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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."]}