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We studied the species of amphibians from Serranía de Las Quinchas, in the mid Magdalena Valley region of Colombia, comparing the data from a recent fieldwork with museum records and literature. We present a list of 50 species of amphibians (Anura and Caudata). In our survey, we recorded 36 species, of which 6 represented new records. One of the most interesting record is that of the genus Oedipina, since this is the first time the genus is recorded for the Cordillera Oriental. The number of species is what would be expected in a humid tropical forest of the mid-Magdalena river valley, given the co-occurrence of amphibian faunas distributed in the sub-Andean, Caribbean, and Chocó biogeographic regions.

We describe a new species of worm salamander (Oedipina (oedopinola) fortunensis, holotype male in Seckenberg, Frankfurt am Main) from Fortuna, west- central Panama.&nbsp

Mental glands and their associated delivery behaviors during courtship are unique to the plethodontid salamanders. Because previous interpretations of the evolution of these features were conducted using older phylogenetic hypotheses, we reanalyzed these traits with newer courtship descriptions and contemporary phylogenetic methods. Using Bayesian ancestral state reconstruction methods that have been developed since the first phylogenetic analyses were conducted in the mid-1990s, we reconstructed mental gland and courtship behavior evolution on a Bayesian phylogeny of the nuclear gene Rag1. The most probable ancestral condition for plethodontids was resolved as presence of a mental gland. Loss of a mental gland occurred in each subfamily and was recovered as the most probable ancestral condition for the Spelerpinae. In contrast, parsimony reconstruction recovered the presence of a mental gland in the ancestor to Spelerpinae with multiple secondary losses. We hypothesize that that absence of a mental gland is possibly ancestral in some clades (i.e., Spelerpinae) and secondary in others (e.g., paedomorphic Eurycea). The most probable ancestral form of the mental gland is likely to be the large pad-type distributed extensively in Plethodontinae and Bolitoglossinae. Desmognathans have the most unique mental glands, occurring in an anterior protrusion or bifurcated form (in Desmognathus wrighti). Fan-shaped mental glands evolved independently in Eurycea and Oedipina. Small pads arose independently in Bolitoglossinae, Plethodontinae, and Spelerpinae. Head-rubbing behavior for mental gland delivery mode was recovered as the most probable and parsimonious ancestral state for the Plethodontidae, with independent losses of this behavior in Plethodontinae and Spelerpinae. Because head-rubbing was observed in outgroups, we hypothesize that head-rubbing behavior predated mental gland evolution. Pulling, snapping, slapping, and biting behaviors evolved independently in the Plethodontinae and Spelerpinae and are not homologous with head-rubbing. All hypotheses of mental gland and courtship evolution invoke homoplasy.

We describe a new species of Oedipina (subgenus Oedipina) from premontane elevations of three isolated mountains in northern Nicaragua. The new cryptic species differs in molecular characters from its closest relatives: Oedipina cyclocauda (an Atlantic lowland species with a distributional range from central Panama to extreme southeastern Nicaragua) and Oedipina pseudouniformis. We regard all Nicaraguan specimens previously referred to as O. pseudouniformis to be conspecific with the new species herein described and restrict O. pseudouniformis as a Costa Rican endemic species. We also record the fourth known locality (and southernmost) of the Nicaraguan endemic O. nica and discuss additional species of Oedipina that are likely to be found in Nicaragua as field research continues in the country.

We describe three new species of the plethodontid salamander genus Oedipina from Honduras. All three are relatively small to moderate sized, elongated and attenuate forms, which are differentiated from each other and from other members of the genus in limb and digital features, size, and body shape. Their distinctiveness is validated by phylogenetic analysis of mtDNA (cytochrome b and 16S) data, which shows each to be strongly differentiated. Furthermore, two of the three species are sister taxa and they comprise a third major clade in the genus, which we recognize as a new subgenus.

Sequences of two mitochondrial genes (385 base pairs of cytochrome b and approximately 520 base pairs of 16S DNA) were gathered for 26 taxa of the Middle American plethodontid salamander genera Nototriton and Oedipina and from three outgroup members of the tribe Bolitoglossini. Phylogenetic analyses of these data reveal well-supported cladistic structure and demonstrate the paraphyly of the moss salamanders of the genus Nototriton, which includes two well-defined clades. One clade, the sister taxon of Oedipina, corresponding to the Costa Rican and Honduran species of the picadoi, richardi, and barbouri groups, retains the name Nototriton. A new name is required for the second clade, the sister taxon of Oedipina plus Nototriton (sensu stricto). This clade, which we name Cryptotriton, is well supported morphologically and includes the species of the nasalis and adelos groups. A new species of Nototriton from Monteverde, Costa Rica, is described as Nototriton gamezi. Species of Oedipina fall into tw...

A new species of Oedipina (subgenus Oedopinola) is described from a cloud forest locality in northwestern Honduras. The new species appears most closely related to Oedopinola gephyra, a cloud forest species from north-central Honduras. It differs from O. gephyra in having a larger size, a jet-black coloration, a row of white flecks ventrolaterally on the body in life, and less interdigital webbing on the hind limbs.

Tail autotomy, an effective antipredator adaptation that enhances survival, may have attendant costs for the surviving prey. One potential cost not previously addressed in long-tailed, lungless salamanders that sacrifice the majority of their body mass by tail autotomy is the substantial reduction in respiratory surface area (RSA). We have tested the respiratory cost of tail autotomy in the bolitoglossine salamander, Oedipina uniformis, by (1) comparing resting and post-exercise oxygen consumption (VO2) in tailed and tailless Oedipina, and (2) comparing the tissue composition of bodies and tails. Metabolic studies indicated no significant effect of tail condition (pre- vs. post-autotomy) on resting VO2, the duration of exercise prior to exhaustion, or post-exercise VO2; aerobic recovery from exercise was also independent of tail condition. Tissue analyses (water and ash content, carbohydrate, lipid, and protein fractions) revealed that Oedipina bodies and tails are similar in composition (both high in muscle, low in lipid); thus the tail is largely a metabolically-homogeneous extension of the body. Our data suggest that tail autotomy in Oedipina does not alter the ratio between metabolically-active tissue mass and RSA, and that a respiratory cost of tail autotomy is of minor importance.

Oedipina nimaso sp. nov. Nimaso Worm Salamander Figure 6 Holotype. UCR 8391, a subadult male from Cerro Nimaso, Prov. Lim��n, Costa Rica, 1093 m, collected by D. Robinson, Federico Bola��os, and Gilbert Barrantes on April 14, 1984. Diagnosis. A small, extremely slender member of Oedipina (Oedopinola), based on having greater than 14 but fewer than 20 trunk vertebrae (Garc��a-Par��s & Wake 2000; McCranie et al. 2008), distinguished from other members of that clade by the combination of its small size, slender habitus, its long pointed snout and very narrow hands and feet with pointed digital tips. Distinguished from Costa Rican and Panamanian members of the clade as follows: from O. carablanca by smaller size, very narrow pes (vs. very broad and webbed in O. carablanca) and manus relative to SVL with reduced numbers of phalanges (0-1 - 2 - 1 manus vs. 1-2 - 3 - 2 in O. carablanca; 0-1 - 2 - 1 - 1 pes vs. 1-2 - 3 -(2,3)- 2 in O. carablanca) and little white dorsal pigment vs. extensive white pigment on head and body in O. carablanca; from O. parvipes and O. maritima by narrower and more syndactylous hands and feet, rounded snout and relatively large and numerous maxillary teeth (max 8 in O. maritima, fewer than 5 in Panamanian O. parvipes); from O. alleni in being much smaller and less robust with much narrower pes and shorter digits, and in having more maxillary teeth (20 vs. 5 or fewer in O. alleni); from O. savagei by being less robust and in having shorter limbs (limb interval 9.5 vs. less than 7 in O. savagei) and narrower pes (1.2 vs. 1.9 in O. savagei), and in lacking white pigment on the back of the head and a dorsal stripe on the trunk; from O. fortunensis by having shorter limbs (limb interval 9.5 vs. 8 in O. fortunensis), narrower pes (1.2 mm vs. 1.7 in O. fortunensis), and a shorter, more pointed head (SL/SG = 6.4 vs. 5.2 in O. fortunensis); and from O. complex by having a longer tail (SVL/TL less than 0.73 vs. 0.89 in O. complex), broader head (SVL/HW 8.6 vs. 10.2 in O. complex) and narrower pes with a long, pointed third toe (vs. short rounded toe in O. complex). Description. A diminutive, slender species compared to other Oedopinola. Sole specimen, the holotype, has a SL of 30.8 mm. Holotype very slender (Fig. 6) with narrow head (SL/HW 8.6) and rather long snout (SL/SG = 6.4) that is more rounded than pointed. Eyes small and only slightly protuberant. Limbs long and slender (SL/ HLL = 4.5) and the right hindlimb is missing. Manus and pes flat and very narrow (SL/FW = 25.7), with digits poorly defined and fused together. Longest digit with long, sharply pointed tip that is slightly bent in a preaxial direction. Relatively numerous maxillary teeth relatively large and single premaxillary tooth is long and hooked, suggesting that individual is near or at sexual maturity. Measurements (in mm), limb interval and tooth counts of the male holotype (Table 2). HW 3.6, SG 4.8, HD 1.6, EW 0.4, EL 1.1, ES 1.3, ED 0.7, IC 1.0, IO 1.2, length of groove extending posteriorly from eye 0.8, distance between nuchal groove and gular fold 1.4, SF 7.3, IN 1.0, external naris to snout 0.4, SP 0.3, SL 30.8, SAV 26.6, AX 19.8, LI 9.5, TL 42.0 (tail broken at tip, only slightly longer in life), tail width at base 2.1, tail depth at base 1.8, FLL 6.2, HLL 6.9, HAW 1.0, FW 1.2, T 5 0.8, T 3 1.1, parotoid width 1.4, parotoid length 2.9, nostril diameter 0.6. Number of teeth: PMT 1, MT 9 / 11, VT 6 / 6. Coloration of the holotype in alcohol. Brown to reddish-brown generally, with small amount of whitish pigment on head, mainly concentrated between eyes. Venter lighter than other surfaces and covered with tiny whitish patches or speckles. Habitat and range. Known only from the type locality. The locality is essentially mature forest within a sparsely inhabited indigenous reserve. The forest is very humid tropical forest in a transition zone to premontane forest. Etymology. The scientific name is a noun in apposition and refers to the type locality at Cerro Nimaso. Remarks. This specimen is badly desiccated. Osteological information has been derived from radiographs. There are 18 trunk vertebrae, one sacral, two caudosacral and 31 caudal vertebrae, with the very tip of the tail missing. The phalangeal formula for the manus is 0-1 - 2 - 1 and for the pes is 0-1 - 2 - 1 - 1. The basic formula for Oedipina is 1-2 - 3 - 2, and 1-2 - 3 - 3 - 2 (Wake 1966, Brame 1968, Garc��a-Par��s & Wake 2000) so there has been a considerable reduction in this species. Some other members of Oedopinola from Costa Rica and Panama also have relatively few phalanges. The last trunk vertebra that bears ribs is number 16. Mesopodial elements are unmineralized cartilage. A preorbital process is present on the vomers. Nasals are moderately protuberant and are the anterior-most skeletal bones. The skull bones are fully articulated and well-developed, suggesting a near adult state of development, so this is likely a miniaturized species compared to other members of its genus., Published as part of Boza-Oviedo, Eduardo, Rovito, Sean M., Chaves, Gerardo, Garc��a-Rodr��guez, Adri��n, Artavia, Luis G., Bola��os, Federico & Wake, David B., 2012, Salamanders from the eastern Cordillera de Talamanca, Costa Rica, with descriptions of five new species (Plethodontidae: Bolitoglossa, Nototriton, and Oedipina) and natural history notes from recent expeditions, pp. 36-61 in Zootaxa 3309 on pages 50-51, DOI: 10.5281/zenodo.211943, {"references":["Garcia-Paris, M. & Wake, D. B. (2000) Molecular phylogenetic analysis of relationships of the tropical salamander genera Oedipina and Nototriton, with descriptions of a new genus and three new species. Copeia, 2000, 42 - 70.","McCranie, J. R., Vieites, D. R., & Wake, D. B. (2008) Description of a new divergent lineage and three new species of Honduran salamanders of the genus Oedipina (Caudata, Plethodontidae). Zootaxa, 1930, 1 - 17.","Wake, D. B. (1966) Comparative osteology and evolution of the lungless salamanders, Family Plethodontidae. Memoirs of the Southern California Academy of Sciences, 9, 1 - 111.","Brame, A. H. Jr. (1968) Systematics and evolution of the Mesoamerican salamander genus Oedipina. Journal of Herpetology, 2, 1 - 64."]}

Oedipina nimaso sp. nov. Nimaso Worm Salamander Figure 6 Holotype. UCR 8391, a subadult male from Cerro Nimaso, Prov. Limón, Costa Rica, 1093 m, collected by D. Robinson, Federico Bolaños, and Gilbert Barrantes on April 14, 1984. Diagnosis. A small, extremely slender member of Oedipina (Oedopinola), based on having greater than 14 but fewer than 20 trunk vertebrae (García-París & Wake 2000; McCranie et al. 2008), distinguished from other members of that clade by the combination of its small size, slender habitus, its long pointed snout and very narrow hands and feet with pointed digital tips. Distinguished from Costa Rican and Panamanian members of the clade as follows: from O. carablanca by smaller size, very narrow pes (vs. very broad and webbed in O. carablanca) and manus relative to SVL with reduced numbers of phalanges (0-1 - 2 - 1 manus vs. 1-2 - 3 - 2 in O. carablanca; 0-1 - 2 - 1 - 1 pes vs. 1-2 - 3 -(2,3)- 2 in O. carablanca) and little white dorsal pigment vs. extensive white pigment on head and body in O. carablanca; from O. parvipes and O. maritima by narrower and more syndactylous hands and feet, rounded snout and relatively large and numerous maxillary teeth (max 8 in O. maritima, fewer than 5 in Panamanian O. parvipes); from O. alleni in being much smaller and less robust with much narrower pes and shorter digits, and in having more maxillary teeth (20 vs. 5 or fewer in O. alleni); from O. savagei by being less robust and in having shorter limbs (limb interval 9.5 vs. less than 7 in O. savagei) and narrower pes (1.2 vs. 1.9 in O. savagei), and in lacking white pigment on the back of the head and a dorsal stripe on the trunk; from O. fortunensis by having shorter limbs (limb interval 9.5 vs. 8 in O. fortunensis), narrower pes (1.2 mm vs. 1.7 in O. fortunensis), and a shorter, more pointed head (SL/SG = 6.4 vs. 5.2 in O. fortunensis); and from O. complex by having a longer tail (SVL/TL less than 0.73 vs. 0.89 in O. complex), broader head (SVL/HW 8.6 vs. 10.2 in O. complex) and narrower pes with a long, pointed third toe (vs. short rounded toe in O. complex). Description. A diminutive, slender species compared to other Oedopinola. Sole specimen, the holotype, has a SL of 30.8 mm. Holotype very slender (Fig. 6) with narrow head (SL/HW 8.6) and rather long snout (SL/SG = 6.4) that is more rounded than pointed. Eyes small and only slightly protuberant. Limbs long and slender (SL/ HLL = 4.5) and the right hindlimb is missing. Manus and pes flat and very narrow (SL/FW = 25.7), with digits poorly defined and fused together. Longest digit with long, sharply pointed tip that is slightly bent in a preaxial direction. Relatively numerous maxillary teeth relatively large and single premaxillary tooth is long and hooked, suggesting that individual is near or at sexual maturity. Measurements (in mm), limb interval and tooth counts of the male holotype (Table 2). HW 3.6, SG 4.8, HD 1.6, EW 0.4, EL 1.1, ES 1.3, ED 0.7, IC 1.0, IO 1.2, length of groove extending posteriorly from eye 0.8, distance between nuchal groove and gular fold 1.4, SF 7.3, IN 1.0, external naris to snout 0.4, SP 0.3, SL 30.8, SAV 26.6, AX 19.8, LI 9.5, TL 42.0 (tail broken at tip, only slightly longer in life), tail width at base 2.1, tail depth at base 1.8, FLL 6.2, HLL 6.9, HAW 1.0, FW 1.2, T 5 0.8, T 3 1.1, parotoid width 1.4, parotoid length 2.9, nostril diameter 0.6. Number of teeth: PMT 1, MT 9 / 11, VT 6 / 6. Coloration of the holotype in alcohol. Brown to reddish-brown generally, with small amount of whitish pigment on head, mainly concentrated between eyes. Venter lighter than other surfaces and covered with tiny whitish patches or speckles. Habitat and range. Known only from the type locality. The locality is essentially mature forest within a sparsely inhabited indigenous reserve. The forest is very humid tropical forest in a transition zone to premontane forest. Etymology. The scientific name is a noun in apposition and refers to the type locality at Cerro Nimaso. Remarks. This specimen is badly desiccated. Osteological information has been derived from radiographs. There are 18 trunk vertebrae, one sacral, two caudosacral and 31 caudal vertebrae, with the very tip of the tail missing. The phalangeal formula for the manus is 0-1 - 2 - 1 and for the pes is 0-1 - 2 - 1 - 1. The basic formula for Oedipina is 1-2 - 3 - 2, and 1-2 - 3 - 3 - 2 (Wake 1966, Brame 1968, García-París & Wake 2000) so there has been a considerable reduction in this species. Some other members of Oedopinola from Costa Rica and Panama also have relatively few phalanges. The last trunk vertebra that bears ribs is number 16. Mesopodial elements are unmineralized cartilage. A preorbital process is present on the vomers. Nasals are moderately protuberant and are the anterior-most skeletal bones. The skull bones are fully articulated and well-developed, suggesting a near adult state of development, so this is likely a miniaturized species compared to other members of its genus.

Oedipina nica sp. nov. Nicaraguan Highland Worm Salamander Figure 1 Oedipina cyclocauda (in part). K��hler 1999, 2001. Holotype. MVZ 263774, an adult male, from El Gobiado, Reserva Natural Cerro Datanl��-El Diablo, 13 ��09���N, 85 �� 52 ���W, 1420 m above sea level (a.s.l.), Dept. Jinotega, Nicaragua, collected 30 August 2009 by S. M. Rovito, T. J. Papenfuss, J. Sunyer, and L. A. Obando (original field number SMR 842). Paratypes (5). UF 156445, from Reserva Natural Cerro Kilamb��, 13 �� 34 ���N, 85 �� 42 ���W, 1625 m a.s.l., Dept. Jinotega, Nicaragua; UF 156446 ��� 47, from Reserva Natural Cerro Kilamb��, 13 �� 35 ���N, 85 �� 43 ���W, 1660 m a.s.l., Dept. Jinotega, Nicaragua; UF 156453 ��� 54, from Reserva Natural Macizos de Pe��as Blancas, 13 �� 17 ���N, 85 �� 43 ���W, 1515 m a.s.l., Dept. Jinotega, Nicaragua. Referred specimens (12). SMF 78736, from Camp El Hielo, Reserva Natural Cerro Kilamb��, 1490 m a.s.l., Dept. Jinotega, Nicaragua; SMF 78737, 78739��� 40, from Camp 2, Reserva Natural Cerro Kilamb��, 13 �� 35.25 'N, 85 �� 41.50 'W, 1360 m a.s.l., Dept. Jinotega, Nicaragua; UF 156443 ��� 44, from Reserva Natural Cerro Kilamb��, 13 �� 34 ���N, 85 �� 42 ���W, 1625 m a.s.l., Dept. Jinotega, Nicaragua; UF 156448 ��� 50, from Reserva Natural Cerro Kilamb��, 13 �� 35 ���N, 85 �� 43 ���W, 1660 m a.s.l., Dept. Jinotega, Nicaragua; UF 156451 ��� 52, two tails dropped by escaped adults from Reserva Natural Cerro Kilamb��, 13 �� 34 ���N, 85 �� 42 ���W, 1600 m a.s.l., Dept. Jinotega, Nicaragua; UF 156455 from Reserva Natural Macizos de Pe��as Blancas, 13 �� 17 ���N, 85 �� 43 ���W, 1515 m a.s.l., Dept. Jinotega, Nicaragua. Diagnosis. A slender, small-sized (largest specimen 48.5 mm SL) species assigned to the genus Oedipina based on the presence of a high number (19���20) of costal grooves between the short limbs, very narrow manus and pes, long tail (tail length ca 2 x SL), overall morphological similarity to other Oedipina species, and molecular phylogenetic evidence. This species is a member of the subgenus Oeditriton (based on molecular data; Fig. 2) and is distinguished from the only other members of that clade (O. kasios and O. quadra) by its more slender habitus and fewer vomerine teeth. Further distinguished from other Nicaraguan members of the genus Oedipina as follows: from O. collaris by being much smaller and more slender, having short limbs and narrow manus and pes, and having a short, rather bluntly tipped snout; from O. cyclocauda by being smaller and more slender without nearly pad-like feet; from O. pseudouniformis by being more slender with shorter, less robust limbs and narrower manus and pes, and by having almost uniformly dark brownishblack coloration with no distinct light pigmentation on the upper parts of the proximal segment of the limbs. Description of the holotype. The male holotype is 39.2 mm SL and is judged to be an adult by the presence of small papillae in the anterior margins of the vent and a small mental gland behind the anterior tip of the mandible. The small head is rounded in dorsal profile, and somewhat flattened in lateral profile with a relatively blunt snout and distinct nasolabial protuberances. Snout to posterior angle of vent 9.8 times head width and 6.5 times head length. Nostrils are small but conspicuous under magnification. The small eyes are inconspicuous and do not extend beyond the lateral margins of the head. The suborbital groove does not intercept the lip line. The skin of the dorsal surface of the head behind the eyes is depressed in a pattern that suggests the presence of a frontoparietal fenestra. There is a single slightly enlarged premaxillary tooth that is located distinctly anterior to the maxillary teeth. Maxillary teeth total 41. The small vomerine teeth are in a short, fairly straight series and total 11. There are 20 costal grooves between the small limbs with a limb interval of 13 intercostal folds. Hands and feet are tiny, narrow, and elongated. Digit I is fused with digit II and digit IV is fused with digit III on the forelimbs, with only a part of a phalangeal segment of digit III free between digits II���III on the forelimbs. Digit I is fused with digit II and digit V is fused with digit IV on the hind limbs; both outer digits are extremely short on the hind limbs. The triangular tip of digit III of the hind limb is distinct from the tiny rounded tips of digits II and IV, but less than a full phalangeal segment of that digit is free of the others. Subdigital pads are nearly imperceptible. Digits on forelimbs in order of decreasing length are III���II���IV���I; digits on hind limbs are III���II���IV���(V���I). The tail is round in cross section and tapers gradually to a narrow tip; the tail is almost twice SL. Measurements (in millimeters) of the holotype. Head width 4.0; snout to gular fold (head length) 6.0; head depth at posterior angle of jaw 1.8; eyelid width 1.1; eyelid length 1.5; eye to nostril 0.6; anterior rim of eye to snout 1.2; horizontal orbital diameter 1.0; interorbital distance 1.7; distance separating eyelids 0.7; nostril diameter 0.2; snout projecting beyond mandible 0.4; distance from eye to distal end of postorbital groove 1.9; snout to posterior angle of vent (SL) 39.2; snout to anterior angle of vent 37.0; snout to forelimb 9.4; axilla to groin 26.0; limb interval 13; shoulder width 2.7; tail length 77.0; tail width at base 2.4; tail depth at base 2.4; forelimb length (to tip of longest digit) 3.7; hind limb length (to tip of longest digit) 4.5; forelimb foot width 0.6; hind limb foot width 0.9; free length of longest digit 0.2. Coloration of the holotype in alcohol. Dorsal and lateral surfaces of the head, body, and tail are black. Costal grooves on the body and tail are the same color as the body or slightly lighter dark gray, but at the deepest part of the groove pigment is lacking so the grooves are prominent. Ventral surfaces of the head, body, and tail are slightly paler black than those dorsal surfaces. Dorsal and ventral surfaces of the limbs are the same as for the body. Tiny white speckles are most visible behind the eye and over the shoulder, then in an irregular, narrow line along the dorsolateral part of the body to the hind limbs, but they are few in number and obscure. White spots are a little larger in the gular and chest areas ventrally. The nasolabial protuberances are unpigmented and contrast with the surrounding tissue. Variation. Variation based on study of the holotype and three male (UF 156445, 156447, 156453), one female (UF 156446), and one juvenile (UF 156454) paratypes: the four males, judged to be mature based on having inconspicuous mental glands and anterior vent margins bearing fine pinnae, range from 38.5���44.5 SL; the female is 48.5 SL. Tails are present in the holotype, three male paratypes, and the juvenile and all taper gradually to slender tips. SL is from 0.86 (adult) to 0.52 (holotype) times tail length. SL in males is 9.8���11.8 times head width and 9.9 in the single female. Limbs are relatively short; number of costal folds between adpressed limbs is 13���14 in males and 13 in the female. There are 19���20 costal grooves between the limbs in the males and 20 in the female. Feet are narrow; 0.8���1.1. The digits are joined to their neighbors, with only the tips of the longest toes free. Males have 1���2 premaxillary teeth, the female has 4. Maxillary teeth range from 41���47 in males; 48 in the female. Males have 9���16 vomerine teeth; the female 18. If one adds the referred specimens, the size range of the adult males increases to 35.3���48.5 SL, and that of females from 37.5���48.5 SL. These specimens are consistent in measurements, tooth and costal groove counts, and coloration with the type series. SMF 78738, from Reserva Natural Cerro Kilamb�� (44.0 SL), fits within the range of the new species in all measurements (see discussion). Variation of coloration based on color photographs taken in life during daytime. Coloration is in general uniformly dark brownish-black with no distinguishing features. The upper side of the proximal segment of the limbs is the same color. The costal grooves are very evident because of the lack of pigment at their centers. Details for particular individuals follow. Adult male paratype (UF 156447; Fig. 1 b): Dorsum of body and tail blackish brown becoming a faded lighter brown on the dorsal and lateral surfaces of head extending to the chin. A lighter brown speckling is faintly noticeable dorsally from the head becoming imperceptible towards the base of the tail. Ventral surfaces of the body are blackish brown, similar in coloration to the tail. Limbs faded brown similar to color of head. Skin creases such as costal grooves, gular fold, and those around the paratoid region lack pigmentation and appear light grey. Iris dark rust brown. Adult male (UF 156450): Dorsum of body and tail blackish brown becoming faded lighter brown on the anterior and lateral surfaces of head. Gold speckling mixed with some white is located on the dorsal surface of the head and body. Speckling is most prominent on the top of the head becoming sparser posteriad, not extending beyond base of tail. Dorsolaterally above the anterior limbs a slight concentration of tiny white speckles forms a short, irregular line. Below the extent of the speckling ventrolaterally coloration is same as that of tail. Limb coloration is same as that of anterior and lateral surface of head. Iris dark rust brown. Juvenile paratype (UF 156454; Fig. 1 d): Dorsal and lateral surfaces of body, head, tail, and limbs are a uniform jet black. There is a faint overlay of tiny dirty white flecks along the dorsum of head, body, and tail giving a somewhat ashy appearance. This flecking is most concentrated dorsolaterally giving the impression of an irregular line extending from behind the jaw to the hind limb. Flecking is also present around the mouth and chest area. Larger white speckles are mixed sparsely and intermittently along the head, body, tail, and limbs. Habitat and distribution. This species is known to occur from 1360���1660 m elevation at three isolated localities in the Lower Montane Moist Forest formation (Holdridge 1967) in north-central Nicaragua (Fig. 3). The holotype was collected in a small patch of secondary cloud forest (Fig. 4 a���b) mostly surrounded by cattle pastures and agricultural land. It was found while raking during the daytime inside a moderately large rotten log. The paratypes and UF referred specimens from Reserva Natural Cerro Kilamb�� were collected in undisturbed, primary cloud forest (Fig. 4 c) during both diurnal and nocturnal surveys. All were found underneath logs, rocks, and other debris in areas along the mountain ridge where the substrate comprised a spongy mass of plant roots and mosses. The SMF referred specimens from Reserva Natural Cerro Kilamb�� were also collected in undisturbed, primary cloud forest during diurnal surveys. Most of them were found inside dead basal ramifications of giant ferns. The paratypes from Reserva Natural Macizos de Pe��as Blancas were collected in undisturbed, primary cloud forest (Fig. 4 d). One was uncovered during the daytime beneath a fallen log, and two others were found active on the forest floor around the campsite at 2130 h. Those found active within the camp were likely dislodged or disturbed by raking and uprooting logs that occurred during set-up of the campsite earlier the same day. Phylogenetic relationships. Phylogenetic analysis of sequence data from two mitochondrial genes (16 S and cyt b) confirms the monophyly of this taxon (ML bootstrap support = 99, Bayesian posterior probability = 1.0) and its position as sister species to Oedipina kasios in the Oeditriton clade (Fig. 2). A sister-species relationship between O. kasio s and O. nica is strongly supported (ML bootstrap support = 93, Bayesian posterior probability = 0.99) as is the monophyly of an Oeditriton clade containing O. kasio s, O. nica, and O. quadra (ML bootstrap support = 96, Bayesian posterior probability = 1.0). Oeditriton is the sister clade to the remaining Oedipina, which are divided into the subgenera Oedipina and Oedopinola (Fig. 2), which supports previous phylogenetic hypotheses for Oedipina (Garc��a-Par��s & Wake 2000; McCranie et al. 2008). All wellsupported phylogenetic relationships are congruent between the ML and Bayesian results. Etymology. The specific name nica is a short name for Nicaraguan (Nicarag��ense in Spanish) and is a colloquial word used in Central America to denote Nicaraguan people. The name alludes to the country of origin of the type specimens. Conservation status. Using the criteria established by IUCN for evaluating threatened species, Oedipina nica should be classified as Endangered (EN B 2 ab[iii]), due to its limited distribution (known only from three isolated highland forest areas with a total extent of less than 500 km 2) and the continued loss of habitat at these localities., Published as part of Sunyer, Javier, Wake, David B., Townsend, Josiah H., Travers, Scott L., Rovito, Sean M., Papenfuss, Theodore J., Obando, Lenin A. & K��hler, Gunther, 2010, A new species of worm salamander (Caudata: Plethodontidae: Oedipina) in the subgenus Oeditriton from the highlands of northern Nicaragua, pp. 29-39 in Zootaxa 2613 on pages 32-35, DOI: 10.5281/zenodo.197858, {"references":["Kohler, G. (1999) The amphibians and reptiles of Nicaragua - a distributional checklist with keys. Courier Forschungsinstut Senckenberg, 213, 1 - 212.","Kohler, G. (2001) Anfibios y Reptiles de Nicaragua. Herpeton. Offenbach, Germany.","Holdridge, L. R. (1967) Life Zone Ecology. Tropical Science Center. San Jose, Costa Rica.","McCranie, J. R., Vieites, D. R. & Wake, D. B. (2008) Description of a new divergent lineage and three new species of Honduran salamanders of the genus Oedipina (Caudata, Plethodontidae). Zootaxa, 1930, 1 - 17."]}

We describe a new species of Oedipina (subgenus Oeditriton) from the highlands of north-central Nicaragua. The new species is most closely related to Oedipina kasios, a premontane species from north-central Honduras, and O. quadra, a lowland species from northern and eastern Honduras. It differs from O. kasios and O. quadra in molecular characters and by having a more slender habitus and fewer vomerine teeth. We also discuss the status of the additional populations of Oedipina from Nicaragua.

Oedipina nica sp. nov. Nicaraguan Highland Worm Salamander Figure 1 Oedipina cyclocauda (in part). Köhler 1999, 2001. Holotype. MVZ 263774, an adult male, from El Gobiado, Reserva Natural Cerro Datanlí-El Diablo, 13 °09’N, 85 ° 52 ’W, 1420 m above sea level (a.s.l.), Dept. Jinotega, Nicaragua, collected 30 August 2009 by S. M. Rovito, T. J. Papenfuss, J. Sunyer, and L. A. Obando (original field number SMR 842). Paratypes (5). UF 156445, from Reserva Natural Cerro Kilambé, 13 ° 34 ’N, 85 ° 42 ’W, 1625 m a.s.l., Dept. Jinotega, Nicaragua; UF 156446 – 47, from Reserva Natural Cerro Kilambé, 13 ° 35 ’N, 85 ° 43 ’W, 1660 m a.s.l., Dept. Jinotega, Nicaragua; UF 156453 – 54, from Reserva Natural Macizos de Peñas Blancas, 13 ° 17 ’N, 85 ° 43 ’W, 1515 m a.s.l., Dept. Jinotega, Nicaragua. Referred specimens (12). SMF 78736, from Camp El Hielo, Reserva Natural Cerro Kilambé, 1490 m a.s.l., Dept. Jinotega, Nicaragua; SMF 78737, 78739– 40, from Camp 2, Reserva Natural Cerro Kilambé, 13 ° 35.25 'N, 85 ° 41.50 'W, 1360 m a.s.l., Dept. Jinotega, Nicaragua; UF 156443 – 44, from Reserva Natural Cerro Kilambé, 13 ° 34 ’N, 85 ° 42 ’W, 1625 m a.s.l., Dept. Jinotega, Nicaragua; UF 156448 – 50, from Reserva Natural Cerro Kilambé, 13 ° 35 ’N, 85 ° 43 ’W, 1660 m a.s.l., Dept. Jinotega, Nicaragua; UF 156451 – 52, two tails dropped by escaped adults from Reserva Natural Cerro Kilambé, 13 ° 34 ’N, 85 ° 42 ’W, 1600 m a.s.l., Dept. Jinotega, Nicaragua; UF 156455 from Reserva Natural Macizos de Peñas Blancas, 13 ° 17 ’N, 85 ° 43 ’W, 1515 m a.s.l., Dept. Jinotega, Nicaragua. Diagnosis. A slender, small-sized (largest specimen 48.5 mm SL) species assigned to the genus Oedipina based on the presence of a high number (19–20) of costal grooves between the short limbs, very narrow manus and pes, long tail (tail length ca 2 x SL), overall morphological similarity to other Oedipina species, and molecular phylogenetic evidence. This species is a member of the subgenus Oeditriton (based on molecular data; Fig. 2) and is distinguished from the only other members of that clade (O. kasios and O. quadra) by its more slender habitus and fewer vomerine teeth. Further distinguished from other Nicaraguan members of the genus Oedipina as follows: from O. collaris by being much smaller and more slender, having short limbs and narrow manus and pes, and having a short, rather bluntly tipped snout; from O. cyclocauda by being smaller and more slender without nearly pad-like feet; from O. pseudouniformis by being more slender with shorter, less robust limbs and narrower manus and pes, and by having almost uniformly dark brownishblack coloration with no distinct light pigmentation on the upper parts of the proximal segment of the limbs. Description of the holotype. The male holotype is 39.2 mm SL and is judged to be an adult by the presence of small papillae in the anterior margins of the vent and a small mental gland behind the anterior tip of the mandible. The small head is rounded in dorsal profile, and somewhat flattened in lateral profile with a relatively blunt snout and distinct nasolabial protuberances. Snout to posterior angle of vent 9.8 times head width and 6.5 times head length. Nostrils are small but conspicuous under magnification. The small eyes are inconspicuous and do not extend beyond the lateral margins of the head. The suborbital groove does not intercept the lip line. The skin of the dorsal surface of the head behind the eyes is depressed in a pattern that suggests the presence of a frontoparietal fenestra. There is a single slightly enlarged premaxillary tooth that is located distinctly anterior to the maxillary teeth. Maxillary teeth total 41. The small vomerine teeth are in a short, fairly straight series and total 11. There are 20 costal grooves between the small limbs with a limb interval of 13 intercostal folds. Hands and feet are tiny, narrow, and elongated. Digit I is fused with digit II and digit IV is fused with digit III on the forelimbs, with only a part of a phalangeal segment of digit III free between digits II–III on the forelimbs. Digit I is fused with digit II and digit V is fused with digit IV on the hind limbs; both outer digits are extremely short on the hind limbs. The triangular tip of digit III of the hind limb is distinct from the tiny rounded tips of digits II and IV, but less than a full phalangeal segment of that digit is free of the others. Subdigital pads are nearly imperceptible. Digits on forelimbs in order of decreasing length are III–II–IV–I; digits on hind limbs are III–II–IV–(V–I). The tail is round in cross section and tapers gradually to a narrow tip; the tail is almost twice SL. Measurements (in millimeters) of the holotype. Head width 4.0; snout to gular fold (head length) 6.0; head depth at posterior angle of jaw 1.8; eyelid width 1.1; eyelid length 1.5; eye to nostril 0.6; anterior rim of eye to snout 1.2; horizontal orbital diameter 1.0; interorbital distance 1.7; distance separating eyelids 0.7; nostril diameter 0.2; snout projecting beyond mandible 0.4; distance from eye to distal end of postorbital groove 1.9; snout to posterior angle of vent (SL) 39.2; snout to anterior angle of vent 37.0; snout to forelimb 9.4; axilla to groin 26.0; limb interval 13; shoulder width 2.7; tail length 77.0; tail width at base 2.4; tail depth at base 2.4; forelimb length (to tip of longest digit) 3.7; hind limb length (to tip of longest digit) 4.5; forelimb foot width 0.6; hind limb foot width 0.9; free length of longest digit 0.2. Coloration of the holotype in alcohol. Dorsal and lateral surfaces of the head, body, and tail are black. Costal grooves on the body and tail are the same color as the body or slightly lighter dark gray, but at the deepest part of the groove pigment is lacking so the grooves are prominent. Ventral surfaces of the head, body, and tail are slightly paler black than those dorsal surfaces. Dorsal and ventral surfaces of the limbs are the same as for the body. Tiny white speckles are most visible behind the eye and over the shoulder, then in an irregular, narrow line along the dorsolateral part of the body to the hind limbs, but they are few in number and obscure. White spots are a little larger in the gular and chest areas ventrally. The nasolabial protuberances are unpigmented and contrast with the surrounding tissue. Variation. Variation based on study of the holotype and three male (UF 156445, 156447, 156453), one female (UF 156446), and one juvenile (UF 156454) paratypes: the four males, judged to be mature based on having inconspicuous mental glands and anterior vent margins bearing fine pinnae, range from 38.5–44.5 SL; the female is 48.5 SL. Tails are present in the holotype, three male paratypes, and the juvenile and all taper gradually to slender tips. SL is from 0.86 (adult) to 0.52 (holotype) times tail length. SL in males is 9.8–11.8 times head width and 9.9 in the single female. Limbs are relatively short; number of costal folds between adpressed limbs is 13–14 in males and 13 in the female. There are 19–20 costal grooves between the limbs in the males and 20 in the female. Feet are narrow; 0.8–1.1. The digits are joined to their neighbors, with only the tips of the longest toes free. Males have 1–2 premaxillary teeth, the female has 4. Maxillary teeth range from 41–47 in males; 48 in the female. Males have 9–16 vomerine teeth; the female 18. If one adds the referred specimens, the size range of the adult males increases to 35.3–48.5 SL, and that of females from 37.5–48.5 SL. These specimens are consistent in measurements, tooth and costal groove counts, and coloration with the type series. SMF 78738, from Reserva Natural Cerro Kilambé (44.0 SL), fits within the range of the new species in all measurements (see discussion). Variation of coloration based on color photographs taken in life during daytime. Coloration is in general uniformly dark brownish-black with no distinguishing features. The upper side of the proximal segment of the limbs is the same color. The costal grooves are very evident because of the lack of pigment at their centers. Details for particular individuals follow. Adult male paratype (UF 156447; Fig. 1 b): Dorsum of body and tail blackish brown becoming a faded lighter brown on the dorsal and lateral surfaces of head extending to the chin. A lighter brown speckling is faintly noticeable dorsally from the head becoming imperceptible towards the base of the tail. Ventral surfaces of the body are blackish brown, similar in coloration to the tail. Limbs faded brown similar to color of head. Skin creases such as costal grooves, gular fold, and those around the paratoid region lack pigmentation and appear light grey. Iris dark rust brown. Adult male (UF 156450): Dorsum of body and tail blackish brown becoming faded lighter brown on the anterior and lateral surfaces of head. Gold speckling mixed with some white is located on the dorsal surface of the head and body. Speckling is most prominent on the top of the head becoming sparser posteriad, not extending beyond base of tail. Dorsolaterally above the anterior limbs a slight concentration of tiny white speckles forms a short, irregular line. Below the extent of the speckling ventrolaterally coloration is same as that of tail. Limb coloration is same as that of anterior and lateral surface of head. Iris dark rust brown. Juvenile paratype (UF 156454; Fig. 1 d): Dorsal and lateral surfaces of body, head, tail, and limbs are a uniform jet black. There is a faint overlay of tiny dirty white flecks along the dorsum of head, body, and tail giving a somewhat ashy appearance. This flecking is most concentrated dorsolaterally giving the impression of an irregular line extending from behind the jaw to the hind limb. Flecking is also present around the mouth and chest area. Larger white speckles are mixed sparsely and intermittently along the head, body, tail, and limbs. Habitat and distribution. This species is known to occur from 1360–1660 m elevation at three isolated localities in the Lower Montane Moist Forest formation (Holdridge 1967) in north-central Nicaragua (Fig. 3). The holotype was collected in a small patch of secondary cloud forest (Fig. 4 a–b) mostly surrounded by cattle pastures and agricultural land. It was found while raking during the daytime inside a moderately large rotten log. The paratypes and UF referred specimens from Reserva Natural Cerro Kilambé were collected in undisturbed, primary cloud forest (Fig. 4 c) during both diurnal and nocturnal surveys. All were found underneath logs, rocks, and other debris in areas along the mountain ridge where the substrate comprised a spongy mass of plant roots and mosses. The SMF referred specimens from Reserva Natural Cerro Kilambé were also collected in undisturbed, primary cloud forest during diurnal surveys. Most of them were found inside dead basal ramifications of giant ferns. The paratypes from Reserva Natural Macizos de Peñas Blancas were collected in undisturbed, primary cloud forest (Fig. 4 d). One was uncovered during the daytime beneath a fallen log, and two others were found active on the forest floor around the campsite at 2130 h. Those found active within the camp were likely dislodged or disturbed by raking and uprooting logs that occurred during set-up of the campsite earlier the same day. Phylogenetic relationships. Phylogenetic analysis of sequence data from two mitochondrial genes (16 S and cyt b) confirms the monophyly of this taxon (ML bootstrap support = 99, Bayesian posterior probability = 1.0) and its position as sister species to Oedipina kasios in the Oeditriton clade (Fig. 2). A sister-species relationship between O. kasio s and O. nica is strongly supported (ML bootstrap support = 93, Bayesian posterior probability = 0.99) as is the monophyly of an Oeditriton clade containing O. kasio s, O. nica, and O. quadra (ML bootstrap support = 96, Bayesian posterior probability = 1.0). Oeditriton is the sister clade to the remaining Oedipina, which are divided into the subgenera Oedipina and Oedopinola (Fig. 2), which supports previous phylogenetic hypotheses for Oedipina (García-París & Wake 2000; McCranie et al. 2008). All wellsupported phylogenetic relationships are congruent between the ML and Bayesian results. Etymology. The specific name nica is a short name for Nicaraguan (Nicaragüense in Spanish) and is a colloquial word used in Central America to denote Nicaraguan people. The name alludes to the country of origin of the type specimens. Conservation status. Using the criteria established by IUCN for evaluating threatened species, Oedipina nica should be classified as Endangered (EN B 2 ab[iii]), due to its limited distribution (known only from three isolated highland forest areas with a total extent of less than 500 km 2) and the continued loss of habitat at these localities.

Oedipina kasios sp. nov. Muralla Worm Salamander Figure 5 Oedipina cyclocauda (part). McCranie & Wilson 2002, McCranie & Casta��eda 2007. Holotype. MVZ 232825, an adult female, from near Quebrada Pinol, 15 ��07���N, 86 �� 44 ���W, Parque Nacional La Muralla, 1190 m a.s.l., Dept. Olancho, Honduras, collected 19 July 2001 by C. Sheehy, J. H. Townsend, and L. D. Wilson. Paratypes (5). USNM 343455 ��� 56, confluence of quebradas Pinol and Las Cantinas, 15 ��09���N, 86 �� 43 ���W, Parque Nacional La Muralla, 950 m a.s.l., Dept. Olancho, Honduras; MVZ 232826, same data as for holotype, except 1230 m a.s.l.; USNM 530580, Cerro de Enmedio, 15 ��06���N, 86 �� 41 ���W, Parque Nacional La Muralla, 1780 m a.s.l., Dept. Olancho, Honduras; USNM 343457, Quebrada de la Escaleras, 15 �� 12 ���W, 86 �� 41 ���W, Parque Nacional La Muralla, 950 m a.s.l., Dept. Olancho, Honduras. Referred material (1). USNM 530581 (juvenile), same data as paratype USNM 530580. Diagnosis. A small (maximum known size 46.8 mm SL) species distinguished from Oedipina cyclocauda of Costa Rica (the species with which this population was previously tentatively identified in McCranie & Wilson 2002 and McCranie & Casta��eda 2007) by having the tail nearly rectangular in cross section throughout its length (versus nearly round for most or all of its length in O. cyclocauda; McCranie pers. obs., also see Taylor 1952); from O. quadra by having fewer maxillary teeth, silver white dorsolateral spots and flecks visible to the unaided eye in life, and a smaller maximum SL; from all described Honduran species of the subgenus Oedipina (O. ignea, O. stuarti and O. taylori by its much smaller size (or inferred smaller size, see below); from the third new species described herein by inferred smaller size and better defined digits; from all described Honduran species of the subgenus Oedopinola (O. elongata, O. gephyra and O. tomasi) by having 19���20 costal grooves. In addition to these features this species is distinct from all others that have been studied in having phylogenetically distinct mtDNA haplotypes (Fig. 1). Description. This is a small, slender species; adult SL for three males is 35.7���37.5, x = 36.9 mm; for three females 33.0��� 46.8, x = 40.6 mm. The head is small, narrow, dorsally flattened, with a broadly rounded snout. SL averages 12.5 times head width in three males (12.1���12.8) and 12.9 in three females (12.2���13.4). SL averages 6.6 times head length in three males (6.5���6.7) and 7.3 in three females (6.5���8.2). Nostrils are small but conspicuous under magnification. Nasolabial protuberances are weakly developed in males and inconspicuous in females. Eyes are small and barely extend beyond the lateral margins of the head and are directed frontolaterally. Male mental glands are not evident. The suborbital groove does not intercept the lip line. There are 1���2 slightly enlarged premaxillary teeth in males that lie anteriorly near the lip line. There are 4���5 premaxillary teeth in females that are located well within the mouth and in line with the small maxillary teeth. Maxillary teeth number 26���38 (x = 33.3) in three males and 30���38 (x = 34.3) in three females. Vomerine teeth are 10���15 (x = 12.3) in three males and 16���18 (x = 17.3) in three females; the small teeth are in an arched series. There are 19���20 costal grooves between the small limbs with a limb interval of 13 in three males and 10.5 ��� 13.0 in three females. Hands and feet are tiny, narrow and elongated. Digit I is fused with digit II and digit IV is fused with digit III on the forelimbs, with 1.0��� 1.5 segments (phalanges) of digit III free between digits II��� III on the forelimbs. Digit I is fused with digit II and digit V is fused with digit IV on the hind limbs, with 1.5 ��� 2.0 segments on both sides of digit III free on the hind limbs (Fig. 3). Protruding digital tips are acutely rounded to bluntly rounded and bear weak subdigital pads. Digits on the forelimbs in order of decreasing length are III���II���IV���I; digits on the hind limbs are III���II���IV���V���I. The tail is nearly rectangular in cross section, tapering on distal one-third of its length, and long (1.6���1.8 + times SL in three). Measurements (in millimeters), limb interval and tooth counts of the male holotype. Head width 3.2; snout to gular fold (head length) 5.1; head depth at posterior angle of jaw 1.7; eyelid width 0.8; eyelid length 1.2; eye to nostril 0.5; anterior rim of eye to snout 1.2; horizontal orbital diameter 0.6; interorbital distance 2.4; distance separating eyelids 1.2; nostril diameter 0.2; snout projecting beyond mandible 0.3; distance from eye to distal end of postorbital groove 1.4; snout to posterior angle of vent (SL) 41.9; snout to anterior angle of vent 39.2; snout to forelimb 7.9; axilla to groin 28.4; limb interval 12; shoulder width 2.2; tail length 76 + (tip removed for tissue sample); tail width at base 1.8; tail depth at base 1.9; forelimb length (to tip of longest digit) 3.3; hind limb length 3.5; forelimb foot width 0.6; hind limb foot width 0.8; free length of longest digit on hind limb 0.1. Numbers of teeth: premaxillary 5; maxillary 35; vomerine 16. Coloration of the holotype in alcohol. Dorsal and lateral surfaces of the head, body, and tail are medium brown with a grayish tinge on the body and tail. Tiny white spots or flecks are present on the lateral surface of the body. Costal grooves on the body and tail are pale brown. Ventral surfaces of the head, body, and tail are slightly paler brown than the dorsal surfaces and without a grayish tinge. Dorsal and ventral surfaces of the limbs are the same as for the body, except that the soles of all four feet are pale brown to cream. Tiny, generally pale brown iridophores are visible throughout under magnification, except on the soles of the feet. Color variability. The paratypes closely resemble the holotype in coloration, except that one (USNM 530580) is slightly darker brown overall and some (i.e., MVZ 232826) have more tiny white spots or flecks on the lateral surface of the body. McCranie & Wilson (2002: 152) described the color in life of two males (USNM 343455 ��� 56) as: ���all dorsal surfaces Grayish Brown (20), although that of head a slightly paler shade of brown; dorsolateral surface of body with silver-white flecks visible to unaided eye; ventral and subcaudal surfaces similar to dorsal surface of head.��� Those same authors also described color in life of a female (USNM 530580) as: ���all dorsal surfaces Jet Black (89), although that of head a slightly paler shade of black; dorsolateral surface of body with numerous scattered olive-green and white flecks visible to unaided eye; ventral and subcaudal surfaces Dark Neutral Gray (83), except chin somewhat paler; iris black.��� Habitat and distribution. Salamanders of this species were collected from 950���1780 m elevation in the Premontane Wet Forest and Lower Montane Wet Forest formations of Holdridge (1967). All specimens were taken in pristine broadleaf forest ranging from partial to complete canopy cover. Most were found under rotten logs, but two were inside a rotten log that also contained four Nototriton lignicola. All known localities are within Parque Nacional La Muralla in northwestern Olancho (Fig. 4). Comments. The specimen referred to this species is a juvenile and was not useful in preparing the description. Etymology. The specific name kasios is Greek for sister and is used in reference to this species being the sister species of Oedipina quadra in our phylogenetic analysis. The name is used as a noun in apposition., Published as part of Mccranie, James R., Vieites, David R. & Wake, David B., 2008, Description of a new divergent lineage and three new species of Honduran salamanders of the genus Oedipina (Caudata, Plethodontidae), pp. 1-17 in Zootaxa 1930 on pages 11-13, DOI: 10.5281/zenodo.184828, {"references":["McCranie, J. R. & Wilson, L. D. (2002) The Amphibians of Honduras. Society for the Study of Amphibians and Reptiles, Ithaca, New York, 625 pp.","McCranie, J. R. & Castaneda, F. E. (2007) Guia de Campo de los Anfibios de Honduras. Bibliomania!, Salt Lake City, 304 pp.","Taylor, E. H. (1952) The salamanders and caecilians of Costa Rica. University of Kansas Science Bulletin, 34 (12), 695 - 791.","Holdridge, L. R. (1967) Life Zone Ecology. Revised Edition. Tropical Science Center, San Jose, Costa Rica, 206 pp."]}

Oedipina quadra sp. nov. Honduran Lowland Worm Salamander Figure 2 Oedipina cyclocauda (part). Brame 1968, McCranie & Wilson 2002, McCranie & Casta��eda 2007. Holotype. MVZ 257761, an adult male, from Urus Tingni Kiamp, 14 �� 55 ���N, 84 �� 41 ���W, tributary of upper portion of R��o Warunta, 160 m above sea level (a.s.l.), Dept. Gracias A Dios, Honduras, collected 7 February 2006 by J. R. McCranie. Paratypes (29). MVZ 257755, USNM 563379 ��� 80, same data as holotype, except collected on different dates; MVZ 257756, USNM 563378, Warunta Tingni Kiamp, 14 �� 55 ���N, 84 �� 41 ���W, 150 m a.s.l., Dept. Gracias A Dios, Honduras; USNM 560949, between Urus Tingni Kiamp and Warunta Tingni Kiamp, 190 m a.s.l., Dept. Gracias A Dios, Honduras; USNM 560948, Cabeceras de R��o Rus Rus, 14 �� 53 ���N, 84 �� 40 ���W, 190 m a.s.l., Dept. Gracias A Dios, Honduras; MVZ 232824, MVZ 257757, Kaska Tingni, 14 �� 48 ���N, 84 �� 46 ���W, 70 m a.s.l., Dept. Gracias A Dios, Honduras; USNM 534115 ��� 19, Quebrada Mach��n, 15 �� 19 ���N, 85 �� 17 ���W, 540 m a.s.l., Dept. Col��n, Honduras; CM 68241, LSUMZ 33608, Cerro Calentura S of Trujillo, Dept. Col��n, Honduras; USNM 343452, confluence of r��os Yanguay and Wamp��, 15 ��03���N, 85 ��08���W, 110 m a.s.l., Dept. Olancho, Honduras; USNM 343453, confluence of r��os Sausa and Wamp��, 15 ��04���N, 85 ��06���W, 100 m a.s.l., Dept. Olancho, Honduras; USNM 343454, confluence of Quebrada Siksatara and R��o Wamp��, 15 ��04���N, 85 ��02���W, 95 m a.s.l., Dept. Olancho, Honduras; USNM 316539, 7.4 km SE of La Ceiba, 260 m a.s.l., Dept. Atl��ntida, Honduras; LACM 4702 ���04, LSUMZ 21327, mountain S of Corozal, 200���250 m a.s.l., Dept. Atl��ntida, Honduras; SMF 77486, USNM 530579, MVZ 257758 - 60, Parque Nacional Pico Bonito Centro de Visitantes (also called Estaci��n Forestal de CURLA), 15 �� 42 ���N, 86 �� 51 ���W, 120���500 m a.s.l., Dept. Atl��ntida, Honduras. Referred material (4). LSUMZ 21326 (juvenile), same data as paratype LSUMZ 21327; UMMZ 58610 (juvenile), R��o Claura, Dept. Col��n, Honduras; BMNH 1985.1229 ��� 30 (both poorly preserved), Quebrada Limoncito, 250 m a.s.l., Dept. Col��n, Honduras. Diagnosis. A moderate-sized (maximum known size 55.5 mm SL) species distinguished from Oedipina cyclocauda of Costa Rica (the species with which this population was previously tentatively identified in McCranie & Wilson 2002 and McCranie & Casta��eda 2007) by having the tail nearly rectangular in cross section throughout its length (versus nearly round for most or all of its length in O. cyclocauda; McCranie pers. obs., also see Taylor 1952); from the second new species described herein by having more maxillary teeth, a larger maximum SL, and by lacking silver white dorsolateral spots and flecks in life; from all described Honduran species of the subgenus Oedipina (O. ignea, O. stuarti, O. taylori and the third species described herein) in the following ways: from ignea in having a nearly rectangular tail and smaller size; from stuarti in lacking pale brown to dirty white small glandular spots on the head and body and by having a smaller size; from taylori in having 19���20 costal grooves, maxillary teeth, and a smaller size; from the third new species described below by inferred smaller size and better defined digits; from all described Honduran species of the subgenus Oedopinola (O. elongata, O. gephyra and O. tomasi) by having 19���20 costal grooves. In addition to these features this species is distinct from all others that have been studied in having phylogentically distinct mtDNA haplotypes (Fig. 1). Description. This is a moderately sized, slender species; adult SL for seven males is 36.1���43.7, x = 41.6 mm; for 16 females 35.6���55.5, x = 45.0 mm. The head is small, narrow, and dorsally flattened, with a broadly rounded snout. SL averages 11.6 times head width in seven males (9.8���13.1) and 12.2 in 14 females (10.7��� 13.7). SL averages 6.4 times head length in seven males (5.6���6.7) and 6.7 in 14 females (5.7���7.7). Nostrils are small but conspicuous under magnification. Nasolabial protuberances are weakly developed in males and inconspicuous in females. Eyes are small and barely extend beyond the lateral margins of the head and are directed frontolaterally. Male mental glands are small, barely evident, located anteriorly on lower jaw. The suborbital groove does not intercept the lip line. There are 1���3 slightly enlarged premaxillary teeth in males that lie only slightly anteriorly to the maxillary teeth. There are 2���6 premaxillary teeth in females that are located well within the mouth and in line with the small maxillary teeth. Maxillary teeth number 37���47 (x = 42.3) in seven males and 28���53 (x = 43.0) in 13 females. Vomerine teeth are 15���20 (x = 17.5) in six males and 16���22 (x = 19.5) in 11 females; the small teeth are in an arched series. There are 19���20 costal grooves between the small limbs with a limb interval of 11.0���13.0 in four males and 14 females. Hands and feet are tiny, narrow and elongated. Digit I is fused with digit II and digit IV is fused with digit III on the forelimbs, with about 1.0 segments (phalanges) of digit III free between digits II���III on the forelimbs. Digit I is fused with digit II and digit V is fused with digit IV on the hind limbs, with about 1.0���2.0 segments on both sides of digit III free on the hind limbs (Fig. 3). Protruding digital tips are acutely rounded to bluntly rounded and bear weak subdigital pads. Digits on the forelimbs in order of decreasing length are III���II���IV���I; digits on the hind limbs are III���II��� IV ���V���I. The tail is nearly rectangular in cross section in almost all (somewhat rounded basally in MVZ 257759), tapering on distal one-third of its length, and long (1.3 [regenerated]��� 2.4 times SL in 13). Measurements (in millimeters), limb interval and tooth counts of the male holotype. Head width 3.2; snout to gular fold (head length) 6.8; head depth at posterior angle of jaw 2.2; eyelid width 0.8; eyelid length 1.0; eye to nostril 0.8; anterior rim of eye to snout 1.2; horizontal orbital diameter 0.5; interorbital distance 3.1; distance separating eyelids 1.3; nostril diameter 0.3; snout projecting beyond mandible 0.3; distance from eye to distal end of postorbital groove 2.0; snout to posterior angle of vent (SL) 42.5; snout to anterior angle of vent 39.3; snout to forelimb 9.8; axilla to groin 26.9; limb interval 12 1 / 2; shoulder width 2.3; tail length 93; tail width at base 2.2; tail depth at base 2.2; forelimb length (to tip of longest digit) 2.2; hind limb length 3.2; forelimb foot width 0.9; hind limb foot width 1.2; free length of longest digit on hind limb 0.1. Numbers of teeth: premaxillary 2; maxillary 38; vomerine 15. Coloration of the holotype in alcohol. Dorsal and lateral surfaces of the head, body, and tail are black. Costal grooves on the body and tail are dirty white. Ventral surfaces of the head, body, and tail are slightly paler black than those dorsal surfaces. Dorsal and ventral surfaces of the limbs are the same as for the body. Tiny, generally white to dirty white iridophores are visible throughout under magnification. There is a large patch of skin on the left side of the body where the black pigment is missing, perhaps the result of an old injury or a skin disease. Color variability. The paratypes closely resemble the holotype, with the exception of the abnormal loss of pigment in the holotype, noted above. McCranie & Wilson (2002: 152) described color in life for one female (USNM 343452) as: ���dorsal surfaces of head, body, and tail black; lateral surfaces of head and body brownish black; lateral surface of tail black; dorsal surfaces of limbs and ventral and subcaudal surfaces brownish black.��� Habitat and distribution. Salamanders of this species were collected from 70���540 m elevation in the Lowland Moist Forest formation of Holdridge (1967). Specimens were taken in either pristine broadleaf forest with a closed canopy or in secondary broadleaf forest with a partially open canopy. All were collected while raking through leaf litter during the day and at night. The known distribution of this species extends from just south of La Ceiba, Atl��ntida, eastward and southeastward to the R��o Coco drainage system of northeastern Honduras (Fig. 4). The species probably occurs in suitable habitat on the Nicaraguan side of the R��o Coco as well. Comments. A few specimens are treated as referred material. We are confident of their assignment but they are either juveniles or poorly preserved and hence of no use in our description. Etymology. The specific name quadra is a Latin adjective for square and alludes to the nearly rectangular tail characteristic of this species., Published as part of Mccranie, James R., Vieites, David R. & Wake, David B., 2008, Description of a new divergent lineage and three new species of Honduran salamanders of the genus Oedipina (Caudata, Plethodontidae), pp. 1-17 in Zootaxa 1930 on pages 6-10, DOI: 10.5281/zenodo.184828, {"references":["McCranie, J. R. & Wilson, L. D. (2002) The Amphibians of Honduras. Society for the Study of Amphibians and Reptiles, Ithaca, New York, 625 pp.","McCranie, J. R. & Castaneda, F. E. (2007) Guia de Campo de los Anfibios de Honduras. Bibliomania!, Salt Lake City, 304 pp.","Taylor, E. H. (1952) The salamanders and caecilians of Costa Rica. University of Kansas Science Bulletin, 34 (12), 695 - 791.","Holdridge, L. R. (1967) Life Zone Ecology. Revised Edition. Tropical Science Center, San Jose, Costa Rica, 206 pp."]}

Oedipina leptopoda sp. nov. Narrow-footed Worm Salamander Oedipina cyclocauda (part), Brame 1968, Good & Wake 1998, Garc��a-Par��s & Wake 2000, McCranie & Wilson 2002, McCranie & Casta��eda 2007. Holotype. MVZ 167772, a subadult female from 32 km (road) W of Yoro on road to Moraz��n, 15.267480 N, 87.434820 W, Dept. Yoro, Honduras, collected 8 January 1979 by E. J. Koford and J. F. Lynch. Paratype. MVZ 171078, adult male from Monta��a de Yoro, 6.6 km (road) S of Yoro, 15.073650 N, 87.1333 W, Dept. Yoro, Honduras. Referred material (1). FMNH 34683 (subadult), Portillo Grande, Dept. Yoro, Honduras. Diagnosis. A small to moderate sized (holotype is a subadult at 29.6 mm SL) member of Oedipina. Distinguished from Oedipina cyclocauda of Costa Rica (the species with which this population was previously identified by Good & Wake 1998 and Garc��a-Par��s & Wake 2000) by its inferred larger size and narrower hands and feet with less discrete digits. It differs from other described Honduran species of the subgenus Oedipina (O. ignea, O. stuarti, and O. taylori) in the following ways: from ignea in having more maxillary teeth and narrower hands and feet; from stuarti in being smaller, having much narrower hands and feet, and lacking glandular spots on its head and body; from taylori in having a broader head, narrower hands and feet, shorter body and tail, and 19-20 costal grooves; from members of the subgenus Oeditriton (O. quadra and O. kasios) in being larger (based on the size of the paratype), having relatively smaller limbs and hands and feet, and a rounded body and tail; from all described Honduran species of the subgenus Oedopinola (O. elongata, O. gephyra and O. tomasi) by having 19���20 costal grooves. In addition to these features, this species differs in allozymic (Good & Wake 1998) and mtDNA characters (Fig. 1) from all members of the genus in which those characters are known Description. This is a slender species of moderate size, judging from the fact that the paratype is an adult male, the smaller sex in this genus, at a size larger than that of several other species in the subgenus. The head is relatively broad (SL 7.4-9.8 times head width), rounded rather than flattened, and with a narrowly rounded snout (expanded and broadly rounded in the paratype, an adult male 47.9 SL). Nostrils are small and conspicuous. Nasolabial protuberances are moderate. Eyes are of moderate size and barely extend beyond the lateral margins of the head. The suborbital groove does not intercept the lip line. There are 5 small premaxillary teeth in the holotype and a single slightly larger tooth in the adult male paratype. The teeth are near the front but inside of the mouth, well separated from the maxillary teeth. There are 29 maxillary teeth in the holotype and 40 in the paratype. The holotype has 10 vomerine teeth and the paratype has 17. The vomerine teeth are small and in an arched series. The holotype and paratype have 19 costal grooves between the slender limbs, with a limb interval of 12 in both specimens. Hands and feet are extraordinarily narrow, small and flattened (Fig. 3). In the holotype digits can be discerned only because there are ill-defined grooves on the dorsal surfaces that suggest where digital borders lie. All of the digits are tightly fused, all the way to their tips in the holotype but there are small, rounded, independent tips of the three longest digits of both forelimbs and hind limbs of the paratype. In the holotype the longest digit is pointed, and it extends less then 0.2 mm from the ends of the neighboring digits. The holotype lacks a tail and has been almost completely eviscerated (for tissue sample used in allozyme and DNA studies). The paratype has a rather stout, round tail that is broken 44.4 mm from its base, and it doubtless greatly exceeded SL in the intact animal. Measurements (in millimeters), limb interval and tooth counts of the female holotype. Head width 4.0; snout to gular fold (head length) 5.4; head depth at posterior angle of jaw 2.1; eyelid width 0.5; eyelid length 1.3; eye to nostril 1.2; anterior rim of eye to snout 1.3; horizontal orbital diameter 1.0; interorbital distance 1.7; distance separating eyelids 1.1; nostril diameter 0.2; snout projecting beyond mandible 0.2; distance from eye to distal end of postorbital groove 1.5; snout to posterior angle of vent (SL) 29.6; snout to anterior angle of vent 28.3; snout to forelimb 7.5; axilla to groin 18.2; limb interval 12; shoulder width 2.7; tail length 33.2; tail width at base 1.5; tail depth at base 1.8; forelimb length (to tip of longest digit) 3.9; hind limb length 4.2; forelimb foot width 0.6; hind limb foot width 0.8; free length of longest digit on hind limb 0.2. Numbers of teeth: premaxillary 5; maxillary 29; vomerine 10. Coloration of the holotype and paratype in alcohol. The holotype, while apparently well preserved, appears to be faded and the overall coloration is now moderate brown. The head, especially the snout, is significantly paler than the rest of the specimen. The paratype is all black, darker dorsally and laterally than ventrally. The swollen snout and the nasolabial protuberances are much lighter, pale gray. The small and inconspicuous mental gland is pale gray. Habitat and distribution. The species is known only from three specimens collected many years ago. Field notes are precise with respect to locality, two from west and one from south of Yoro (Fig. 4), but vague with respect to habitat. The estimated elevation for these specimens is from ca. 700 to 1300 m a.s.l. Comments. The specimen referred to this species is a poorly preserved subadult with sufficient morphological detail to make it a likely member of this species, and its collecting site is also appropriate for this assignment. The paratype is an intact specimen with good preservation of color, whereas the holotype is eviscerated and faded but otherwise well preserved. However, because the two specimens are not syntopic and because we have molecular data only from the holotype, we elected to use the specimen for which we have the best documentation as the holotype. Etymology. The specific name leptopoda is derived from the Greek lepto- for small or fine and podos for foot, and is used as an adjective in reference to the very small and narrow hands and feet of this species., Published as part of Mccranie, James R., Vieites, David R. & Wake, David B., 2008, Description of a new divergent lineage and three new species of Honduran salamanders of the genus Oedipina (Caudata, Plethodontidae), pp. 1-17 in Zootaxa 1930 on pages 13-15, DOI: 10.5281/zenodo.184828, {"references":["Good, D. A. & Wake, D. B. (1998; dated 1997) Phylogenetic and taxonomic implications of protein variation in the Mesoamerican salamander genus Oedipina (Caudata: Plethodontidae). Revista de Biologia Tropical, 48 (3), 1185 - 1208.","Garcia-Paris, M. & Wake, D. B. (2000) Molecular phylogenetic analysis of relationships of the tropical salamander genera Oedipina and Nototriton, with descriptions of a new genus and three new species. Copeia, 2000 (1), 42 - 70.","McCranie, J. R. & Wilson, L. D. (2002) The Amphibians of Honduras. Society for the Study of Amphibians and Reptiles, Ithaca, New York, 625 pp.","McCranie, J. R. & Castaneda, F. E. (2007) Guia de Campo de los Anfibios de Honduras. Bibliomania!, Salt Lake City, 304 pp."]}

Fossorial salamanders typically have elongate and attenuated heads and bodies, diminutive limbs, hands and feet, and extremely elongate tails. Batrachoseps from California, Lineatriton from eastern México, and Oedipina from southern México to Ecuador, all members of the family Plethodontidae, tribe Bolitoglossini, resemble one another in external morphology, which has evolved independently. Whereas Oedipina and Batrachoseps are elongate because there are more trunk vertebrae, a widespread homoplasy (parallelism) in salamanders, the genus Lineatriton is unique in having evolved convergently by an alternate “giraffe-neck” developmental program. Lineatriton has the same number of trunk vertebrae as related, nonelongated taxa, but individual trunk vertebrae are elongated. A robust phylogenetic hypothesis, based on sequences of three mtDNA genes, finds Lineatriton to be deeply nested within a clade characterized by generalized ecology and morphology. Lineatriton lineolus , the only currently recognized taxon in the genus, shows unanticipated genetic diversity. Surprisingly, geographically separated populations of L. lineolus are not monophyletic, but are sister taxa of different species of the morphologically generalized genus Pseudoeurycea. Lineatriton , long thought to be a unique monospecific lineage, is polyphyletic. Accordingly, the specialized morphology of Lineatriton displays homoplasy at two hierarchical levels: ( i ) with respect to other elongate lineages in the family (convergence), and ( ii ) within what is currently recognized as a single taxon (parallelism). These evolutionary events are of adaptive significance because to invade the lowland tropics salamanders must be either arboreal or fossorial; the repeated evolution of elongation and attenuation has led to multiple lowland invasions.

Several recent publications have made recommendations for changes in the taxonomy of plethodontid salamanders. Here formal taxonomic proposals are made, in accordance with the Code, regarding family-group taxa. Subfamilies Hemidactyliinae and Plethodontinae are recognized, the former with four tribes and the latter with five tribes. Genera are assigned to the tribes, and subgenera are recognized in the genera Batrachoseps, Bolitoglossa, Hydromantes, Oedipina and Plethodon.

Although plethodontid salamanders are characterized by a diversity of life histories, several aspects of their breeding biology are highly conserved. Two of these are solitary oviposition and attendance of clutches by females until hatching. We infer evolutionary changes in these reproductive features based on the discovery of oviposition sites and eggs of two species, Batrachoseps nigriventris and B. wrighti. Batrachoseps nigriventris deviates from the ancestral reproductive pattern in two ways: most clutches are laid at communal oviposition sites and female care has been lost. In contrast, B. wrighti may deviate from the ancestral pattern only in the loss of maternal care. A review of the reproductive biology of Batrachoseps and other plethodontids reveals that female attendance of eggs was lost in the ancestor of all Batrachoseps, whereas communal oviposition likely originated following the basal split within the genus. Additionally, communal oviposition and abandonment of clutches have evolved in two other plethodontid clades: Nototriton/Oedipina and Hemidactylium. Studies of taxa that have lost parental care may give insight into factors favoring the maintenance of parental care in other lineages. LETHODONTID salamanders are characterized by a diversity of life histories, in

A new genus and species of plethodontid bolitoglos- sin: salamander is described from material collected in northwestern Guatemala. Bradytriton silus new genus, new species, is unique in a cornbination of structural characteristics that includes a laterally cornpressed tail, stocky body with no clearly defined neck, and short, slender limbs bearing syndactylous hands and feet. To diagnose the new genus, an analysis of the entire neotropical assemblage of pleth- od(intid salamanders was undertaken. Approximately 138 species bel Jng to the supergenus Bolitoglossa and 1 1 genera are recognized. Th: genus Chiropterotriton is shown to be polyphyletic; thus, two new genera are described. Nototriton new genus, includes the picadoi group of Chiropterotriton beta. Dendrotriton new genus, includes the hrc meliacia group of Chiropterotriton beta. Those species formerly cal ed Chiropterotriton alpha remain as the sole representatives of the genus. Eight of the eleven neotropical genera are shown to be monophyletic. Of the three exceptional genera, both Dendrotriton and Nototriton are nonparaphyletic relative to all genera except Oed- ipiaa. but Dendrotriton is easily distinguished from Oedipina. No- tot,iton may be paraphyletic relative to Oedipina, but the two genera car be readily distinguished on the basis of major differences in ecc logy and shape. Pseudoeurycea comprises morphologically gen- eralized species that may not be far removed from the morphology off he ancestral stock ofthe entire neotropical group. Only Nyctanolis and Chiropterotriton are more plesiomorphic than Pseudoeurycea. Lists of species assigned to the genera are provided. Potential rela- tio iships within the supergenus Bolitoglossa are discussed, but par- alldism and convergence have been so extensive that no definitive sta ement concerning generic affinities is possible.

Mental hedonic gland clusters of 18 species of eastern United States plethodontids were studied by gross dissection and light microscopy. In the Plethodontinae examined, the glands are largely or completely posterior to the mandibular symphysis. In the Plethodontini, the glands are relatively short, vertically oriented in a circular cluster, and bordered dorsally by the dense inner layer of the dermis, the stratum compactum. Within the Hemidactylini genus Eurycea, the glands are more elongate and excretory ducts only occur ventral to and immediately posterior of the mandibular symphysis. The glands then pass horizontally between the intermandibularis posterior muscle ventrally and the geniohyoideus muscle dorsally. The stratum compactum is reduced to a thin sheath over the glands. Reduced or specialized glands resembling those of the Plethodontini occur in the hemidactyliines Hemidactylium, Typhlotriton, and E. multiplicata. From literature reports, the Bolitoglossini contain seven genera with glands like the Plethodontini and one (Oedipina) with glands like Eurycea. In the Desmognathinae, except for Desmognathus wrighti, the excretory ducts are in a fleshy protrusion anterior to the mandibular symphysis, and the short glands extend posteriorly, terminating ventral to and just posterior of the mandibular symphysis. In D. wrighti the gland cluster bifurcates anteriorly with elongate glands then extending posteriorly between the dermis and throat musculature along the medial borders of the dentaries. The excretory ducts in D. wrighti pass dorsally around the anterior ends of the dentaries.