San Blas MVZ 210404 AF199210 AF199154 Oedipina poelzi Costa Rica: Alajuela MVZ 206398 AF199223 AF199171 Oedipina pseudouniformis Costa Rica: Cartago MVZ 203749 AF199227 AF199178 Oedipina quadra Honduras: Gracias a Dios MVZ 232824 FJ196865 FJ196871 Oedipina savagei Costa Rica: Puntarenas UCR 14587 AF199209 AF199152 Oedipina stenopodia Guatemala: San Marcos MVZ 163649 AF199228 AF199181 Oedipina taylori Guatemala: Zacapa USCG 1134 HM068304 HM068302 Oedipina tomasi Honduras: Cortés UF 150066 JN190929 JN190935 MVZ 258037 HM068305 - Oedipina uniformis Costa Rica: Cartago MVZ 203751 AF199230 AF199190 Nototriton barbouri Honduras: Yoro UF 156538 GU971733 GU971734

Sequence alignment and analyses. A dataset containing all available sequences of Oedipina was generated from a combination of published data available from the NCBI (http://www.ncbi.nlm.nih.gov/) and newly generated sequence data (Table 1). Sequences were trimmed to 491 bp for 16S and 385 bp for cyt b to match the length of available sequences from multiple genera. Sequences were aligned using ClustalW (Thompson et al. 1994). Pairwise sequence divergence was estimated under the Kimura-2 parameter using the program package MEGAv5

NEW SPECIES OF WORM SALAMANDER FROM HONDURAS Zootaxa 2990 © 2011 Magnolia Press · 61 (Tamura et al. 2011); Kimura-2 parameter was selected for use to maintain consistency with previously published sequence divergence estimates for Oedipina (García-París & Wake 2000, McCranie et al. 2008). Bayesian and maximum likelihood (ML) phylogenetic analyses were performed on the 876 bp concatenated dataset. We partitioned the dataset by gene (16S and cyt b) and by codon position (1st, 2nd, 3rd) for cyt b, with best-fit models for each partition determined using the Akaike Information Criterion implemented in MrModeltest2.2 (Nylander 2004). The model GTR+I+ was selected for 16S; for cyt b GTR+Γ was selected for 1st position, K(80)+I+Γ for 2nd position, and HKY for 3rd position. Bayesian analysis was performed using MrBayes 3.1.2 (Huelsenbeck & Ronquist 2001), and consisted of two parallel runs of four Markov chains (three heated, one cold) run for 10 x 106 generations and sampled every 1000 generations, with a random starting tree and the first 20% of samples dis- carded as burnin. Cumulative and sliding window plots of split frequencies and correlation of split frequencies in first versus second runs were visualized using Geneious v5.4 (Drummond et al. 2011) to assess convergence around posterior quantities. The remaining 8,001 post-burnin trees from both runs were used to generate a 50% majority rule consensus tree. Maximum likelihood analysis was carried out in RaxML v7.2.6 (Stamatakis 2006), with 1000 bootstrap replicates run on the concatenated and partitioned dataset under the GTR-GAMMA substitu- tion model; the dataset was partitioned by gene (16S and cyt b) and by codon position for cyt b.

Systematics

Oedipina petiola sp. nov. Figures 2–3

Oedipina gephyra. McCranie 1996: 29, García-París & Wake 2000: 45 (part), McCranie & Wilson 2002: 155 (part), McCranie & Castañeda 2005: 7, McCranie 2006b: 7 (part), McCranie & Castañeda 2007: 116 (part), McCranie et al. 2008: 4, Sunyer et al. 2010: 31.

Holotype. USNM 343462, subadult male, from south slope of Cerro Búfalo, 15°39’N, 86°48’W, Parque Nacional Pico Bonito, Cordillera Nombre de Dios, 1580 m elev., Atlántida, Honduras, collected 18 February 1995, by James R. McCranie. Diagnosis. A moderate-sized (holotype is a subadult male with 42.6 mm SVL; Figure 2) member of Oedipina, subgenus Oedopinola. Distinguished from Oedipina (Oedopinola) gephyra and Oedipina (Oedopinola) tomasi by having narrower hind feet with less discrete digits (Figure 3; hind foot width / SVL 0.019 in O. petiola versus 0.024–0.028 in four adult males, 0.021–0.028 in ten adult females, and 0.028 in one subadult O. gephyra and 0.028 in one adult male O. tomasi; discrete digital groves in O. petiola versus distinct grooves in O. gephyra and O. tomasi; no transverse grooves at proximal edges of digits in O. petiola versus distinct grooves at proximal edges of digits in O. gephyra and O. tomasi). Oedipina petiola differs from the other described Honduran species of the subgenus Oedopinola (O. elongata) in having narrower hind feet (hind foot width / SVL 0.019 versus 0.032 in one male O. elongata), in having 34 maxillary teeth (maxillary teeth usually absent in O. elongata), and in lacking distinct pale markings (distinct pale markings present on top of head and anterior portion of body in O. elongata). The new species differs from all other Honduran species of Oedipina (O. ignea, O. kasios, O. leptopoda, O. quadra, O. stuarti, and O. taylori) by having 17 costal groves (19–21 in those six species). In addition, O. petiola differs in mtDNA from all other members of the genus Oedipina in which that data is known (Table 1), being 4.0–4.2% divergent for 16S and 6.6% divergent for cyt b from its closest known relative, O. gephyra (Table 2). Description of holotype. Subadult male with SVL of 42.6 mm. Snout truncate in dorsal aspect, bluntly rounded in profile; head width 3.8, SVL / head width 11.2, head width / SVL 0.089; head length 6.9 mm, SVL / head length 6.2; nostrils small, situated near tip of snout; distinct nasolabial groove extending from lower-posterior margin of each nostril to lip; labial protuberances small; canthus rostralis moderately arched; eyes not protuberant, not visible beyond margin of jaw when viewed from below; suborbital groove distinct; postorbital groove shallow, extending posteriorly from eye before turning sharply ventrally to connect with gular fold, another branch proceed- ing sharply ventrally just posterior to mandible, extending across throat anterior to gular fold; no dermal glands on head or body; mental gland only faintly indicated; 34 maxillary teeth, extending posteriorly to a point two-thirds distance through length of orbit; two enlarged premaxillary teeth, located posterior to lip and in line with maxillary teeth; 20 vomerine teeth, in long, single, arched series, extending beyond outer edge of internal nares; axillary-

62 · Zootaxa 2990 © 2011 Magnolia Press MCCRANIE & TOWNSEND groin length 26.0 mm; 17 costal grooves (18 trunk vertebrae inferred); tail incomplete, thick, broadly rounded at base, barely constricted at base; tail width 2.4 mm, SVL / tail width 17.8; tail depth 2.6 mm, SVL / tail depth 16.4; postilliac gland not visible; limbs short, 11 costal folds not covered when fore- and hind limbs adpressed along side of body; hind limb length 6.5 mm, SVL / hind limb length 6.6; hind foot width 0.8 mm, SVL / hind foot width 53.3, hind foot width / SVL 0.019; digits I and II and III and IV on forelimbs fused and without indentation in webbing between digits II and III; digits I and II fused on hind limbs, dorsal grooves discrete between digits II–IV and absent at proximal edges of all digits; relative length of digits I

FIGURE 2. Subadult male holotype of Oedipina petiola (USNM 343462; SVL = 42.6 mm) in life.

Color in life: dorsum of body and tail Jet Black (89), that of head and limbs slightly paler black; ventral surface of body slightly paler black than dorsum of body; subcaudal surface Jet Black. Color in alcohol: dorsum of head and body grayish black, that of tail black, that of limbs brownish gray; ventral surface of head pale brown with black mottling; gular fold pale brown; ventral surface of body grayish black, that of tail black; ventral surfaces of limbs brown; tiny white iridophores present on body, especially laterally. Habitat and distribution. The single known specimen was under a log during February 1995 in primary broadleaf cloud forest (Lower Montane Wet Forest formation of Holdridge 1967) at 1580 m elevation. Other salamanders found at the same time and locality were representatives of the Nototriton barbouri and Bolitoglossa porrasorum complexes. The single specimen of the Nototriton was also under a log and the Bolitoglossa were common in arboreal situations. The collection site was reached by climbing from the Quebrada de Oro at 950 m elevation along the side of a large landslide. The site is located about 50 m W of the edge of the landslide. A return trip to the type locality for three days during late May 1996 failed to find additional specimens of the Oedipina. Comment. One anonymous reviewer suggested that the foot morphology of the subadult holotype of the new species might differ from the foot morphology of O. gephyra because the former is known only from a subadult with a SVL of 42.6 mm. Therefore, we compared the foot morphology of O. petiola to a subadult (USNM 343458) of O. gephyra from that species’ type locality. That subadult O. gephyra (SVL 35.7 mm) has a hind foot width / SVL of 0.028 (at the upper level of that ratio for adult O. gephyra) and better developed digital grooves compared with the larger holotype of O. petiola. Etymology. The name petiola is from Latin petiolus, meaning diminutive foot, and refers to the narrow feet in the holotype. Phylogenetic relationships. Both Bayesian and ML analyses (Figure 4) support Oedipina petiola as the sister species to O. gephyra (bootstrap [bs] = 96, posterior probability [pp] = 1.0), forming a clade with their sister taxon O. tomasi (bs = 100, pp = 1.0). Both analyses also recovered clades corresponding to the subgenera Oedipina, Oeditriton, and Oedopinola (Figure 4), and support inclusion of O. petiola and the rest of the O. gephyra group in Oedopinola. Although only a single sample of O. petiola is available, intraspecific variation in its sister species is very low (0.0–0.2% for 16S, 0.0% for cyt b) compared with the interspecific divergence between O. petiola and O. gephyra (4.0–4.2% for 16S, 6.6% for cyt b). Based on sequence divergence (Table 2), O. petiola is well differenti- ated from the remaining species of the subgenus Oedopinola (5.9–10.1% for 16S, 16.1– 25.3% for cyt b).

NEW SPECIES OF WORM SALAMANDER FROM HONDURAS Zootaxa 2990 © 2011 Magnolia Press · 63 64 · Zootaxa 2990 © 2011 Magnolia Press MCCRANIE & TOWNSEND FIGURE 3. Comparison of foot morphology between the holotype of Oedipina petiola (USNM 343462; SVL = 42.6 mm) and subadult (USNM 343458; SVL = 35.7 mm) and adult (USNM 530585; SVL = 54.1 mm) O. gephyra; A) dorsal (left) and ventral (right) views of right hind foot of O. petiola, B) dorsal (left) and ventral (right) views of right hind foot of a subadult O. gephyra, C) dorsal (left) and ventral (right) views of right hind foot of an adult O. gephyra.

Discussion

The type locality of Oedipina (Oedopinola) petiola lies in the nuclear zone of Parque Nacional Pico Bonito. McCranie & Castañeda (2005) demonstrated that 19 species of amphibians and reptiles endemic to Honduras occurred in that park. Two of these 19 Honduran endemics remain known only from the environs of that park (Craugastor chrysozetetes and C. cruzi). McCranie et al. (2008) demonstrated that the population of Oedipina cyclocauda reported from this park by McCranie & Castañeda (2005) actually represented O. quadra, another species not known outside of Honduras. With the addition of O. petiola, 21 species of Honduran endemics are now known from Parque Nacional Pico Bonito, three of which are not known from any locality outside this park.

NEW SPECIES OF WORM SALAMANDER FROM HONDURAS Zootaxa 2990 © 2011 Magnolia Press · 65 FIGURE 4. Bayesian phylogram based on concatenated and partition 16S and cyt b dataset, showing phylogenetic hypothesis for the genus Oedipina and its three subgenera and relationships of O. petiola sp. nov. Maximum likelihood bootstrap values and posterior probabilities less than 60 and 0.6 are not shown.

Much of the nuclear zone of Parque Nacional Pico Bonito, including the type locality of Oedipina petiola, is still covered with primary broadleaf forest. The windward sides of this mountainous park are visible from the city of La Ceiba and many smaller nearby towns. The interest of providing potable water to the human residents of the area is the driving force behind the protection of these forests. However, the current countrywide trend of increased deforestation in Honduras is also occurring on the leeward sides of these mountains, including those of the nuclear zone of the park. The “authorities” said to be responsible for the continued protection of the forests inside this national park remain oblivious to this deforestation. The first author has made nine trips to the leeward side of the mountains inside the park in the vicinity of Quebrada de Oro, the last being during 2003. Each trip has revealed additional deforestation inside the nuclear zone on that side of the park. The type locality of O. petiola lies on this leeward side of the park above Quebrada de Oro. Although primary forest remains in the vicinity of the type locality, the continued deforestation occurring in the immediate area is a concern for the long time survival of the forests and fauna. Oedipina petiola is classified as Critically Endangered using the criteria developed by the IUCN (B2ab[iii]).

66 · Zootaxa 2990 © 2011 Magnolia Press MCCRANIE & TOWNSEND Brame (1968), in the first major revision of the genus Oedipina, recognized only 15 named species in the genus. With the description of O. petiola, that number now stands at 31, with most additional species being recognized in the last 15 years. Some of these additions represent species names resurrected from the synonymy of another nominal form (i.e. Good & Wake 1997; Brame’s concept of O. uniformis has been shown to include at least five named species, which was to be expected because of the extreme variation between foot morphology demonstrated by his drawings). However, several of these additions are a result of field workers discovering new populations representing undescribed species. The recent advances in molecular systematics have proven to be a powerful tool in the recognition of evolutionary species of Oedipina, despite the conserved external morphology found in most members of the genus. Molecular studies by García-París & Wake (2000) and McCranie et al. (2008) have identified three monophyletic groups within Oedipina. García-París & Wake (2000) resurrected the name Oedopinola Hilton as a subgenus for the new clade their study recovered, thus Oedipina Keferstein became the name for the other subgenus. The McCranie et al. (2008) study identified a third clade, for which they proposed the subgeneric name Oeditriton, That study also recovered a phylogeny that confirmed that the subgenera Oedipina and Oedopinola each formed a monophyletic group. The recovered phylogeny (Figure 4) also confirms that each of these three subgenera forms a monophyletic group. The genus Oedipina has a geographical distribution extending from Chiapas, Mexico, to northern Ecuador. Although Honduras is in the northern portion of this geographical distribution, it contains members of all three subgenera. In fact, the subgenus Oeditriton was thought to be endemic to Honduras until the recent discovery of a population of that subgenus in northern Nicaragua (Sunyer et al. 2010). The description of O. petiola represents the seventh species of Oedipina described since 2005 (the fifth from Honduras). Additional undescribed species of Oedipina are likely to be discovered in Honduras in the near future.

Acknowledgments

Leonel Archaga of Corporación Hondureña de Desarrollo Forestal (AFE-COHDEFOR), Tegucigalpa, issued the collecting and export permits (Dictamen DAPVS-47–95) under which the holotype of the new species was collected. Mario R. Espinal was helpful in acquiring these permits. Steve W. Gotte and James A. Poindexter (USNM) provided the loan of the new species and comparative specimens. Poindexter also took the photographs used in Fig- ure 3 and Gotte provided additional information for one specimen. The field team of Jason Butler, Lorraine Ketzler, John Slapcinsky, Nathaniel Stewart, and Larry Wilson assisted in collecting samples of O. gephyra in 2008; that fieldwork was supported by a grant from the Critical Ecosystem Partnership Fund and carried out under permits AFE-COHDEFOR Resolucion GG-MP-055-2006 and Dictamen DAPVS 0091-2006 issued by Carla Cárcamo de Martínez and Iris Acosta O. (Departamento de Areas Protegidas y Vida Silvestre); we thank Arie Sanders, José Mora-B., and Fredy Membreño from the Centro Zamorano de Biodiversidad for providing logistical support for JHT’s 2008 fieldwork. Ileana Luque-Montes kindly translated the abstract. Laboratory work was carried out in the University of Florida WEC/SFRC Molecular Ecology Lab, and we thank James D. Austin and Jason M. Butler for support in the lab.

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Appendix I. Comparative specimens examined.

Oedipina gephyra Honduras: Yoro—2.5 airline km NNE of La Fortuna, USNM 343458–61, 530582–85.

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