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Nototriton kenorum sp. nov. Kens’ moss salamander Holotype. UCR 23685 (Figs. 11–13), an adult female from Costa Rica: Provincia de Limón: Cantón de Guácimo: Distrito de Guácimo: along edge of small road on the northern slopes of the Turrialba Volcano, approximately 9 kilometers south of Jiménez, ca. 750 m a.s.l., collected by Brian Kubicki on 10 May 2014. Paratopotypes. UCR 23686 and UCR 23687, same collection data as the holotype. UCR 23688, same locality data as the holotype, but collected by Brian Kubicki on 23 July 2016. Generic Placement. Assigned to the genus Nototriton due to having fewer than 14 costal grooves, reduced manus and pes that are longer than wide, and the molecular evidence (16S, COI, and cyt b mtDNA distances) presented herein. Subgeneric Placement. Nototriton kenorum is assigned to the subgenus Nototriton due to its known geographic distribution (endemic to Costa Rica), its small hands and feet with the majority of the lengths of fingers II, III, and IV and toes II, III, IV, and V being free of extended palmar or plantar tissue, and by the molecular evidence (16S, COI, and cyt b mtDNA distances) presented herein. Diagnosis. The combination of the following characteristics can be used to distinguish Nototriton kenorum from the other described species of the subgenus Nototriton: (1) having small hands and feet, with at least the distal phalanges on the fingers II, III, and IV and toes II, III, IV, and V being free of the margin of the palmar and plantar tissue; (2) tiny nostril openings, both RNW and LNH not known to exceed 0.09 mm; (3) a wide head, with HeW known to exceed 4.5 mm; (4) very narrow hands and feet, narrowest among the known members of the subgenus Nototriton within Costa Rica, HaW not known to exceed 34.1 % of HeW and FoW not known to exceed 43.2% of HeW; (5) 16S, COI, and cyt b mtDNA distances. Description of holotype. Adult female having a total length of 73.5 mm and SL of 33.5 mm. Head slightly wider than neck and shoulders (HeW 4.4 mm, NeW 4.1 mm, ShW 4.0 mm), with greatest width of head just posterior to the articulation of the jaws; snout raised anterodorsally, spadate to rounded in dorsal outline, and rounded in profile; snout relatively short (SnL 1.1 mm, 3.3 % of SL), with nearly terminal non-protruding tiny nostrils (LNH 0.09 mm, RNW 0.09 mm) directed anterolaterally; internarial area convex in dorsal outline. Eyes relatively large (EW = 145 % of SnL), protruding beyond dorsal and ventral outline of head, directed anterolaterally, with a distinct suborbital groove. Top of head flat and smooth, tapering slightly toward anterior terminus, lacking contrasting interorbital or other dermal structures. Canthus rostralis weak to rounded along posterior half, indiscernible along anterior half; intercanthal area flat to slightly convex; and loreal region flat to slightly concave. No obvious cirri (nasolabial protuberances) present, but nasolabial grooves very weakly discernible on tip of snout; nasolabial grooves start at lateral margins of nares and extend ventrally at first, but take a drastic backwards curve about halfway to lip; nasolabial grooves terminate just prior to reaching margin of upper lip. Gular fold well-defined, starting on dorsolateral portion of neck, below postorbital groove, wrapping around posterolateral section of head at a slightly anterior angle and crossing venter as a smooth and weak anterior-oriented curve. Nuchal grooves weakly evident, starting above postorbital groove and converging medially at occiput. Postorbital grooves weakly defined, starting at the posterior corner of eyes and traveling horizontally along the inferior margin of parotoid glands, terminating at gular fold on the lateral portion of neck. Horizontal mandibular grooves weakly discernible, but vertical mandibular groove welldefined. Horizontal mandibular groove starting at corner of mouth and extending horizontally to vertical portion of mandibular groove. Mandibular groove existing as a single solid structure, starting along inferior margins of postorbital grooves and traveling vertically to and crossing gular region, just posterior of inner mandibular grooves. Inner mandibular grooves weakly evident, principally discernible for a small section, running medially parallel to posterior portion of mandibular bones, becoming indiscernible near anterior tip of lower jaw. Pair of very weakly, but discernible, raised parotoid glands present on dorsolateral margin of head between orbits, postorbital grooves, and occiput. Snout not evidently protruding beyond anterior margin of lower lip in lateral view. No mental gland visible under skin of anterior intermandibular region. Arms relatively long and slender overall (FLL = 4.8 mm, 14.3 % of SL), without noticeable hypertrophied forearm compared to upper arm. Hands very small and slender (HaL = 1.7 mm, 26.6 % of VGS; HaW = 1.5 mm, 34.1 % of HeW). Fingers II, III, and IV short and thick, protruding freely, with at least ultimate and all to part of penultimate phalanges being free beyond interdigital tissue margin (LF2 0.62 mm, LF3 0.53 mm); no discernible indentation at interdgital space between fingers I and II. Tips of fingers rounded, with terminal pads discernible on ventral surface. Palmar surfaces appearing to be smooth overall. Dorsal surfaces of hands with discernible interdigital grooves, especially between fingers II-III and III-IV, that start at interdigital tissue margins and cross onto metacarpal region. Relative lengths of fingers on right hand I Coloration in life. The ground color of the dorsal and lateral surfaces of the head and trunk consisted of a nearly uniform fleshy-orange coloration. There were not many secondary markings on the dorsal and lateral surfaces of the head and trunk, but a small concentration of larger irregular dark brown markings were visible on the snout, nuchal region, and hips. Additionally, there were numerous very fine white chromatophores scattered thourghout the dorsal dorsal and lateral surfaces of the head and body. The iris was bright orangish-red with a dark brownishmaroon reticulation. The upper surfaces of the arms and legs were very similar in coloration and pattern to that of the abovementioned chromatic characteristics of the dorsal and lateral surfaces of the head and body, but there appeared to be a slightly brighter orange hue on the dorsal surfaces of the limbs. The dorsal and dorsolateral surfaces of the tail were nearly uniform fleshy-orange, but were similar to that of the limbs in having a slightly more brighter orange hue. Additionally, there were some irregular dark brown patches, one very evident patch was found about one third down (anterior towards posterior orientation) the length of the tail. As on the rest of the dorsal and lateral surfaces of the holotype, numerous very fine white chromatophores were visible scattered randomly throughout the tail. The ventrolateral surfaces of the head, limbs, body, and tail were contrasted from the superior surfaces by consisting of a concentration of fine white or pale fleshy-orange markings on a darker brownish-gray background. The gular patch was slightly more pale in comparison to the other ventral surfaces. The palmar and plantar surfaces were pale gray, lacking discernible chromatophores, however a red reticulated pattern was discernible under the skin, formed by a network of blood-filled capillaries. Coloration in ethanol. After more than five years in ethanol (70%), the overall coloration of the holotype has darkened throughout and contains a principal dark brown tone on the dorsal surfaces of the head and body. Two pale tannish-yellow patches are visible on the posterior dorsolateral surfaces of the head, where the parotoid glands are located. The dorsal surface of the entire tail is also a pale tannish-yellow coloration, with the exception of a darker brownish gray transversal band, located about one third down (anterior towards posterior orientation) the length of the tail. The ventral surfaces of the body and tail are darker brownish-gray. The ventral surface of the head has an evident paler tone compared to that of the rest of the body and tail. Measurements (in mm), limb interval, and percentages of the holotype. SL 33.5; total length 73.5; tail length 40.0; ShW 4.0; HeW 4.4; NeW 4.1; EW 1.6; SnL 1.1; JSL 3.9; LGFS 7.0; LNH 0.09; RNW 0.09; IND 1.1; NLP 0.5; ICD 2.0; HLL 5.3; FLL 4.8; TW 4.4; VGS 6.4; FSL 8.6; UHL 3.6; AGL 19.3; VL 1.9; HaW 1.5; HaL 1.7; LF2 0.62; LF3 0.53; WF3 0.40; FoW 1.9; FoL 2.3; LT2 0.78; LT3 0.65; WT3 0.43. Limb interval six. Measurements in related percentages: VGS/SL 19.1 %; IND/HeW 25.0 %; AGL/SL 57.6 %; HeW/SL 13.1 %; Hew/AGL 22.8 %; SnL/ HeW 25.0 %; LNH/HeW 2.0 %; LNH/SL 0.27 %; RNW/HeW 2.0 %; RNW/SL 0.27 %; HLL/SL 15.8 %; FLL/SL 14.3 %; HaL/VGS 26.6 %; FoL/VGS 35.9 %; Haw/HeW 34.1 %; FoW/HeW 43.2 %; LT2/FoL 33.9 %; LF2/HaL 36.5 %; WT3/FoW 22.6 %; WF3/HaW 26.7 %. Noteworthy variation. The paratopoype UCR 23688 had discernible nasolabial protruberences that extended slightly beyond the upper lip margin, where as the holotype and other two paratopotypes did not have evident nasolabial protruberences.Additionally, UCR 23688 completely lacked any discernible nostril openings, even when viewed under a disecting scope at 30x magnification. Although we did not perform any internal examination to determine the sexes of the type series specimens, we feel that it is very likely that UCR 23688 is a male and the holotype UCR 23685 and the other two paratopotypes (UCR 23686 and UCR 23687) are females. Measurements (in mm), limb intervals, and percentages of the paratopotypes. SL 27.5–31.3; ShW 3.2– 4.1; HeW 3.9–4.8; NeW 3.4–4.2; EW 1.3–1.5; SnL 0.9–1.2; JSL 3.3–4.1; LGFS 6.0–7.0; LNH 0.00–0.09; RNW 0.00–0.09; IND 0.9–1.1; NLP 0.4–0.7; ICD 1.7–2.1; HLL 4.8–6.0; FLL 4.3–5.0; TW 3.3–4.4; VGS 5.1–5.9; FSL 7.5–8.2; UHL 3.1–3.8; AGL 16.5–17.8; VL 1.6–1.9; HaW 1.1–1.5; HaL 1.3–1.8; LF2 0.65–0.78; LF3 0.47–0.68; WF3 0.28–0.43; FoW 1.4–1.9; FoL 1.8–2.3; LT2 0.50–0.87; LT3 0.71–0.87; WT3 0.28–0.43. Limb intervals six. Measurements in related percentages: VGS/SL 18.4–18.8 %; IND/HeW 21.4–25.6 %; AGL/SL 56.9–58.2 %; HeW/ SL 14.2–15.3 %; Hew/AGL 22.5–27.0 %; SnL/HeW 21.4–25.0 %; LNH/HeW 0.0–2.1 %; LNH/SL 0.0–0.31 %; RNW/HeW 0.0–1.9 %; RNW/SL 0.0–0.28 %; HLL/SL 17.5–19.2 %; FLL/SL 15.6–16.0 %; HaL/VGS 25.5–30.5 %; FoL/VGS 35.3–39.0 %; Haw/HeW 28.2–31.3 %; FoW/HeW 35.9–39.6 %; LT2/FoL 21.7–43.5 %; LF2/HaL 43.3–50.0 %; WT3/FoW 17.5–22.6 %; WF3/HaW 23.3–28.6 %. Etymology. The specific epithet “ kenorum ” is a patronym, used as a noun in the plural masculine genitive case. This taxon is named in honor of Kenneth Marc Kubicki (1946–2008), the father of the senior author who was always very supportive of BK’s interests, including a life-long passion with nature (especially fishes and amphibians) and natural history. Additionally, this taxon is named for Kenneth Marc Kubicki-Reyes (2009–), the son of BK and AR; to symbolize our hope and optimism with the continued study, protection, and appreciation of our world’s natural resources, which are soon going to fall into the hands of our children and their future generations. Habitat and natural history observations. Nototriton kenorum has been found exclusively within moss growing on the trunks and branches of trees. More than twenty different individuals of N. kenorum have been observed by the senior author in the wild, and all were found within moss in arboreal situations. Likely N. kenorum can also inhabit moss and other microhabitats that offer similar physical parameters, growing on rocks and terrestrial surfaces, but until now none have been discovered to inhabit such sites. Distribution. Nototriton kenorum is endemic to Costa Rica, and currently only known to inhabit a very small section of unusually low elevation cloud forest habitat within the lower portion of the Tropical Premontane Rainforest life zone (Holdridge 1967) along the northern slopes of the Turrialba Volcano. The known altitudinal range for N. kenorum is from ca. 775–900 masl. This is the only species of Costa Rican moss salamander that is thus far known to exclusively inhabit elevations below 1000 masl., Published as part of Kubicki, Brian, Reyes, Aura & Arias, Erick, 2022, Revised taxonomy and distributions of Costa Rican moss salamanders (Caudata: Plethodontidae: Nototriton), with descriptions of new taxa, pp. 451-496 in Zootaxa 5194 (4) on pages 477-481, DOI: 10.11646/zootaxa.5194.4.1, http://zenodo.org/record/7156853, {"references":["Holdridge, L. R. (1967) Life Zone Ecology. Revised edition. Tropical Science Center, San Jos, 206 pp."]}

Nototriton vereh sp. nov. Vereh moss salamander Holotype. UCR 23681 (Figs. 7–9), an adult female from Costa Rica: Provincia de Cartago: Cantón de Turrialba: Distrito de Chirripó: a private property owned by Brian Kubicki and referred to as the Río Vereh Cloud Forest Reserve, 1400 m a.s.l., collected by Brian Kubicki on 15 October 2016. Paratypes. UCR 23682, an adult female from Costa Rica: Provincia de Limón: Cantón de Turrialba: Distrito de Chirripó: moss on bank of road between Jicotea and Río Vereh, ca. 1250 m a.s.l., collected by Brian Kubicki in the company of Aura Reyes on 2 November 2013. UCR 23683, an adult male from Costa Rica: Provincia de Cartago: Cantón de Turrialba: Distrito de Chirripó: edge of road between Bajo Pacuare and Grano de Oro, ca. 1250 m a.s.l., collected by Brian Kubicki, in the company of Aura Reyes, on 13 April 2013; UCR 23684, an adult male: same locality data as UCR 23683, but collected by Brian Kubicki, in the company of Aura Reyes, on 21 March 2013. Generic Placement. Assigned to the genus Nototriton due to having fewer than 14 costal grooves, reduced manus and pes that are longer than wide, and the molecular evidence (16S, COI, and cyt b mtDNA distances) presented herein. Subgeneric Placement. Nototriton vereh is further assigned to the subgenus Taylorotriton due to the combination of its known geographic distribution (endemic to the central Caribbean region of Costa Rica), its tiny hands and feet that are pad-like in appearance, with at the most the distal phalanx on each digit protruding in a bluntly pointed tip beyond the fleshy palmar and plantar tissue, and again the molecular evidence (16S, COI, and cyt b mtDNA distances) presented herein. Diagnosis. The combination of the following characteristics can be used to distinguish Nototriton vereh from the other described species of the genus Nototriton: (1) having tiny hands and feet, with at the most only the distal phalanges on the fingers and toes free of palmar and plantar tissue; (2) large nostril openings, greater than 0.25 mm in height (Fig. X); (3) 16S, COI, and cyt b mtDNA distances. Comparisons. Since Nototriton vereh is only known to occur within Costa Rica and the phenotypic and molecular evidence presented herein strongly supports it forming part of the newly proposed Taylorotriton subgenus, the phenotypic comparisons presented are only with respect to other members of that subgeneric taxon (i.e. N. richardi, and N. tapanti). Contrasting characteristics for Nototriton vereh are presented in parentheses. Nototriton richardi (Taylor, 1949) can be distinguished from N. vereh by having relatively narrower feet, average FoW = 0.97 mm (average FoW = 0.85 mm); shorter hind limbs, average HLL = 3.7 mm and HLL = 15.7–19.5 % of SL (longer hind limbs, average HLL = 4.1 mm and HLL = 18.0–21.0 % of SL). Nototriton tapanti Good & Wake, 1993 has wider hands and feet, average HaW = 0.9 mm, HaW = 24.1–29.7 % of HeW, average FoW 1.1 = mm, FoW = 31.0–35.1 % of HeW (narrower hands and feet, average HaW = 0.7 mm, HaW = 23.3–27.6 % of HeW, average FoW 0.9 = mm, FoW = 26.7–32.1 % of HeW). Description of holotype. Subadult to adult female having a SL of 18.6 mm. Head slightly wider than neck and shoulders (HeW 2.9 mm, NeW 2.5 mm, ShW 2.5 mm), with greatest width of head just posterior to the articulation of the jaws; snout raised anterodorsally, spadate to rounded in dorsal outline, and rounded in profile; snout relatively short (SnL 0.6 mm, 3.2 % of SL), with nearly terminal non-protruding large nostrils (LNH 0.28 mm, RNW 0.22 mm) directed anterolaterally; internarial area convex in dorsal outline. Eyes relatively large (EW = 183 % of SnL), weakly protruding beyond dorsal and ventral outline of head, directed anterolaterally, with a distinct suborbital groove. Top of head flat and smooth, tapering slightly toward anterior terminus, lacking contrasting interorbital or other dermal structures. Canthus rostralis weakly rounded; intercanthal area flat to slightly convex; and loreal region slightly concave. No obvious cirri (nasolabial protuberances), but nasolabial grooves weakly discernible on tip of snout; nasolabial grooves start at ventrolateral margins of nares and extend ventrally, with a slight outward orientation, and terminate prior to reaching upper lip margin. Gular fold well-defined, starting on dorsolateral portion of neck, below postorbital groove, wrapping around posteriolateral section of head at a slightly anterior angle and crossing venter as a smooth anterior-oriented curve. Nuchal grooves very weakly evident, starting above postorbital groove and converging medially at occiput. Postorbital grooves weakly evident, starting at the posterior corner of eyes and traveling horizontally along the inferior margin of parotoid glands, terminating at gular fold on the lateral portion of neck. Horizontal mandibular groove moderately defined, starting at corner of mouth and extending horizontally to vertical portion of mandibular groove. Mandibular groove not a single continuous structure, but rather two independent structures on each side of head. Vertical portion of mandibular grooves starting along the inferior margins of postorbital grooves and traveling vertically to gular region, terminating at or just medially of intersections with inner mandibular grooves. Inner mandibular groove not a single continuous structure, but rather two independent and laterally postioned structures. Inner mandibular grooves moderately evident, starting at anterior termini or margin of mandibular grooves, running medially parallel to mandibular bones, becoming indiscernible near anterior tip of lower jaw. Pair of weak, but discernibly raised parotoid glands present on dorsolateral margin of head between orbits, postorbital grooves, and occiput. Snout protruding beyond anterior margin of lower lip in lateral view. No mental gland visible under skin of anterior intermandibular region. Arms relatively short and slender (FLL = 3.3 mm, 17.7 % of SL), without noticeable hypertrophied forearm compared to upper arm. Hands very small and slender (HaL = 0.9 mm, 22.5 % of VGS; HaW = 0.8 mm, 27.6 % of HeW). Fingers II and III protruding slightly beyond interdigital tissue margin (LF2 0.31 mm, LF3 0.19 mm), fingers I and IV with minimal indentation at interdgitial spaces, barely receding proximally beyond tips of latter mentioned digits. Finger III most free of interdigital tissue, with about entire distal phalanx protruding. Tips of fingers rounded (fingers I, II, IV) to acutely rounded (Finger III). Terminal pads weakly discernible on ventral surface of fingers. Palmar surfaces appearing to be smooth overall, but with an evident dermal crease extending transversally along proximal margin of tip of Finger III. Dorsal surfaces of hands with discernible interdigital grooves that start at interdigital tissue margins and cross metacarpal region. Relative lengths of fingers on right hand I Coloration in life. The ground color of the dorsal and dorsolateral surfaces of the head and trunk consisted of a mixture of reddish-brown earthy tones. Secondary coloration on the dorsal and dorsolateral surfaces of the head and trunk consisted of larger irregular black markings (especially concentrated along the dorsolateral region of the trunk and dorsal surface of the head), numerous closely spaced darker chevron marks visible along upper dorsum of the trunk, and numerous fine to very fine pale orange to white patches of chromatophores scattered randomly throughout the dorsal surface. The iris was bright orange with a dark brownish-black reticulation. The upper surfaces of the arms and legs were a mixture of chromatophore patches of irregular shapes and sizes, ranging in color from pale orange, pinkish-orange, dark brown, and black. Additionally, there were some fine to very fine patches of white chromatophores scattered randomly throughout the dorsal surface of the limbs. The dorsal and dorsolateral surfaces of the tail were nearly uniform reddish-orange, with some fine to very fine patches of white chromatophores scattered randomly throughout. The ventrolateral surfaces of the body and tail were contrasted from the superior surfaces by consisting of a concentration of fine white or pale markings on a dark gray background. The ventral surfaces of the head, body, limbs, and tail were pale brown to gray with numerous fine to very fine white spots and irregular markings scattered throughout. Just posterior to the cloaca, there was a concentration of contrasting pale orangish spots and irregular markings. The ventral surfaces of the limbs and gular patch were slightly paler in comparison to the ventral surfaces of the body and tail. The palmar and plantar surfaces were pale grayish-white. Coloration in ethanol. After more than 5 years in ethanol (70%), the overall coloration of the holotype has darkened throughout and contains a principal dark brown tone. Measurements (in mm), limb interval, and percentages of the holotype. SL 18.6; ShW 2.5; HeW 2.9; NeW 2.5; EW 1.1; SnL 0.6; JSL 2.5; LGFS 4.1; LNH 0.28; RNW 0.22; IND 0.7; NLP 0.4; ICD 1.4; HLL 3.9; FLL 3.3; TW 2.4; VGS 4.0; FSL 5.4; UHL 2.2; AGL 10.2; VL 1.2; HaW 0.8; HaL 0.9; LF2 0.31; LF3 0.19; WF3 0.22; FoW 0.9; FoL 1.2; LT2 0.4; LT3 0.22; WT3 0.22. Limb interval 4. Measurements in related percentages: VGS/SL 21.5 %; IND/HeW 24.1 %; AGL/SL 54.8 %; HeW/SL 15.6 %; Hew/AGL 28.4 %; SnL/ HeW 20.7 %; LNH/HeW 9.7 %; LNH/SL 1.5 %; RNW/HeW 7.6 %; RNW/SL 1.2 %; HLL/SL 21.0 %; FLL/SL 17.7 %; HaL/VGS 22.5 %; FoL/VGS 30.0 %; Haw/HeW 27.6 %; FoW/HeW 31.0 %; LT2/FoL 33.3 %; LF2/HaL 34.4 %; WT3/FoW 24.4 %; WF3/HaW 27.5%. Noteworthy variation. The male paratypes (UCR 23683 and UCR 23684) had more truncate snouts, and much more defined and protruding cirri or nasalabial protruberances, sexually dimorphic features that are often observed among plethodontids, including bolitoglossines. Measurements (in mm), limb intervals, and percentages of the paratypes. SL 20.8–22.8; ShW 2.4–2.5; HeW 2.6–3.0; NeW 2.3–2.5; EW 1.1–1.2; SnL 0.8–0.9; JSL 2.6–2.8; LGFS 4.4–4.9; LNH 0.25–0.28; RNW 0.22– 0.28; IND 0.6–0.8; NLP 0.3–0.5; ICD 1.3–1.5; HLL 4.1; FLL 3.5–4.1; TW 2.5–2.7; VGS 3.7–4.2; FSL 5.7–6.4; UHL 2.3–2.5; AGL 11.4–13.1; VL 1.5–1.7; HaW 0.7; HaL 1.0; LF2 0.28–0.40; LF3 0.19–0.25; WF3 0.22; FoW 0.8–0.9; FoL 1.2; LT2 0.31–0.40; LT3 0.22–0.28; WT3 0.22–0.23. Limb intervals 5–6. Measurements in related percentages: VGS/SL 17.1–19.2 %; IND/HeW 23.1–28.6 %; AGL/SL 54.8–57.5 %; HeW/SL 12.0–13.5 %; Hew/ AGL 21.0–24.6 %; SnL/HeW 26.7–32.1 %; LNH/HeW 9.3–10.0 %; LNH/SL 1.2–1.4 %; RNW/HeW 7.7–10.0 %; RNW/SL 1.0–1.3 %; HLL/SL 18.0–19.7 %; FLL/SL 16.2–18.8 %; HaL/VGS 23.8–27.0 %; FoL/VGS 28.6–32.4 %; Haw/HeW 23.3–26.9 %; FoW/HeW 26.7–32.1 %; LT2/HeW 7.7–10.0 %; LT3/FoW 20.6–25.6 %; LT2/FoL 25.8–33.3 %; LF2/HaL 28.0–40.0 %; WT3/FoW 24.4–28.8 %; WF3/HaW 31.4%. Etymology. The specific epithet is a noun in apposition, and refers to the region where the holotype was collected, the upper catchment basin of the Vereh River. Habitat and natural history observations. Nototriton vereh has been found to inhabit moss growing on the ground and on the lower sections of tree trunks (within a meter above the ground). One specimen (UCR 23684) was found within a clump of moss that had fallen from a tree above; this moss had been on the ground for a while judging by its appearance, so it is highly unlikely that this individual had been living within the moss high up in the canopy. We feel that it is most likely that UCR 23684 was living within the terrestrial leaf litter and was taking diurnal refuge within the fallen moss clump when BK and AR discovered it. Two individuals (UCR 23682 and UCR 23683) were found by BK and AR during the day within moss growing on the ground in highly disturbed roadside habitats. The holotype (UCR 23681) was discovered by BK within a section of abandonded pasture in his private property referred to as the Rio Vereh Cloud Forest Reserve. The holotype was discovered during the day within a thick patch of moss (approximately 5 cm in thickness) growing on the lower trunk of a small tree, approximately 50 cm above the ground. Distribution. Nototriton vereh is endemic to Costa Rica, and currently only known to inhabit a very small section of cloud forest within the Tropical Premontane Rainforest life zone (Holdridge 1967) along the northern Caribbean foothills of the Cordillera de Talamanca. The known altitudinal range for N. vereh is from ca. 1250–1400 m a.s.l., Published as part of Kubicki, Brian, Reyes, Aura & Arias, Erick, 2022, Revised taxonomy and distributions of Costa Rican moss salamanders (Caudata: Plethodontidae: Nototriton), with descriptions of new taxa, pp. 451-496 in Zootaxa 5194 (4) on pages 470-475, DOI: 10.11646/zootaxa.5194.4.1, http://zenodo.org/record/7156853, {"references":["Taylor, E. H. (1949) New salamanders of Costa Rica. University of Kansas Science Bulletin, 33, 279 - 288. https: // doi. org / 10.5962 / bhl. part. 16130","Good, D. A. & Wake, D. B. (1993) Systematic studies of the Costa Rican moss salamanders, genus Nototriton, with descriptions of three new species. Herpetological Monographs, 7, 131 - 159. https: // doi. org / 10.2307 / 1466956","Holdridge, L. R. (1967) Life Zone Ecology. Revised edition. Tropical Science Center, San Jos, 206 pp."]}

Taylorotriton subgen. nov. Type species. Parvimolge richardi Taylor, 1949. Diagnosis. Diminutive salamanders with a standard length not known to exceed 27 mm. Possessing tiny hands and feet that are pad-like in appearance, with at the most the distal phalanx on each digit protruding in a bluntly pointed tip beyond the fleshy palmar and plantar tissue. Hand width not known to exceed 30% of head width. Relatively large nostril openings; width of nostril typically exceeding 5% of head width. The members of this subgenus are also diagnosed by evident divergence in their mtDNA sequences. Etymology. Taylorotriton, is formed from Taylor (a surname) and the Greek word Triton (a Greek god of the sea, son of Poseidon and Amphitrite) (Day 2007). Triton is a commonly used word for salamanders, and is a root in several genera and subgenera of bolitoglossine salamanders. We propose this subgeneric name in honor of Edward Harrison Taylor (1889–1978), the legendary alpha-taxonomist who greatly expanded the knowledge and understanding of Costa Rica’s amphibians, including salamanders, during his work in the country especially during middle decades (40s and 50s) of the 20th century. Composition. Only three species are recognized to comprise this subgenus, The two former members of the Nototriton richardi species group (N. richardi and N. tapanti) and a newly proposed species described herein below. Distribution. All three members of the subgenus Taylorotriton are known to be endemic to Costa Rica and restricted to the Caribbean slopes of the Central Volcanic Range and northeastern Talamanca, at elevations from roughly 1300–2250 masl (Fig. 6).

Nototriton (Nototriton) gamezi García-París & Wake, 2000 Monteverde moss salamander Nototriton gamezi García-París & Wake (2000: 55). Nototriton (Nototriton) gamezi: Dubois & Raffaëlli (2012: 141). Holotype. MVZ 207122, by original designation, an adult female from Costa Rica: Provincia de Alajuela: Cantón de San Ramón: Distrito de Peñas Blancas: Monteverde Cloud Forest Reserve: Carril Bosque Eterno at junction with Pantanosa Trail, 1600 m a.s.l. (N 10°19’, W 84°47.5’), collected by D. C. Cannatella, D. A. Good, W. Guindon, and D. B. Wake on 14 August 1987. Etymology. The specific epithet is a patronym, used as a noun in the masculine genitive case, in dedication to Mr. Rodrigo Gámez, the first director of the Instituto Nacional de Biodiversidad (García-París & Wake 2000). Distribution. Nototriton gamezi is currently only known to inhabit the Monteverde Cloud Forest Reserve between roughly 1500–1600 masl, but following further exploration in the surrounding mountains its range will likely extend beyond the reserve., Published as part of Kubicki, Brian, Reyes, Aura & Arias, Erick, 2022, Revised taxonomy and distributions of Costa Rican moss salamanders (Caudata: Plethodontidae: Nototriton), with descriptions of new taxa, pp. 451-496 in Zootaxa 5194 (4) on page 488, DOI: 10.11646/zootaxa.5194.4.1, http://zenodo.org/record/7156853, {"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. https: // doi. org / 10.1643 / 0045 - 8511 (2000) 2000 [0042: MPAORO] 2.0. CO; 2","Dubois, A. & Raffaelli, J. (2012) A new ergotaxonomy of the order Urodela Dumeril, 1805 (Amphibia, Batrachia). Alytes, 28, 77 - 161."]}

Nototriton (Nototriton) costaricense Arias & Kubicki, 2018 Southern moss salamander Holotype. UCR 22900, by original designation, a subadult from Costa Rica: Provincia de Limón: Cantón de Talamanca: Distrito de Telire: Cerro Pat, Parque Internacional La Amistad, ca. 1500 m a.s.l., obtained by Erick Arias on 11 March 2015. Etymology. The specific epithet refers to the Spanish word meaning Costa Rican, “costaricense”. The name represents the fact that the holotype and species was discovered in Costa Rica. Given the close proximity of the type locality to the Costa Rica-Panama border, we speculate that one day it may indeed be discovered within Panama as well (Arias & Kubicki 2018). Distribution. Nototriton costaricense is known from a single site within the Lower Montane Rain Forest life zone (Holdridge 1967) along the mid-elevation slopes of the extreme southeastern Cordillera de Talamanca, within Parque Internacional La Amistad, in the vicinity of Cerro Pat, ca. 1500 m (Arias & Kubicki 2018)., Published as part of Kubicki, Brian, Reyes, Aura & Arias, Erick, 2022, Revised taxonomy and distributions of Costa Rican moss salamanders (Caudata: Plethodontidae: Nototriton), with descriptions of new taxa, pp. 451-496 in Zootaxa 5194 (4) on page 488, DOI: 10.11646/zootaxa.5194.4.1, http://zenodo.org/record/7156853, {"references":["Arias, E. & Kubicki, B. (2018) A new moss salamander, genus Nototriton (Caudata: Plethodontidae), from the Cordillera de Talamanca, in the Costa Rica-Panama border region. Zootaxa, 4369 (4), 487 - 500. https: // doi. org / 10.11646 / zootaxa. 4369.4.2","Holdridge, L. R. (1967) Life Zone Ecology. Revised edition. Tropical Science Center, San Jos, 206 pp."]}

Nototriton maximo sp. nov. Maximo’s moss salamander Holotype. UCR 23689 (Fig. 17), an adult female from Costa Rica: Provincia de Guanacaste: Cantón de Tiláran: Distrito de Tronadora: edge of road approximately 3 kilometers NNW of the entrance to the Santa Elena Reserve, Monteverde, ca. 1425 m a.s.l., collected by Brian Kubicki in the company of Aura Reyes on 22 February 2013. Paratopotype. UCR 23690, same collection data as the holotype. Paratypes. UCR 23691 Provincia de Guanacaste: Cantón de Tiláran: Distrito de Tronadora: edge of road approximately 200 meters NNE of the entrance of the Santa Elena Reserve, Monteverde, ca. 1600 m a.s.l., collected by Brian Kubicki in the company of Aura Reyes on 26 April 2013. UCR 23692 same locality data as UCR 23691, but collected by Brian Kubicki in the company of Aura Reyes on 5 July 2016. UCR 23693, a subadult from Costa Rica: Provincia de Alajuela: Cantón de Upala: Distrito de Aguas Claras: moss on tree trunk about 3 meters above the ground, northeastern slope of the Rincon de la Vieja Volcano, Rincon de la Vieja National Park, ca. 1450 m a.s.l., collected by Brian Kubicki in the company of Aura Reyes on 17 December 2013. Generic Placement. Assigned to the genus Nototriton due to having fewer than 14 costal grooves, reduced manus and pes that are longer than wide, and the molecular evidence (16S, COI, and cyt b mtDNA distances) presented herein. Subgeneric Placement. Nototriton maximo is assigned to the subgenus Nototriton due to its known geographic distribution (endemic to Costa Rica), its small hands and feet with the majority of the lengths of fingers II, III, and IV and toes II, III, IV, and V being free of extended palmar or plantar tissue, and by the molecular evidence (16S, COI, and cyt b mtDNA distances) presented herein. Diagnosis. The combination of the following characteristics can be used to distinguish Nototriton maximo from the other described species of the subgenus Nototriton: (1) having small hands and feet, with at least the distal phalanges on the fingers II, III, and IV and toes toes II, III, IV, and V being free of the margin of the palmar and plantar tissue; (2) typically shorter front and hind limbs, HLL known to range from 4.1–5.2 mm and FLL known to range from 4.0– 4.8 mm; (3) 16S, COI, and cyt b mtDNA distances. Description of holotype. Adult female having a total length of 71.8 mm and SL of 31.9 mm. Head slightly wider than neck and shoulders (HeW 4.0 mm, NeW 3.7 mm, ShW 3.7 mm), with greatest width of head just posterior to the articulation of the jaws; snout raised anterodorsally, spadate to rounded in dorsal outline, and rounded to truncate in profile; snout relatively short (SnL 1.2 mm, 3.8 % of SL), with nearly terminal non-protruding small nostrils (LNH 0.12 mm, RNW 0.12 mm) directed anterolaterally; internarial area convex in dorsal outline. Eyes relatively large (EW = 133 % of SnL), protruding well beyond dorsal and ventral outline of head, directed anterolaterally, with a distinct suborbital groove. Top of head flat and smooth, tapering toward anterior terminus, lacking contrasting interorbital or other dermal structures. Canthus rostralis distinct; intercanthal area flat to slightly convex; and loreal region flat to slightly concave. No obvious cirri (nasolabial protuberances) present. Nasolabial grooves discernible on tip of snout, starting at lateral margins of nares and extending nearly vertically (with a slight outward angle) to a point just above upper lip margin. Gular fold well-defined, starting on dorsolateral portion of neck, below weakly discernible postorbital groove, wrapping around posteriolateral section of head at a slightly anterior angle and crossing venter as a smooth and weak anterior-oriented curve. Nuchal grooves very weak to indiscernible. Postorbital grooves very weak, nearly indiscernible, starting at the posterior corner of eyes and traveling horizontally along the inferior margin of parotoid glands, terminating at gular fold on the lateral portion of neck. Horizontal mandibular grooves nearly indiscernible, starting at corners of mouth and extending horizontally to vertical mandibular groove. Vertical mandibular grooves discernible, but weak starting along inferior margins of postorbital grooves and extending vertically to gular region, terminating at junction with inner mandibular grooves. Inner mandibular grooves evident, running medially parallel to mandibular bones, becoming indiscernible near anterior tip of lower jaw. Pair of nearly indiscernible and weakly raised parotoid glands present on dorsolateral margin of head among orbits, postorbital grooves, and occiput. Snout not evidently protruding beyond anterior margin of lower lip in lateral view. No mental gland visible under skin of anterior intermandibular region. Arms relatively long and slender overall (FLL = 4.8 mm, 15.0 % of SL), without noticeable hypertrophied forearm compared to upper arm. Hands very small and slender (HaL = 1.7 mm, 32.1 % of VGS; HaW = 1.6 mm, 40.0 % of HeW). Fingers II, III, and IV long and slender, protruding freely, with at least ultimate and penultimate phalanges being free beyond interdigital tissue margin (LF2 0.71 mm, LF3 0.71 mm); a weak, but evident indentation at interdgitial space between fingers I and II. Tips of fingers rounded, with terminal pads discernible on ventral surfaces, especially fingers II and III. Palmar surfaces appearing to be smooth overall. Dorsal surfaces of hands with discernible interdigital grooves that start at interdigital tissue margins and cross onto metacarpal region. Relative lengths of fingers on right hand I Legs relatively long and slender overall (HLL 5.2 mm, 16.3 % of SL), with lower leg being very slightly thicker than upper leg. Feet very small and slender (FoL 2.1 mm, 39.6 % of VGS; FoW 2.0 mm, 52.5 % of HeW). Toes II, III, IV and V protruding freely, with at least ultimate (Toe V) and penultimate (toes II, III, and IV) phalanges being free beyond interdigital tissue margin (LT2 0.79 mm, LT3 0.68 mm); very minimal to indiscernible indentation at interdgitial space between toes I and II. Tips of toes rounded, with terminal pads discernible on ventral surfaces. Plantar surfaces appearing to be smooth overall, without any evident dermal creases. Dorsal surfaces of feet with discernible interdigital grooves, especially between toes II-III, III-IV, and IV-V that start at interdigital tissue margins and cross onto metatarsal region. Relative lengths of toes on right foot I Body subcylindrical (slightly wider than high) in cross section, and relatively robust (TW = 4.4 mm; TW = 22.8 % of AGL). Between axilla and groin, 11 costal grooves visible, 13 if counting axillary and inguinal grooves; costal grooves most visible on ventral and lateral portions of body. Adpressed limbs separated by six costal folds; 12 costal folds total between axilla and groin. Slight middorsal depression extends longitudinally along length of body, starting at base of head (occiput) and becoming indiscernible on anterior portion of tail. Tail long, 39.9 mm in length, cylindrical in cross section, having a very slight constriction at base, and evenly tapering to pointed terminus; in life, no caudal grooves were discernible. Skin on surfaces of head, body, limbs, and tail smooth. Coloration in life. The ground color of the dorsal and lateral surfaces of the head and trunk reddish-orange. There were several pale fleshy colored dash-like markings on the lateral and dorsolateral surface of the body.Additionally, there were some scattered dark brown to black small irregular markings and tiny white spots on the dorsal and lateral surfaces of the head and body. The iris was dark reddish-orange with dark brownish-black reticulation. The upper surfaces of the arms and legs were very similar in overall coloration and pattern to that of the abovementioned chromatic characteristics of the dorsal and lateral surfaces of the head and body. The dorsal and dorsolateral surfaces of the tail were uniform bright reddish-orange. The ventrolateral surfaces of the head, limbs, body, and tail were contrasted from the superior surfaces by consisting of a concentration of fine white or pale fleshy-orange markings on a brown to brownish-orange background. The area directly surrounding the cloaca had a concentration of bright orange chromatophores. Coloration in ethanol. After more than eight years in ethanol (70%), the overall ground coloration of the holotype has shifted to a pale fleshy tan dorsally. Throughtout the dorsal and dorsolateral surfaces of the head and body there are also numerous fine to medium-sized darker markings and spots; on the dorsolateral section of each side of the trunk, there are a series of darker dash-like markings. The ventral surfaces of the head, body, limbs, and tail are darker brownish-gray with very small pale patches scattered throughout. Measurements (in mm), limb interval, and percentages of the holotype. SL 31.9; total length 71.8; tail length 39.9; ShW 3.7; HeW 4.0; NeW 3.7; EW 1.6; SnL 1.2; JSL 3.7; LGFS 6.4; LNH 0.12; RNW 0.12; IND 1.0; NLP 0.5; ICD 1.8; HLL 5.2; FLL 4.8; TW 4.4; VGS 5.3; FSL 8.0; UHL 3.6; AGL 19.3; VL 1.9; HaW 1.6; HaL 1.7; LF2 0.71; LF3 0.71; WF3 0.34; FoW 2.0; FoL 2.1; LT2 0.79; LT3 0.68; WT3 0.37. Limb interval six. Measurements in related percentages: VGS/SL 16.6 %; IND/HeW 25.0 %; AGL/SL 60.5 %; HeW/SL 12.5 %; Hew/AGL 20.7 %; SnL/ HeW 30.0 %; LNH/HeW 3.0 %; LNH/SL 0.38 %; RNW/HeW 3.0 %; RNW/SL 0.38 %; HLL/SL 16.3 %; FLL/SL 15.0 %; HaL/VGS 32.1 %; FoL/VGS 39.6 %; Haw/HeW 40.0 %; FoW/HeW 52.5 %; LT2/FoL 37.6 %; LF2/HaL 41.8 %; WT3/FoW 18.5 %; WF3/HaW 21.3 %. Noteworthy variation. The paratype (UCR 23693) has discernible differences in the overall structure of the hands and feet, which are especially evident while comparing the digits to those of other members of the type series. The digits on the hands and feet of UCR 23693 are shorter and more tapered distally than on the other type series specimens; this could be related to ontogenesis, given the smaller size (22.0 mm) of UCR 23693, but UCR 23690, with a SL of 22.5 mm, has clearly visible differences in the structure of the digits from UCR 23693. The overall digit structure of UCR 23690 is similar to that of UCR 23689, UCR 23691, and UCR 23692. Measurements (in mm), limb intervals, and percentages of the paratopotypes. SL 22.0–28.5; ShW 2.6– 3.4; HeW 3.1–4.0; NeW 2.9–3.5; EW 1.1–1.3; SnL 0.8–1.1; JSL 2.7–3.4; LGFS 4.8–5.7; LNH 0.15–0.25; RNW 0.09–0.19; IND 0.7–1.0; NLP 0.3–0.6; ICD 1.6–1.8; HLL 4.1–5.2; FLL 4.0–4.5; TW 2.8–3.9; VGS 4.0–5.2; FSL 6.2–7.5; UHL 2.5–3.4; AGL 12.5–16.9; VL 0.9–1.4; HaW 1.0–1.4; HaL 1.0–1.5; LF2 0.47–0.68; LF3 0.43–0.68; WF3 0.22–0.31; FoW 1.3–1.7; FoL 1.4–2.0; LT2 0.53–0.76; LT3 0.4–0.71; WT3 0.25–0.34. Limb interval five–six. Measurements in related percentages: VGS/SL 18.2–19.1 %; IND/HeW 20.0–25.0 %; AGL/SL 56.8–59.3 %; HeW/ SL 13.4–14.2 %; Hew/AGL 23.3–24.8 %; SnL/HeW 25.8–28.6 %; LNH/HeW 4.0–7.8 %; LNH/SL 0.56–1.11 %; RNW/HeW 2.6–5.0 %; RNW/SL 0.34–.0.71 %; HLL/SL 16.8–19.5 %; FLL/SL 15.8–18.2 %; HaL/VGS 25.0–30.2 %; FoL/VGS 34.9–39.6 %; Haw/HeW 31.3–35.5 %; FoW/HeW 40.6–42.9 %; LT2/FoL 37.9–42.7 %; LF2/HaL 41.3–47.0 %; WT3/FoW 19.2–21.5 %; WF3/HaW 20.0–28.0 %. Etymology. The specific epithet “ maximo ” is a patronym, used as a noun apposition. This taxon is named in honor of Maximiliano “Maximo” Flores (1964–2016), who was a good friend and neighbor of BK and AR. Maximo’s friendship and help with the efforts of the Costa Rican Amphibian Research Center and at the Guayacán Rainforest Reserve, another private property owned by BK, will always be very much appreciated. Maximo was always very willing to join BK during both diurnal and nocturnal field outings, and he accompanied BK and AR on numerous trips throughout the cloud forest regions of Costa Rica searching for moss salamanders. Habitat and natural history observations. Nototriton maximo has been found within moss growing on the ground, fallen and standing tree trunks and accessable branches of trees. Distribution. Nototriton maximo is endemic to Costa Rica, and currently only known to inhabit two very small distinct cloud forest regions in northwestern Costa Rica. The first region is along the continental divide just north and northwest of the Monteverde Cloud Forest Reserve. The second region is along the northern slopes of the Rincón de la Vieja Volcano. The known altitudinal range for N. maximo is from ca. 1425–1600 masl.

Nototriton (Taylorotriton) richardi (Taylor, 1949) Richard’s moss salamander Parvimolge richardi Taylor (1949: 284) Chiropterotriton richardi Wake & Lynch (1976: 60) Nototriton richardi Wake & Elias (1983: 11) Nototrtion (Nototriton) richardi Dubois & Raffaëlli (2012: 141) Holotype. RCT 1436 (Richard C. Taylor field number), by original designation, but currently housed at the Chicago Field Museum of Natural History, FMNH 178295, an adult female from Costa Rica: Provincia de Alajuela: Cantón de Alajuela: Distrito de Sarapiquí: Isla Bonita: American Cinchona Plantation, ca. 6500 feet (ca. 1980 m a.s.l.), collected by Richard Clark Taylor on 1 August 1947. Etymology. The specific epithet is a patronym, used as a noun in the masculine genitive case, in dedication to Richard Clark Taylor (1926–2002), the son of Edward Harrison Taylor (1889–1978). Richard Taylor joined his father while searching for reptiles and amphibians during an expedition to Costa Rica in 1947, and collected the hototype of this taxon. Distribution. Nototriton richardi is endemic to Costa Rica and is only known to inhabit the Caribbean slopes of the Cordillera Volcanica Central at elvations from roughly 1300–1800 m a.s.l. (Savage 2002). Specimens have been collected from within a swath ranging from the northwestern slopes of the Irazu Volcano to the eastern flanks of Juan Castro Blanco National Park., Published as part of Kubicki, Brian, Reyes, Aura & Arias, Erick, 2022, Revised taxonomy and distributions of Costa Rican moss salamanders (Caudata: Plethodontidae: Nototriton), with descriptions of new taxa, pp. 451-496 in Zootaxa 5194 (4) on page 475, DOI: 10.11646/zootaxa.5194.4.1, http://zenodo.org/record/7156853, {"references":["Taylor, E. H. (1949) New salamanders of Costa Rica. University of Kansas Science Bulletin, 33, 279 - 288. https: // doi. org / 10.5962 / bhl. part. 16130","Wake, D. B. & Lynch, J. F. (1976) The distribution, ecology, and evolutionary history of plethodontid salamanders in tropical America. Scientific Bulletin Natural History Museum of Los Angeles Country, 25, 1 - 65.","Wake, D. B. & Elias, P. (1983) New genera and a new species of Central American salamanders, with a review of the tropical genera (Amphibia, Caudata, Plethodontidae). Contributions in Science. Natural History Museum of Los Angeles County, 345, 1 - 19. https: // doi. org / 10.5962 / p. 208170","Dubois, A. & Raffaelli, J. (2012) A new ergotaxonomy of the order Urodela Dumeril, 1805 (Amphibia, Batrachia). Alytes, 28, 77 - 161.","Savage, J. M. (2002) The Amphibians and Reptiles of Costa Rica: A Herpetofauna between Two Continents, between Two Seas. The University of Chicago Press, Chicago, Illinois, 934 pp."]}

Nototriton lateomuscus sp. nov. San Ramon moss salamander Holotype. UCR 23694 (Figs. 14–16), an adult female from Costa Rica: Provincia de Alajuela: Cantón de San Ramón: Distrito de Angeles: Edge of road, approximately 3 kilometers west of Balsa, ca. 1200 m a.s.l., collected by Brian Kubicki in the company of Aura Reyes on 21 October 2016. Paratopotypes. UCR 23695, same collection data as the holotype. UCR 23696, same locality data as the holotype, but collected by Brian Kubicki in the company of Aura Reyes on 12 October 2013. Generic Placement. Assigned to the genus Nototriton due to having fewer than 14 costal grooves, reduced manus and pes that are longer than wide, and the molecular evidence (16S, COI, and cyt b mtDNA distances) presented herein. Subgeneric Placement. Nototriton lateomuscus is assigned to the subgenus Nototriton due to its known geographic distribution (endemic to Costa Rica), its small hands and feet with the majority of the lengths of fingers II, III, and IV and toes II, III, IV, and V being free of extended palmar or plantar tissue, and by the molecular evidence (16S, COI, and cyt b mtDNA distances) presented herein. Diagnosis. Nototriton lateomuscus is distinguished from the other described species of the subgenus Nototriton by its16S, COI, and cyt b mtDNA distances. Comparisons. Since the molecular evidence presented herein strongly supports N. lateomuscus forming part of clade with N. abscondens and N. gamezi, and due to the fact that all three of these species have adjoining distributions, morphological comparisons are only going to be focused on distinguishing N. abscondens and N. gamezi from N. lateomuscus. Contrasting characteristics for Nototriton lateomuscus are presented in parentheses. Nototriton abscondens (Taylor, 1948) can be distinguished from N. lateomuscus by having relatively smaller nostril openings, average LNH = 0.16 mm and RNW = 0.16 (average LNH = 0.21 mm and RNH = 0.2 mm); wider internarial distance, average IND = 0.9 mm (average IND = 0.7 mm). Nototriton gamezi (García-París & Wake, 2000) has larger nostril openings, average LNH = 0.29 mm and RNH = 0.24 mm (average LNH = 0.21 mm and RNH = 0.2 mm). Description of holotype. Adult female having a total length of 66.2 mm and SL of 29.3 mm. Head slightly wider than neck and shoulders (HeW 4.0 mm, NeW 3.3 mm, ShW 3.4 mm), with greatest width of head just posterior to the articulation of the jaws; snout raised anterodorsally, spadate to rounded in dorsal outline, and rounded in profile; snout relatively short (SnL 1.0 mm, 3.4 % of SL), with nearly terminal non-protruding large nostrils (LNH 0.25 mm, RNW 0.25 mm) directed anterolaterally; internarial area convex in dorsal outline. Eyes relatively large (EW = 130 % of SnL), protruding beyond dorsal outline of head, directed anterolaterally, with a distinct suborbital groove. Top of head flat and smooth, tapering slightly toward anterior terminus, lacking contrasting interorbital or other dermal structures. Canthus rostralis rounded; intercanthal area flat to slightly convex; and loreal region flat to slightly concave. No obvious cirri (nasolabial protuberances) present, but nasolabial grooves weakly discernible on tip of snout; nasolabial grooves start at lateral margins of nares and extend ventrally with a slight outward orientation; nasolabial grooves start at lateral margins of nares and extend ventrally at first, but take a drastic backwards curve about halfway to lip; nasolabial grooves terminate just prior to reaching margin of upper lip. Gular fold welldefined, starting on lateral portion of neck, below postorbital groove, wrapping around posterolateral section of head at a slightly anterior angle and crossing venter as a smooth and anterior-oriented curve. Nuchal grooves very weak to indiscernible. Postorbital grooves discernible, starting at posterior corner of eyes and traveling horizontally along the inferior margin of parotoid glands, terminating at gular fold on the lateral portion of neck. Horizontal mandibular grooves indiscernible, but vertical mandibular groove well-defined.Vertical mandibular grooves starting at inferior margins of postorbital grooves and extending vertically to and crossing onto gular region, but becoming indiscernible just medially of inner mandibular grooves. Inner mandibular grooves evident, nearly forming a single curving structure, but become indiscernible at anterior portion of gular patch. Pair of very weakly but discernibly raised parotoid glands present on dorsolateral margin of head between orbits, postorbital grooves, and occiput. Upper lip protruding slightly beyond edge of lower lip in ventral and lateral views; no mental gland discernible under skin of anterior intermandibular region. Arms relatively long and slender overall (FLL = 4.9 mm, 16.7 % of SL), without noticeable hypertrophied forearm compared to upper arm. Hands small and slender (HaL = 1.6 mm, 29.1 % of VGS; HaW = 1.4 mm, 25 % of HeW). Fingers II, III, and IV protruding freely, with at least ultimate and all to part of penultimate phalanges being free beyond interdigital tissue margin (LF2 0.74 mm, LF3 0.56 mm). Finger I with minimal indentation at interdgitial space. Tips of fingers rounded, with terminal pads discernible on ventral surfaces. Palmar surfaces appearing to be smooth overall, but with weakly discernible depressions radiating out from palmar section to interdigital margins between fingers. Dorsal surfaces of hands with discernible interdigital depressions or grooves, especially between fingers II-III and III-IV, that start at interdigital tissue margins and cross onto metacarpal region. Relative lengths of fingers on right hand I Coloration in life. The ground color of the dorsal and lateral surfaces of the head and trunk consisted of a brownish-orange coloration. Scattered throughout the dorsal and lateral surfaces of the head and trunk there were numerous dark brownish-black markings of irregular size and shape, in addition to numerous very fine white chromatophores. The parotoid glands where a contrasted paler tone compared to the surrounding dorsal surfaces. The iris was bright copper-orange with a black reticulation. The upper surfaces of the arms and legs are very similar in coloration and pattern to that of the above-mentioned chromatic characteristics of the dorsal and lateral surfaces of the head and body. The dorsal and dorsolateral surfaces of the anterior third of the tail were a paler shade of reddish-orange, but the posterior two-thirds of the tail was similar to dorsal surfaces of the trunk. There were also some very fine white chromatophores scattered randomly throughout the tail, but at a much lesser concentration compared to the body and head. The ventral and ventrolateral surfaces of the head, limbs, and body were contrasted from the superior surfaces by consisting of a concentration of fine white chromatophores on a darker brownish-gray background. The gular patch was slightly more pale in comparison to the ventral surfaces of the trunk. The palmar and plantar surfaces were pale grayish-brown, lacking evident chromatophores. Directly surrounding the cloaca there was a concentration of bright orange chromatophores. The anterior third of the vental surface of the tail had an overall similar pattern and coloration to that of the ventral surface of the trunk. The ventral surface along the posterior two-thirds of the tail had a dark reddish orange ground color with numerous dark brown markings of irregular size and shape in addition to a relatively large concentration of very fine white chromatophores. Coloration in ethanol. After more than a year and a half in ethanol (70%), the overall coloration of the holotype has changed to principally consisting of a mixture of tan and brown mottling. Two pale tannish-yellow patches are visible on the posterior dorsolateral surfaces of the head, where the parotoid glands are located. The dorsal surface of the anterior third of the tail is also pale tannish-yellow. The ventral surfaces of the body and tail are dark brownishgray. The ventral surface of the head has an evident paler tone compared to that of the rest of the body and tail. Measurements (in mm), limb interval, and percentages of the holotype. SL 29.3; total length 66.2; tail length 36.9; ShW 3.4; HeW 4.0; NeW 3.3; EW 1.3; SnL 1.0; JSL 3.4; LGFS 6.0; LNH 0.25; RNW 0.25; IND 0.8; NLP 0.5; ICD 1.6; HLL 5.9; FLL 4.9; TW 3.3; VGS 5.5; FSL 8.0; UHL 3.3; AGL 16.7; VL 1.9; HaW 1.4; HaL 1.6; LF2 0.74; LF3 0.56; WF3 0.31; FoW 1.6; FoL 2.1; LT2 0.9; LT3 0.68; WT3 0.31. Limb interval 4.5. Measurements in related percentages: VGS/SL 18.8 %; IND/HeW 20.0 %; AGL/SL 57.0 %; HeW/SL 13.7 %; Hew/AGL 24.0 %; SnL/ HeW 25.0 %; LNH/HeW 6.3 %; LNH/SL 0.85 %; RNW/HeW 6.3 %; RNW/SL 0.85 %; HLL/SL 20.1 %; FLL/SL 16.7 %; HaL/VGS 29.1 %; FoL/VGS 38.2 %; Haw/HeW 35.0 %; FoW/HeW 40.0 %; LT2/FoL 42.9 %; LF2/HaL 46.3 %; WT3/FoW 19.4 %; WF3/HaW 22.1 %. Measurements (in mm), limb intervals, and percentages of the paratopotypes. SL 19.2–26.6; ShW 2.5–3.0; HeW 3.0–3.3; NeW 2.5–3.1; EW 1.2–1.3; SnL 0.6–1.0; JSL 2.4–3.3; LGFS 4.5–5.7; LNH 0.12–0.25; RNW 0.14– 0.22; IND 0.7; NLP 0.3–0.4; ICD 1.4–1.7; HLL 4.5–4.9; FLL 3.9–4.6; TW 2.5–3.4; VGS 3.9–4.9; FSL 5.7–7.5; UHL 2.4–3.2; AGL 11.4–15.1; VL 1.6; HaW 0.9–1.1; HaL 1.1–1.2; LF2 0.37–0.57; LF3 0.34–0.4; WF3 0.22–0.23; FoW 1.1–1.5; FoL 1.4–1.6; LT2 0.53–0.65; LT3 0.37–0.56; WT3 0.25. Limb interval 4–6. Measurements in related percentages: VGS/SL 18.4–20.3 %; IND/HeW 21.2–23.3 %; AGL/SL 56.8–59.4 %; HeW/SL 12.4–15.6 %; Hew/ AGL 21.9–26.3 %; SnL/HeW 20.0–30.3 %; LNH/HeW 3.6–8.3 %; LNH/SL 0.45–1.3 %; RNW/HeW 4.2–7.3 %; RNW/SL 0.53–1.1 %; HLL/SL 18.4–23.4 %; FLL/SL 17.3–25.5 %; HaL/VGS 24.5–28.2 %; FoL/VGS 32.7–35.9 %; Haw/HeW 30.0–33.3 %; FoW/HeW 36.7–45.5 %; LT2/FoL 37.9–40.6 %; LF2/HaL 33.6–47.5 %; WT3/FoW 16.7–22.7 %; WF3/HaW 18.2–25.6 %. Etymology. The specific epithet is formed from the Latin words lateo, which means to lie hidden, and muscus, which means moss. This taxon is named due to the fact that it often “lies hidden” within moss. Habitat and natural history observations. Nototriton lateomuscus is known only from three specimens that have been found exclusively within moss growing on the trunks and branches of trees. Distribution. Nototriton lateomuscus is endemic to Costa Rica, and currently only known to inhabit a single site on the extreme eastern margin of the Tilaran mountains (ca. 1200 masl), north of the city of San Ramon., Published as part of Kubicki, Brian, Reyes, Aura & Arias, Erick, 2022, Revised taxonomy and distributions of Costa Rican moss salamanders (Caudata: Plethodontidae: Nototriton), with descriptions of new taxa, pp. 451-496 in Zootaxa 5194 (4) on pages 481-485, DOI: 10.11646/zootaxa.5194.4.1, http://zenodo.org/record/7156853, {"references":["Taylor, E. H. (1948) New Costa Rican salamanders. Proceedings of Biological Society of Washington, 61, 177 - 180.","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. https: // doi. org / 10.1643 / 0045 - 8511 (2000) 2000 [0042: MPAORO] 2.0. CO; 2"]}

Subgenus Nototriton Dubois & Raffaëlli, 2012 Type species. Spelerpes picadoi Stejneger, 1911. Diagnosis. Diminutive to small salamanders with a standard length not known to exceed 38 mm, but with adult sizes typically exceeding 25 mm. Possessing small hands and feet that are longer than wide, with the hand width not known to exceed 2.0 mm and foot width not known to exceed 2.5 mm. Digits of hands and feet, excluding Finger I and Toe I, free of the palmar and plantar tissue for the majority of their lengths, with most digits having the ultimate and parts of the penultimate phalanges free of interdigital tissue. The members of this subgenus are also diagnosed by evident divergence in their mtDNA sequences. Etymology. Nototriton, is formed from the Greek words notos (meaning south) and Triton, the name of a Greek sea-god. The term “triton” is commonly used word for salamanders, and is a root in several genera and subgenera of bolitoglossine salamanders (Wake & Elias 1983). Composition. We recognize nine species to comprise this subgenus; the following six formerly recognized members of the Nototriton picadoi species group (N. abscondens, N. costaricense, N. gamezi, N. guanacaste, N. picadoi, and N. saslaya) and three newly proposed taxa described herein below. With the addition of fresh material collected by BK from near the type locality of Nototriton major that strongly agreed morphologically with the diagnositic characteristics of that taxon (Good and Wake 1993) we were able make a detailed taxonomic review of this species. Nototriton major was previously known from just a single specimen ever collected, the holotype, and no genetic material was available until now for analysis. Following our comparison of the morphological characteristics, genetics, and geographic proximity of known sites, we concluded that N. major can best be recognized as a junior synonym of Nototriton picadoi. Additionally, following our detailed review of the situation regarding Nototriton matama, in similar manner, we propose that this taxon also be recognized as a junior synonym of N. picadoi, due to its very low genetic distance and weakly supported morphological diagnostics that are more of a result of poorly preserved specimens than actual characteristics present on living or well preserved individuals. Distribution. The nine members of the subgenus Nototriton are known to have a disjunct distribution, with N. saslaya only being known from two high-elevation sites in northern Nicaragua, whereas the remaining eight species are known to be restricted to the Caribbean slopes of Costa Rica. The members of this subgenus are known from elevations ranging between roughly 750–2500 masl (Fig. 10).

Nototriton (Nototriton) abscondens (Taylor, 1948) Concealed moss salamander Chiropterotriton abscondens Taylor (1948: 177). Nototriton abscondens: Good & Wake (1993: 139). Nototriton (Nototriton) abscondens: Dubois & Raffaëlli (2012: 141). Holotype. RCT 1414 (Richard C. Taylor field number), by original designation, but currently housed at the Chicago Field Museum, FMNH 178285. An adult female from Costa Rica: Provincia de Alajuela: Cantón de Alajuela: Distrito de Sarapiquí: Isla Bonita: American Cinchona Plantation, ca. 5500 feet (ca. 1675 m a.s.l.), collected by Richard Clark Taylor on 1 August, 1947. Etymology. The specific epithet comes from the Latin prefix ab - (meaning from or away) and condo (meaning hidden). Distribution. Nototriton abscondens is endemic to Costa Rica and is only known to inhabit the Caribbean slopes of the Cordillera Volcánica Central at elvations from roughly 1000–2500 masl (Savage 2002). Specimens have been collected from within a swath ranging from the northern slopes of the Turrialba Volcano to the western flanks of Juan Castro Blanco National Park., Published as part of Kubicki, Brian, Reyes, Aura & Arias, Erick, 2022, Revised taxonomy and distributions of Costa Rican moss salamanders (Caudata: Plethodontidae: Nototriton), with descriptions of new taxa, pp. 451-496 in Zootaxa 5194 (4) on page 488, DOI: 10.11646/zootaxa.5194.4.1, http://zenodo.org/record/7156853, {"references":["Taylor, E. H. (1948) New Costa Rican salamanders. Proceedings of Biological Society of Washington, 61, 177 - 180.","Good, D. A. & Wake, D. B. (1993) Systematic studies of the Costa Rican moss salamanders, genus Nototriton, with descriptions of three new species. Herpetological Monographs, 7, 131 - 159. https: // doi. org / 10.2307 / 1466956","Dubois, A. & Raffaelli, J. (2012) A new ergotaxonomy of the order Urodela Dumeril, 1805 (Amphibia, Batrachia). Alytes, 28, 77 - 161.","Savage, J. M. (2002) The Amphibians and Reptiles of Costa Rica: A Herpetofauna between Two Continents, between Two Seas. The University of Chicago Press, Chicago, Illinois, 934 pp."]}

Nototriton (Nototriton) guanacaste Good & Wake, 1993 Guanacaste moss salamander Nototriton guanacaste Good & Wake (1993: 138). Nototriton (Nototriton) guanacaste: Dubois & Raffaëlli (2012: 141). Holotype. MVZ 207111, by original designation, an adult male from Costa Rica: Provincia de Guanacaste: Cantón de Liberia: Distrito de Mayorga: Guanacaste National Park: upper slopes of the Cacao Volcano, 1580 m a.s.l., collected by D. C. Cannattela and D. A. Good on 24 August 1987 (Good & Wake 1993). Etymology. The specific epithet is a noun in apposition and was chosen in celebration of establishment of Guanacaste National Park, where this species is known to be restricted (Good & Wake 1993). Distribution. Nototriton guanacaste is currently only known to inhabit two small patches of cloud forest within Guanacaste National Park, both located on the upper slopes of the Orosi and Cacao volcanoes between roughly 1400–1650 masl (Savage 2002)., Published as part of Kubicki, Brian, Reyes, Aura & Arias, Erick, 2022, Revised taxonomy and distributions of Costa Rican moss salamanders (Caudata: Plethodontidae: Nototriton), with descriptions of new taxa, pp. 451-496 in Zootaxa 5194 (4) on page 489, DOI: 10.11646/zootaxa.5194.4.1, http://zenodo.org/record/7156853, {"references":["Good, D. A. & Wake, D. B. (1993) Systematic studies of the Costa Rican moss salamanders, genus Nototriton, with descriptions of three new species. Herpetological Monographs, 7, 131 - 159. https: // doi. org / 10.2307 / 1466956","Dubois, A. & Raffaelli, J. (2012) A new ergotaxonomy of the order Urodela Dumeril, 1805 (Amphibia, Batrachia). Alytes, 28, 77 - 161.","Savage, J. M. (2002) The Amphibians and Reptiles of Costa Rica: A Herpetofauna between Two Continents, between Two Seas. The University of Chicago Press, Chicago, Illinois, 934 pp."]}

Nototriton (Nototriton) picadoi (Stejneger, 1911) Picado’s moss salamander Spelerpes picadoi Stejneger (1911: 285). Oedipus picadoi: Dunn (1924: 99). ? Pseudoeurycea picadoi: Taylor (1949: 279). Chiropterotriton picadoi: Wake & Lynch (1976: 59). Nototriton picadoi: Wake & Elias (1983: 11). Nototriton major Good & Wake (1993: 137). Nototriton matama Boza-Oviedo et al. (2012: 48). Nototriton (Nototriton) picadoi: Dubois and Raffaëlli (2012: 141). Holotype. USNM 48280, by original designation, gender not provided in the description: Provincia de Cartago: Cantón de El Guarco: Distrito de San Isidro: La Estrella, ca. 2000 m a.s.l. (Picado 1913), Collected by Clodomido Picado (Stejneger 1911). Etymology. The specific epithet is a patronym, used as a noun in the masculine genitive case, in dedication to Dr. Clodomiro Picado Twight (1887–1944), who was a very distinguished Costa Rican scientist and the person that collected the holotype of this taxon. Distribution. Nototriton picadoi is endemic to Costa Rica and is only known to inhabit the Caribbean slopes of extreme eastern portion of the Cordillera Volcánica Central, specifically along the eastern flank of the Turrialba Volcano into the northern and eastern sections of the Cordillera de Talamanca. The reported altitudinal range for this taxon is from 1200–2200 masl (Savage 2002), but during this study the lower limit was discovered to extend down to 900 masl (BK personal observation). Noteworthy observations or remarks. Due to the very low genetic distances, adjoining distributional ranges, and weakly supported morphological diagnostics, we herein recognized N. major (Good & Wake, 1993) and N. matama (Boza-Oviedo et al., 2012) as junior synonyms under N. picadoi., Published as part of Kubicki, Brian, Reyes, Aura & Arias, Erick, 2022, Revised taxonomy and distributions of Costa Rican moss salamanders (Caudata: Plethodontidae: Nototriton), with descriptions of new taxa, pp. 451-496 in Zootaxa 5194 (4) on page 489, DOI: 10.11646/zootaxa.5194.4.1, http://zenodo.org/record/7156853, {"references":["Stejneger, L. H. (1911) Descriptions of three new batrachians from Costa Rica and Panama. Proceedings of the United States National Museum, 41, 285 - 288. https: // doi. org / 10.5479 / si. 00963801.1857.285","Dunn, E. R. (1924) New salamanders of the genus Oedipus with a synoptical key. Field Museum Natural History, Zoological Series, 12 (7), 95 - 100.","Taylor, E. H. (1949) New salamanders of Costa Rica. University of Kansas Science Bulletin, 33, 279 - 288. https: // doi. org / 10.5962 / bhl. part. 16130","Wake, D. B. & Lynch, J. F. (1976) The distribution, ecology, and evolutionary history of plethodontid salamanders in tropical America. Scientific Bulletin Natural History Museum of Los Angeles Country, 25, 1 - 65.","Wake, D. B. & Elias, P. (1983) New genera and a new species of Central American salamanders, with a review of the tropical genera (Amphibia, Caudata, Plethodontidae). Contributions in Science. Natural History Museum of Los Angeles County, 345, 1 - 19. https: // doi. org / 10.5962 / p. 208170","Good, D. A. & Wake, D. B. (1993) Systematic studies of the Costa Rican moss salamanders, genus Nototriton, with descriptions of three new species. Herpetological Monographs, 7, 131 - 159. https: // doi. org / 10.2307 / 1466956","Boza-Oviedo, E., Rovito, S. M., Chaves, G., Garcia-Rodriguez, A., Artavia, L. G., Bolanos, F. & Wake, D. 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. Zootaxa, 3309 (1), 36 - 61. https: // doi. org / 10.11646 / zootaxa. 3309.1.2","Dubois, A. & Raffaelli, J. (2012) A new ergotaxonomy of the order Urodela Dumeril, 1805 (Amphibia, Batrachia). Alytes, 28, 77 - 161.","Savage, J. M. (2002) The Amphibians and Reptiles of Costa Rica: A Herpetofauna between Two Continents, between Two Seas. The University of Chicago Press, Chicago, Illinois, 934 pp."]}

Nototriton costaricense sp. nov. Southern Moss Salamander (Figures 3���5) Holotype. UCR 22900, a subadult from Costa Rica: Provincia de Lim��n: Cant��n de Talamanca: Distrito de Telire: Cerro Pat, Parque Internacional La Amistad, ca. 1500 m a.s.l., obtained by Erick Arias on 11 March 2015. Generic Placement. Assigned to the genus Nototriton due to having fewer than 14 costal grooves and reduced manus and pes that are longer than wide, and to the subgenus Nototriton based the molecular evidence presented herein. Diagnosis. The combination of the following characteristics can be used to distinguish Nototriton costaricense from other described species in the subgenus Nototriton: (1) having at least the distal phalanges on fingers II, III, and IV in addition to the distal phalanges on toes II, III, IV, and V free of webbing; (2) large nostril openings, greater than 0.3 mm in diameter (Fig. 5); (3) robust habitus, TW = 14.6 % of SL; (4) long hind limbs, HLL = 21.9 % of SL; (4) differentiated from other members of the genus Nototriton, subgenus Nototriton by its 16S, COI, and cyt b mtDNA distances. Comparisons. Since Nototriton costaricense is only known to occur in Costa Rica and molecular evidence strongly supports it forming part of the Nototriton picadoi species group, phenotypic comparisons are presented here only with respect to members of that clade (i.e., N. abscondens, N. gamezi, N. guanacaste, N. major, N. matama, and N. picadoi), which are all endemic to Costa Rica (Boza-Oviedo et al. 2012; Townsend 2016). Contrasting characteristics for Nototriton costaricense are presented in parentheses. Nototriton abscondens (Taylor, 1948) can be distinguished from N. costaricense by having relatively smaller nostrils openings, 0.3���0.8 % of SL (RNW = 1.3 % of SL); relatively narrower head, 12���14 % of SL (broader head, HeW = 15.1 % of SL); slender trunk, TW 8���10 % of SL (thick trunk, TW 14.6 % of SL). Nototriton gamezi Good & Wake, 1993 has relatively shorter hind limbs, 16���17 % of SL (longer hind limb, HLL = 21.9 % of SL); narrower head, 13���14 % of SL (broader head, HeW = 15.1 % of SL); slender trunk, TW 8���10 % of SL (thick trunk, TW 14.6 % of SL). Nototriton guanacaste Garc��a-Par��s & Wake, 2000 has relatively smaller nostril openings, 0.4���0.95 % of SL (RNW = 1.3 % of SL); relatively shorter hind limb, 17���20 % of SL (longer hind limb, HLL = 21.9 % of SL). Nototriton major Good & Wake, 1993 has relatively smaller nostrils openings, 0.3 % of SL (RNW = 1.3 % of SL); shorter snout-gular length, 17.7 % of SL (longer snout-gular length, LGFS = 21 % of SL); relatively narrower head, 11.3 % of SL (broader head, HeW = 15.1 % of SL); slender trunk, TW 9���10 % of SL (trunk robust, TW 14.6 % of SL); relatively shorter hind limb, 19.3 % of SL (longer hind limb, HLL = 21.9 % of SL); relatively shorter fore limb, 14.8 % of SL (longer front limb, FLL = 20.1 % of SL). Nototriton matama Boza-Oviedo et al., 2012 has relatively larger nostrils openings, 2 % of SL (RNW = 1.3 % of SL); pointed outer toe tips (rounded outer toe tips); relatively narrower head, 14 % of SL (broader head, HeW = 15.1 % of SL). Nototriton picadoi (Stejneger, 1911) has a relatively narrower head, 12���14 % of SL (broader head, HeW = 15.1 % of SL); slender trunk, TW 10���11 % of SL (trunk robust, TW 14.6 % of SL); relatively shorter hind limb, 18���19 % of SL (longer hind limb, HLL = 21.9 % of SL); relatively shorter fore limb, 16.9���18.2 % of SL (longer front limb, FLL = 20.1 % of SL). Although Nototriton tapanti Good & Wake, 1993 is not a member of the N. picadoi species group, but we decided to include a phenotypic comparison here due to the fact that no DNA sequences of N. tapanti were available for molecular comparison with N. costaricense. Nototriton tapanti can easily be distinguished from N. costaricense by its syndactylous hands and feet (having at least the distal phalanges on fingers II, III, and IV in and toes II, III, IV, and V free of webbing); tips of free digits pointed and without subterminal pads (rounded tips and terminal pads present). Description of holotype. Subadult having a SL of 21.9 mm. Head slightly wider than neck and shoulders (HeW 3.3 mm, NeW 3.0 mm, ShW 2.9 mm), with greatest width of head just posterior to the articulation of the jaws; snout raised anterodorsally, spadate to rounded in dorsal outline, and rounded in profile; snout relatively short (SnL 0.7 mm, 3.2 % of SL), with nearly terminal non-protruding large nostrils (LNH 0.34 mm, RNW 0.31 mm) directed anterolaterally; internarial area convex in dorsal outline. Eyes relatively large (EW = 142 % of SnL), weakly protruding beyond dorsal and ventral outline of head, directed anterolaterally, with a distinct suborbital groove. Top of head flat and smooth, tapering slightly toward anterior terminus, lacking contrasting interorbital or other dermal structures. Canthus rostralis weakly rounded; intercanthal area flat to slightly convex; and loreal region slightly concave. No obvious cirri (nasolabial protuberances), but nasolabial grooves weakly discernable on tip of snout; nasolabial grooves start at lateral margins of nares and extend ventrally, with a slight outward orientation, and terminate prior to reaching upper lip margin. Gular fold well-defined, starting on dorsolateral portion of neck, and wrapping around lateral section of head and crossing venter as a smooth anterior-oriented curve. Upper lip markedly protruding beyond edge of lower lip in ventral view; and no mental gland visible under skin of anterior intermandibular region. A very weakly discernible groove starts at posterior dorsolateral margin of orbit and extends dorsolaterally to anterior margin of gular fold. Pair of weak, but discernably raised parotoid glands present on dorsolateral margin of head between orbits and occiput. Arms relatively long and slender (FLL = 4.4 mm, 20.1% of SL), without noticeable hypertrophied forearm compared to upper arm. Hands small and slender (HaL = 1.2 mm, 29.3% of VGS; HaW = 1.0 mm, 30.3 % of HeW). Fingers II, III, and IV relatively long (LF2 0.47 mm, LF3 0.47 mm) and robust with rounded tips; fingers II and IV nearly equal in width, but Finger III noticeably wider (WF3 0.22 mm); terminal pads poorly discernable, appearing to be present on ventral tips of fingers I, II, III, and IV; Finger I barely protruding beyond palmar tissue; webbing absent between fingers II and III and fingers III and IV. Palmar surface appearing to be smooth overall, but with an evident dermal crease extending transversally near proximal edge. Dorsal surfaces of hand with well defined interdigital grooves extending across metacarpal region. Relative lengths of fingers on right hand I Coloration in life. The ground color of the dorsal surfaces of the head and trunk consisted of a mixture of pale earthy tones (Light Flesh Color [250], Light Pratt���s Rufous [71], and Flame Scarlet [73]). Secondary coloration on the dorsal surfaces of the head and trunk consisted of irregular dark markings (Marron [39] and Dark Grayish Brown [284]), grayish white dashes, and numerous very fine circular Marron (39) chromatophores. On the posterior half of the parotoid glands there was a pale flesh colored patch. On the center of the head, between the orbits and occiput, there was a concentration of dark (Dark Grayish Brown [284]) irregular markings. On the left parotoid gland there was a dark wedge shaped mark, appearing to be a scar. The upper surfaces of the arms and legs have a Dark Grayish Brown (284) ground color. Secondary colors on the upper surfaces of the arms and legs are in the form of irregular spots either pearly white, Light Buff (2), Light Flesh Color (250), and Flame Scarlet (73). The hands and feet were a pale translucent gray. The dorsal surface of the tails was a mottled mixture of Flame Scarlet (73), Light Flesh Color (250), and Light Buff (2). The dominant color of the dorsal surface of the tail was Flame Scarlet. Additionally, there were numerous white spots and dashes on the dorsal surface of the tail. The colorations on the lateral surfaces of the body and tail were divided by a concentration of white markings or dashes that especially on the trunk formed what could be weakly interpreted as a longitudinal band. The inferior half of the lateral body and tail was Dark Grayish Brown (284) with scattered white markings. The superior half of the body and tail consisted of a mixture of earthy tones typical of the overall adjoining dorsal surfaces. The lateral surfaces of the head, between the orbits and the gular fold, were dominated by Flame Scarlet chromatophores (73); the mandibular region was dark gray with some scattered white markings or spots. The iris was a coppery-red with scattered dark spots and reticulation. The ventral skin of the head, body, and limbs was pale brown to gray with a very fine dark gray to black reticulation. Numerous white spots and irregular markings were also scattered throughout the ventral surfaces of the head and body. On the ventrolateral surface of the head, between the nuchal groove and gular fold, there was a patch of flame Scarlet skin marginally wrapping onto the ventral surface. Overall, the tone of the gular region was noticeably paler than the rest of the ventral surfaces. The palmar and plantar surfaces were pale grayish-white. The ventral surface of the tail is similar in general appearance to that of the body, but the ground color is noticeably a darker tone of gray. Coloration in ethanol. After nearly two years in ethanol (70%), the overall coloration of the holotype has darkened throughout and contains a mixture of four principal colors; Chestnut (30), Flame Scarlet (73), Raw Umber (280), and a pale fleshy gray Measurements (in mm), tooth counts, limb interval, and percentages of the holotype. SL 21.9; Tal 25.0; TL 46.9; ShW 2.9; HeW 3.3; NeW 3.0; EW 1.1; SnL 0.7; JSL 2.5; LGFS 4.6; LNH 0.34; RNW 0.31; IND 0.7; NLP 0.4; ICD 1.6; HLL 4.8; FLL 4.4; TW 3.2; VGS 4.1; FSL 6.4; UHL 2.7; AGL 12.0; VL 1.4; HaW 1.0; HaL 1.2; LF2 0.47; LF3 0.47; WF3 0.22; FoW 1.4; FoL 1.6; LT2 0.71; LT3 0.54; WT3 0.26. Limb interval 5. Measurements in related percentages: VGS/SL 18.7 %; IND /HeW 21.2 %; AGL/SL 54.8 %; HeW/SL 15.1 %; Hew/AGL 27.5 %; SnL/ HeW 21.2 %; LNH/HeW 10.3 %; LNH/SL 1.6 %; RNW/HeW, 9.4 %; RNW/SL 1.4 %; HLL /SL 21.9 %; FLL/SL 20.1 %; HaL/VGS 29.3 %; FoL/VGS 39.0 %; Haw/HeW 30.3 %; FoW/HeW 42.4 %; LT3/HeW 16.4 %. Etymology. The specific epithet refers to the Spanish word meaning Costa Rican, ��� costaricense ���. The name represents the fact that the holotype and species was discovered in Costa Rica. Given the close proximity of the type locality to the Costa Rica-Panama border, we speculate that one day it may indeed be discovered within Panama as well. Habitat and natural history observations. The holotype of Nototriton costaricense was discovered inside a moss mat that was growing approximately one meter above the ground on the trunk of a tree. The type locality is ca. 2.5 km SSW of a peak known as Cerro Pat; the general site is a relatively flat ridge, ranging from 1400���1500 m asl, which is covered by mature forest. Despite the elevation of the site typically being dominated by cloud forest with an abundance of bryophytes and epiphytes, it was nearly devoid of such floral conditions; bryophytes and epiphytes were uncommon there. The type locality is located on a ridgeline that extends out towards the north from the Kamuk massif. Just south of the type locality, the topography of the ridgeline leading towards the Kamuk massif drastically rises until reaching an unnamed peak with an elevation of approximately 1850 m asl. The unnamed peak and its slopes is covered in mature forest, and in contrast to that of the type locality, there is an abundance of bryophytes and other epiphytes covering the trees. Despite an intense search by EA through moss, bromeliads, and leaf litter, in what appeared to be more optimal habitat along the slopes of the unnamed peak, no additional salamanders were found. Inasmuch as Nototriton costaricense is only known from a single specimen, any assumptions about its natural history are highly speculative. With that said, however, like other members of the Nototriton picadoi species group, it is probable that this species is prone to living within moss mats, both in terrestrial and arboreal situations, within cloud forest. All known Costa Rican species of Nototriton are known to principally inhabit moss, unlike congeners of the subgenus Bryotriton found in Nuclear Central America that principally known to inhabit microhabitats such as leaf litter, rotting logs, and bromeliads (Townsend et al. 2010, 2011, 2013; Townsend 2016). Within another patch of moss on the same tree where the holotype of N. costaricense was found, an individual of Bolitoglossa minutula was also discovered (Arias 2017). Other amphibian species found in sympatry with Nototriton costaricense are Craugastor podiciferus, C. aff. underwoodi, Diasporus hylaeformis, Incilius epioticus, Pristimantis caryophyllaceus, and P. pardalis. Distribution. Nototriton costaricense is known from a single site within the Lower Montane Rain Forest life zone (Holdridge 1967) along the mid-elevation slopes of the extreme southeastern Cordillera de Talamanca, within Parque Internacional La Amistad, in the vicinity of Cerro Pat, ca. 1500 m (Fig. 1). Further explorations and encounters with this species are needed before we will be able to further understand its potential range. We speculate that it is very possible that N. costaricense extends into Panama due to the proximity and relatively homogenous habitat lying between the type locality and the border (~ 11 kilometers straight line distance). Nototriton costaricense, if found to inhabit Panama, would represent the first species of this genus to extend beyond Costa Rica. Discussion With the discovery and description of Nototriton costaricense, the number of species known for this genus within Costa Rica is now elevated to nine, and the total diversity of salamanders for the tiny country 52. In the last decade 11 species (23.5% of total) of salamanders were described from Costa Rica (Wake et al. 2007; Garc��a-Par��s et al. 2008; Bola��os & Wake 2009; Boza-Oviedo et al. 2012; Kubicki 2016; Kubicki & Arias 2016), indicating that the inventory of salamanders from the country is incomplete. The bolitoglossines recently described from Costa Rica are only known from a few localities and relatively few specimens, and half of these species are known from the Caribbean slopes of Costa Rica. We would like to emphasize, again, that special attention needs to placed on improving the knowledge of the diversity of not only salamanders, but amphibians in general along the midelevation Caribbean slopes of the Cordillera de Talamanca of Costa Rica. The mid-elevation slopes along the Caribbean versant of Talamanca, due to the difficult terrain and over all limited access, harbor some of the poorest known and least explored forests within Costa Rica; further inventories should be viewed as a priority in this overall region. With our discovery of Nototriton costaricense, the distribution of genus is now known to extend ~ 50 km to the south-southeast from the previously known southern limit of the genus, which was the type locality of N. matama on the southeast terminus of the Matama ridge, Lim��n Province, Costa Rica. Between the type localities of N. matama and N. costaricense, there is an abundance of suitable habitat for the genus, and this area needs to be surveyed further in attempt to learn more about the actual distributional boundaries between these two enigmatic and recently described taxa. The genetic distances between Nototriton costaricense and the other members of the N. picadoi species group range from 1.8���2.4 % for 16S; this distance is relatively low in comparison to those suggested by Fouquet et al. (2007) for Neotropical frogs, in which the authors recommend a minimum of 3% 16S divergence be recognized as threshold to define candidate species. We recently commented that this threshold possibly underestimates the true diversity of Neotropical salamanders (Kubicki & Arias 2016), and that each taxon should be carefully evaluated on an individual basis considering all the phenotypic, geographic, and molecular evidence at hand. Kubicki & Arias (2016) described a well-supported morphologically distinctive new species of Bolitoglossa that had genetic distances as low as 2.0 % (16S) among its closest related taxa. Bolitoglossa splendida Boza-Oviedo et al., 2012 is another example of salamander that is well recognized for its phenotypic uniqueness, but presents once again a relatively low (2.0 % 16S) genetic divergence from of its closest related congeners (Boza-Oviedo et al. 2012). Within the genus Nototriton it has been previously published and justified that low genetic distances are known exist among well-recognized species, i.e. 1.9 % in 16S between N. barbouri and N. limnospectator (Townsend 2016). Rovito et al. (2013) used morphological and molecular (two mitochondrial and one nuclear genes) data to define the, Published as part of Arias, Erick & Kubicki, Brian, 2018, A new moss salamander, genus Nototriton (Caudata: Plethodontidae), from the Cordillera de Talamanca, in the Costa Rica-Panama border region, pp. 487-500 in Zootaxa 4369 (4) on pages 493-498, DOI: 10.11646/zootaxa.4369.4.2, http://zenodo.org/record/1136402, {"references":["Boza-Oviedo, E., Rovito, S. M., Chaves, G., Garcia-Rodriguez, A., Artavia, L. G., Bolanos, F. & Wake, D. 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. Zootaxa, 3309, 36 - 61.","Townsend, J. H. (2016) Taxonomic revision of the moss salamander Nototriton barbouri (Schmidt) (Caudata: Plethodontidae), with description of two new species from the Cordillera Nombre de Dios, Honduras. Zootaxa, 4196 (4), 511 - 528. https: // doi. org / 10.11646 / zootaxa. 4196.4.3","Taylor, E. H. (1948) New Costa Rican salamanders. Proceedings of Biological Society of Washington, 61, 177 - 180.","Good, D. A. & Wake, D. B. (1993) Systematic studies of the Costa Rican moss salamanders, genus Nototriton, with descriptions of three new species. Herpetological Monographs, 7, 131 - 159. https: // doi. org / 10.2307 / 1466956","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. https: // doi. org / 10.1643 / 0045 - 8511 (2000) 2000 [0042: MPAORO] 2.0. CO; 2","Stejneger, L. (1911) Description of three new batrachians from Costa Rica and Panama. Proceedings of the United States National Museum, 41, 285 - 288. https: // doi. org / 10.5479 / si. 00963801.1857.285","Townsend, J. H., Butler, J. M., Wilson, L. D. & Austin, J. D. (2010) A distinctive new species of moss salamander (Caudata: Plethodontidae: Nototriton) from an imperiled Honduran endemism hotspot. Zootaxa, 2434, 1 - 16.","Townsend, J. H., Medina-Flores, M., Murillo, J. L. & Austin, J. D. (2011) Cryptic diversity in Chortis Highland moss salamanders (Caudata: Plethodontidae: Nototriton) revealed using mtDNA barcodes and phylogenetics, with a new species from eastern Honduras. Systematics and Biodiversity, 9, 275 - 287. https: // doi. org / 10.1080 / 14772000.2011.611188","Townsend, J. H., Medina-Flores, M., Reyes-Calderon, O. & Austin, J. D. (2013) A new Nototriton (Caudata: Plethodontidae) from Parque Nacional Montana de Botaderos in northeastern Honduras. Zootaxa, 3666 (3), 358 - 368. https: // doi. org / 10.11646 / zootaxa. 3666.3.6","Arias, E. (2017) New distribution records for two uncommon amphibian species on the Caribbean slopes of the Cordillera de Talamanca, Costa Rica. Mesoamerican Herpetology, 4, 446 - 449.","Holdridge, L. R. (1967) Life Zone Ecology. Tropical Science Center, San Jose, Costa Rica, 206 pp.","Wake, D. B., Savage, J. M. & Hanken, J. (2007) Montane salamanders from the Costa Rica - Panama border region, with descriptions of two new species of Bolitoglossa. Copeia, 2007, 556 - 565.","Garcia-Paris, M., Parra-Olea, G. & Wake, D. B. (2008) Description of a new species of the Bolitoglossa subpalmata group (Caudata: Plethodontidae) from Costa Rica. Herpetological Journal, 18, 23 - 31.","Bolanos, F. & Wake, D. B. (2009) Two new species of montane web-footed salamanders (Plethodontidae: Bolitoglossa) from the Costa Rica-Panama border region. Zootaxa, 1981, 57 - 68.","Kubicki, B. & Arias, E. (2016) A beautiful new yellow salamander, genus Bolitoglossa (Caudata: Plethodontidae), from the northeastern slopes of the Cordillera de Talamanca, Costa Rica. Zootaxa, 4184 (2), 329 - 346. https: // doi. org / 10.11646 / zootaxa. 4184.2.5","Fouquet, A., Gilles, A., Vences, M., Marty, C., Blanc, M. & Gemmell, N. J. (2007) Underestimation of species richness in Neotropical frogs revealed by mtDNA analyses. PLoS one, 2, e 1109. https: // doi. org / 10.1371 / journal. pone. 0001109","Rovito, S. M., Parra-Olea, G., Hanken, J., Bonett, R. M. & Wake, D. B. (2013) Adaptive radiation in miniature: the minute salamanders of the Mexican highlands (Amphibia: Plethodontidae: Thorius). Biological Journal of the Linnean Society, 109, 622 - 643. https: // doi. org / 10.1111 / bij. 12083","Rovito, S. M., Parra-Olea, G., Recuero, E. & Wake, D. B. (2015) Diversification and biogeographical history of Neotropical plethodontid salamanders. Zoological Journal of the Linnean Society, 175, 167 - 188. https: // doi. org / 10.1111 / zoj. 12271","Nicolas, V., Schaeffer, B., Missoup, A. D., Kennis, J., Colyn, M., Denys, C., Tatard, C., Cruaud, C. & Laredo, C. (2012) Assessment of three mitochondrial genes (16 S, Cytb, CO 1) for identifying species in the Praomyini tribe (Rodentia: Muridae). PLoS one, 7, e 36586. https: // doi. org / 10.1371 / journal. pone. 0036586","Johns, G. C. & Avise, J. C. (1998) A comparative summary of genetic distances in the vertebrates from the mitochondrial cytochrome b gene. Molecular Biology and Evolution, 15, 1481 - 1490. https: // doi. org / 10.1093 / oxfordjournals. molbev. a 025875","Meyer, C. P. & Paulay, G. (2005) DNA barcoding: error rates based on comprehensive sampling. PLoS biology, 3, e 422. https: // doi. org / 10.1371 / journal. pbio. 0030422","Clare, E. L., Lim, B. K., Fenton, M. B. & Hebert, P. D. N. (2011) Neotropical bats: estimating species diversity with DNA barcodes. PloS one, 6, e 22648. https: // doi. org / 10.1371 / journal. pone. 0022648"]}

A new salamander belonging to the genus Nototriton, subgenus Nototriton, is described from the Caribbean slopes of the southeastern Cordillera de Talamanca in Costa Rica, within Parque Internacional La Amistad, at an elevation ca. 1500 m a.s.l. This new taxon is distinguished from its congeners by its morphological characteristics and by its differentiation in DNA sequences of the 16S rRNA, cytochrome oxidase subunit I (COI), and cytochrome b mitochondrial genes. This new species represents the southernmost extension known for the genus Nototriton.

Nototriton nelsoni sp. nov. Tex��guat Moss Salamander Figure 5 Nototriton barbouri: McCranie et al. (1993: 387). Nototriton sp. (in part): Townsend et al. (2010: 3). Nototriton sp. B.: Townsend et al. (2011a: 277), Rafa��lli (2013: 368), Townsend et al. (2013: 360). Nototriton sp.: Townsend et al. (2012: 96). Nototriton (Bryotriton) barbouri (in part): Dubois & Raffa��lli (2012: 141), Rafa��lli (2013: 368). Holotype. USNM 578300 (Fig. 5), an adult male from Cerro El Chino (15.525394��N, 87.278672��W), 1,420 m elevation, above La Liberaci��n, Refugio de Vida Silvestre Texiguat, Departamento de Atl��ntida, Honduras; collected 19 June 2010 by B.K. Atkinson, C.A. Cerrato-Mendoza, J.H. Townsend, and L.D. Wilson; original field number JHT 3159. GenBank accession numbers JN377387 (16S), JN377391 (cyt b), JN377403 (COI). Paratypes. Four, all from about 2.5 km (airline) NNE of La Fortuna (15.44��N, 87.31��W), Refugio de Vida Silvestre Texiguat, Departamento de Yoro; one male, USNM 339710, 1,690 m elevation; and two females, USNM 339709, 1,690 m elevation, and USNM 339711, 1,800 m elevation; and one cleared-and-stained specimen, USNM 509333, 1,600 m elevation, GenBank accession number AF199138 (cyt b). Diagnosis. A member of the genus Nototriton diagnosed by possessing 13 costal grooves (>16 costal grooves in Oedipina), the presence of a sublingual fold and hands and feet longer than broad (sublingual fold absent and hands and feet broader than long in Bolitoglossa), and small nares (0.006���0.010 NL/SL; 0.017���0.029 NL/SL in Cryptotriton and Dendrotriton). Phylogenetic analysis supports inclusion of N. nelsoni in the subgenus Bryotriton, belonging to the northern clade with N. brodiei, N. oreadorum, and N. stuarti (Fig. 1). From the other members of the northern clade, N. nelsoni can be distinguished from N. brodiei by having a relatively broader head (HW/SVL 0.127���0.142, versus 0.120 in N. brodiei), relatively longer limbs (FLL/SVL 0.173���0.198 and HLL /SVL 0.203��� 0.223, versus 0.148���0.151 and 0.166���0.180 in N. brodiei), and by having fewer maxillary teeth (38���53, versus 60��� 62 in N. brodiei); from N. oreadorum by having a relatively longer and broader head (HL/SVL 0.185���0.226 and HW/SVL 0.127���0.142, versus 0.178���0.182 and 0.117���0.122 in N. oreadorum) and relatively longer front limbs (FLL/SVL 0.173���0.198, versus 0.158���0.178 in N. oreadorum); and from N. stuarti in having relatively longer hind limbs (HLL /SVL 0.203���0.223, versus 0.178 in N. stuarti) and a greater number of maxillary teeth (38���53, versus 36 in N. stuarti). From the other constituent species of Bryotriton, N. nelsoni can be differentiated from N. barbouri by having a relatively longer tail (TL/SVL 1.144���1.532, versus 0.802���1.165 in N. barbouri) and a lower average number of costal grooves between adpressed limbs (4.9 [�� 1.4], versus 6.2 [��0.9] in N. barbouri); from N. lignicola in having relatively broader hind feet (HFW /SVL 0.044���0.058, versus 0.035���0.046 in N. lignicola) and a lower average number of costal grooves between adpressed limbs (4.9 [�� 1.4], versus 5.9 [��0.5] in N. barbouri); from N. limnospectator in having a relatively broader head (HW/SVL 0.127���0.142, versus 0.111���0.118 in N. limnospectator); from N. mime in having a relatively longer tail (TL/SVL 1.144���1.532, versus 0.698���1.117 in N. mime) and relatively shorter limbs (FLL/SVL 0.173���0.198 and HLL /SVL 0.203���0.223, versus 0.195���0.246 and 0.224���0.254 in N. mime); from N. picucha in having broader hind feet (HFW /SVL 0.044���0.058, versus 0.042��� 0.043 in N. picucha) and a higher average number of costal grooves between adpressed limbs (4.9 [�� 1.4], versus 4.0 [��0.7] in N. picucha); and from N. tomamorum in having feet with well-developed digits (syndactylous feet in N. tomamorum), a greater number of maxillary and vomerine teeth (38���53 maxillary teeth and 17���24 vomerine teeth, versus 26 maxillary teeth and 11 vomerine teeth in N. tomamorum) and smaller nares (NL/SL 0.006���0.010, versus 0.018 in N. tomamorum). Nototriton nelsoni can be distinguished from N. saslaya, the only species of the subgenus Nototriton that occurs in the Chort��s Highlands, by having more maxillary and vomerine teeth (38���53 maxillary teeth and 17���24 vomerine teeth, versus 17���22 maxillary teeth and 3���11 vomerine teeth in N. saslaya). Each of the species of Bryotriton in the northern clade are allopatric with respect to each other, and N. nelsoni can be further distinguished from these species based on model-corrected genetic distances (Table 2), being 2.1% (16S) and 9% (cyt b) divergent from its closest relative, N. stuarti, and 2.3% (16S) and 5.4���5.7% (cyt b) divergent from samples from PN Pico Bonito, the geographically closest population, itself described as a new species below. Description of holotype. An adult male (SVL = 31.9 mm, total length = 68.4 mm) with a slender body and reduced limbs. The head is rounded, slightly broader than the body; nostrils are relatively small (NL/SVL=0.006), and the snout is acutely rounded and of moderate length. Nasolabial grooves are visible but slight, with nasolabial protuberances apparent. The eyes are relatively large and protuberant. The mental gland is not apparent. There are 53 maxillary teeth, 4 slightly enlarged premaxillary teeth in line with the maxillary teeth, and 24 vomerine teeth in two well-defined arches. The limbs are short (CLL/SVL=0.42), with approximately 3.5 costal grooves between the adpressed limbs. The hands and feet are narrow with well-developed digits bearing subdigital pads. The relative length of the digits is IMeasurements of holotype (in mm). SVL 31.9; AG 19.1; TW 3.8; HL 6.9; HW 4.2; TL 41.0; HLL 7.2; FLL 6.4; CLL 13.6; FFW 1.1; HFW 1.5; NL 0.2; eyelid length 1.8; eye width 0.9; interorbital distance 1.6; anterior rim of orbit to snout 1.5; distance separating internal margins of nares 1.4; distance separating external margins of nares 1.8. Color in life of holotype. Description of the coloration of the holotype based on a series of color photographs taken in life (color names and numbers follow K��hler [2012]): dorsal surfaces of paratoid region of head, body, and tail darkly mottled, with dominant ground color being Dark Carmine (61); lateral surfaces more heavily mottled with Brick Red (36), becoming nearly solid Ferruginous (35) near the edge of the ventral coloration, with Sepia (286) spots beginning on the centrolateral portion of costal groove 3 and becoming larger posteriorly, with one large blotch present centrolaterally from costal groove 12 to the hind limb; series of irregular Salmon Color (251) middorsal botches corresponding with the relative position of each vertebra, forming the center of a faint herringbone dorsal pattern; dorsal surfaces of head and dorsal and lateral surfaces of body covered with small, irregular Smoky White (261) and Pearl Gray (262) spots; dorsal surface of head mottled Sepia (286), Dark Carmine (61), and Carmine (64), with Smoky White (261) and Pearl Gray (262) spotting, becoming more profuse on lateral surfaces of head; series of Pearl Gray (262) spotting lining the margin of the upper lip; dorsal surface of limbs Sepia (286) with Dark Carmine (61), Deep Vinaceous (248), and Light Pratt���s Rufous (71) stippling, becoming more profuse towards body; ventral surface of head and body Fuscous (283), with Smoky White (261) and Pearl Gray (262) spots slighting larger than those seen laterally; dorsal and lateral surfaces of tail Dark Carmine (61), with Smoky White (261) and Light Lavender (201) spotting more profuse laterally; ventral ground color of tail Dusky Brown (285), being somewhat darker than body, with scattered Smoky White (261) and Light Lavender (201) spotting; color of iris Peach Red Ruby (70). Osteology. Based on examination of digital radiographs for the holotype (Fig. 6 A), Nototriton nelsoni is a typical member of the genus possessing a single cervical vertebra, 14 trunk vertebrae (13 of which bear ribs), and 2 caudosacral vertebrae; a completely fused skull roof formed through contact of the parietal bones; frontal processes of premaxilla fused at point of origin and separate immediately dorsoposterior to origin; preorbital vomerine processes well-developed into a pair of elongate arches each bearing a single row of numerous teeth, extending beyond the outer margins of the choanae; columella absent; phalangeal formulae 1-2-3-2 and 1-2-3-3-2; penultimate phalanges reduced, exceeded in length by terminal phalanges on digits II, III, and IV of the forelimbs and digits II, III, IV, and V of hind limbs; terminal phalanges slightly expanded at distal tips, expansion more pronounced in digit III of forelimbs and digit III of the hindlimbs; mesopodial elements not mineralized. Examination of radiographs of the three paratypes (USNM 339709���11) generally agreed with those of the holotype; with one paratype (USNM 339710) demonstrating incomplete fusion of the skull with sutures apparent between the frontal and parietal elements, and a small frontoparietal fontanelle. Etymology. The specific epithet is a patronym honoring Dr. Cyril ���Cirilo��� Hardy Nelson-Sutherland, Professor Emeritus and co-founder of the Department of Biology at the Universidad Nacional Aut��noma de Honduras, and co-founder and former director and curator of the university herbarium, which was renamed in his honor in 2011. Dr. Nelson devoted his career to studying the flora of Honduras and providing instruction and guidance to generations of Honduran biology students, a career which culminated with the publication of the 1,576-page magnum opus, the Cat��logo de las Plantas Vasculares de Honduras, Espermatofitas (Nelson- Sutherland 2008). Geographic and ecological distribution. Nototriton nelsoni is known only from highland forest within Refugio de Vida Silvestre Tex��guat, 1,420���1,800 m (Fig. 3). Known localities fall entirely within the Lower Montane Wet Forest formation (Holdridge 1967), with the habitat at the type locality being Broadleaf Cloud Forest and the locality for the paratypes being Mixed Cloud Forest transitioning to Broadleaf Cloud Forest (Townsend 2014). Natural history. The holotype was collected as it crawled out of an arboreal bromeliad approximately 1.5 m above the ground on a rainy night. The collection site was at the top of a small peak at the northern terminus of a ridgeline leading to the highest portion of RVS Tex��guat, and the overstory was somewhat stunted with a palmdominated understory and a dense covering of epiphytic plants. This species has also been found utilizing rotten logs as daytime refugia, in sympatry with the worm salamander Oedipina gephyra (McCranie et al. 1993: 387). The congener N. tomamorum is known from a single specimen collected near a stream just below the locality where the four paratypes of N. nelsoni were collected. Remarks. A photograph of the holotype of Nototriton nelsoni appeared in Townsend et al. (2012: 98) as ��� Nototriton sp.���., Published as part of Townsend, Josiah H., 2016, Taxonomic revision of the moss salamander Nototriton barbouri (Schmidt) (Caudata: Plethodontidae), with description of two new species from the Cordillera Nombre de Dios, Honduras, pp. 511-528 in Zootaxa 4196 (4) on pages 520-523, DOI: 10.11646/zootaxa.4196.4.3, http://zenodo.org/record/168312, {"references":["McCranie, J. R., Wilson, L. D. & Williams, K. L. (1993) A new species of Oedipina (Amphibia: Caudata: Plethodontidae) from northern Honduras. Proceedings of the Biological Socity of Washington, 106, 385 - 389.","Townsend, J. H., Butler, J. M., Wilson, L. D. & Austin, J. D. (2010) A distinctive new species of moss salamander (Caudata: Plethodontidae: Nototriton) from an imperiled Honduran endemism hotspot. Zootaxa, 2434, 1 - 16.","Townsend, J. H., Medina-Flores, M., Murillo, J. L. & Austin, J. D. (2011 a) Cryptic diversity in Chortis Highland moss salamanders (Caudata: Plethodontidae: Nototriton) revealed using mtDNA barcodes and phylogenetics, with a new species from eastern Honduras. Systematics and Biodiversity, 9, 275 - 287. http: // dx. doi. org / 10.1080 / 14772000.2011.611188","Rafaelli, J. (2013) Les Urodeles du Monde, 2 nd ed. Penclen Edition, Plumelec, France.","Townsend, J. H., Medina-Flores, M., Reyes-Calderon, O. & Austin, J. D. (2013) A new Nototriton (Caudata: Plethodontidae) from Montana de Botaderos National Park in northeastern Honduras. Zootaxa, 3666 (3), 358 - 368. http: // dx. doi. org / 10.11646 / zootaxa. 3666.3.6","Townsend, J. H., Wilson, L. D., Medina-Flores, M., Aguilar-Urbina, E., Atkinson, B. K., Cerrato-M., C. A., Contreras-C., A., Gray, L. N., Herrera-B., L. A., Luque-Montes, I. R., McKewy-Mejia, M., Portillo-A., A., Stubbs, A. L. & Austin, J. D. (2012) A premontane hotspot of herpetofaunal endemism on the windward side of Refugio de Vida Silvestre Texiguat, Honduras. Salamandra, 48, 92 - 114.","Dubois, A. & Raffaelli, J. (2012) A new ergotaxonomy of the order Urodela Dumeril, 1805 (Amphibia, Batrachia). Alytes 28, 77 - 161.","Kohler, G. (2012) Color Catalogue for Field Biologists. Herpeton Verlag, Offenbach, 49 pp.","Nelson-Sutherland, C. H. (2008) Catalogo de las Plantas Vasculares de Honduras, Espermatofitas. Secretaria de Recursos Naturales y Ambiente / Guaymuras, Tegucigalpa, xxix + 1576 pp.","Holdridge, L. R. (1967) Life Zone Ecology. Revised Edition. Tropical Science Center, San Jose, Costa Rica, 206 pp.","Townsend, J. H. (2014) Characterizing the Chortis Block Biogeographic Province: Geological, physiographic, and ecological associations and herpetofaunal diversity. Mesoamerican Herpetology, 1, 204 - 252."]}

Nototriton oreadorum sp. nov. Pico Bonito Moss Salamander Figure 6 B Nototriton barbouri: McCranie (1996: 28), Nototriton barbouri (in part): McCranie & Wilson (2002: 143), McCranie & Casta��eda (2005: 7; 2007: 108). Nototriton sp. (in part): Townsend et al. (2010: 3). Nototriton cf. barbouri: K��hler (2011: 78). Nototriton sp. A.: Townsend et al. (2011a: 277), Rafa��lli (2013: 368), Townsend et al. (2013: 360). Nototriton (Bryotriton) barbouri (in part): Dubois & Raffa��lli (2012: 141), Rafa��lli (2013: 368). Holotype. USNM 497552, an adult female from the south slope of Cerro B��falo (15.66��N, 86.79��W), 1,540 m elevation, Parque Nacional Pico Bonito, Departamento de Atl��ntida, Honduras; collected 30 May 1996 by S. Gotte and J.R. McCranie; original field number LDW 10 724; GenBank accession number AF199137 (cyt b). Paratype. USNM 339712, an adult female from Quebrada de Oro (15.64��N, 86.80��W), 1,210 m elevation, Parque Nacional Pico Bonito, Departamento de Atl��ntida, Honduras; GenBank accession numbers AF199201 (16S), AF199136 (cyt b). Diagnosis. A member of the genus Nototriton diagnosed by possessing 13 costal grooves (>16 costal grooves in Oedipina), the presence of a sublingual fold and hands and feet longer than broad (sublingual fold absent and hands and feet broader than long in Bolitoglossa), and small nares (NL/SVL 0.006���0.009; NL/SVL 0.017���0.029 in Cryptotriton and Dendrotriton). Phylogenetic analysis supports inclusion of N. oreadorum in the subgenus Bryotriton, and is recovered as a member of the northern clade with N. brodiei, N. nelsoni, and N. stuarti (Fig. 1). From the other species of the northern clade of Bryotriton, N. nelsoni can be distinguished from N. brodiei in having a relatively shorter tail (TL/SVL 0.117���0.122, versus 1.420���1.440 in N. brodiei), relatively longer front limbs (FLL/SVL 0.158���0.178, versus 0.148���0.151 in N. brodiei), relatively shorter hind limbs (HLL /SVL 0.190��� 0.222, versus 0.166���0.180 in N. brodiei), and fewer maxillary teeth (47���52, versus 60���62 in N. brodiei); from N. nelsoni in having a relatively shorter and narrower head (HL/SVL 0.178���0.182 and HW/SVL 0.117���0.122, versus 0.185���0.226 and 0.127���0.142 in N. nelsoni), relatively shorter front limbs (FLL/SVL 0.158���0.178, versus 0.173��� 0.198 in N. nelsoni), and a higher average number of costal grooves between adpressed limbs (6.0 [�� 0.7], versus 4.9 [��1.4] in N. nelsoni); and from N. stuarti by having a relatively narrower head (HW/SVL 0.117���0.122, versus 0.138 in N. stuarti), shorter tail (TL/SVL 1.111���1.169, versus 1.264 in N. stuarti), longer hind limbs (0.190���0.222, versus 0.178 in N. stuarti), smaller nares (NL/SVL 0.006���0.009, versus 0.012 in N. stuarti), and more maxillary teeth (47���52, versus 36 in N. stuarti). From the remaining species of Bryotriton, N. oreadorum can be differentiated from N. barbouri in having relatively broader hind feet (HFW /SVL 0.056���0.057, versus 0.034���0.052 in N. barbouri); from N. lignicola in having relatively broader hind feet (HFW /SVL 0.056���0.057, versus 0.035���0.046 in N. lignicola); from N. limnospectator in having a higher average number of costal grooves between adpressed limbs (6.0 [�� 0.7], versus 4.4 [��0.8] in N. limnospectator); from N. mime in having a relatively longer and broader head (HL/SVL 0.178��� 0.182 and HW/SVL 0.117���0.122, versus 0.182���0.201 and 0.121���0.138 in N. mime), a relatively longer tail (TL/ SVL 1.111���1.169, versus 0.698���1.117 in N. mime), relatively shorter limbs (FLL/SVL 0.158���0.178 and HLL /SVL 0.190���0.222, versus 0.195���0.246 and 0.224���0.254 for N. mime), more maxillary teeth (47���52, versus 27���46 in N. mime), and a higher average number of costal grooves between adpressed limbs (6.0 [�� 0.7], versus 4.8 [��0.4] in N. mime); and from N. picucha in having a relatively shorter and narrower head (HL/SVL 0.178���0.182 and HW/SVL 0.127���0.142, versus 0.197���0.198 and 0.140���0.148 in N. picucha), shorter limbs (FLL/SVL 0.158���0.178 and HLL / SVL 0.190���0.222, versus 0.195���0.246 and 0.224���0.254 for N. picucha), more maxillary and vomerine teeth (47��� 52 maxillary teeth and 20���22 vomerine teeth, versus 41 maxillary teeth and 16���19 vomerine teeth for N. picucha), and a higher average number of costal grooves between adpressed limbs (6.0 [�� 0.7], versus 4.0 [��0.7] in N. picucha). One species of Bryotriton, N. tomamorum, is unique among members of the subgenus in having syndactylous feet (well-differentiated digits in N. oreadorum) and large nares (NL/SVL 0.018, versus 0.006���0.009 in N. oreadorum). Nototriton oreadorum can be distinguished from N. saslaya, the only species of the subgenus Nototriton found in the Chort��s Highlands, by having more maxillary and vomerine teeth (47���52 maxillary teeth and 20���22 vomerine teeth, versus 17���22 maxillary teeth and 3���11 vomerine teeth in N. saslaya). The four species of the northern clade of Bryotriton are all allopatric, and are further differentiated from all other species of Nototriton in the Chort��s Highlands by model-corrected genetic distances (Table 2), being 1.5% (16S) and 6% (cyt b) divergent from its closest relative, N. brodiei, and 2.3% (16S) and 5.4���5.7% (cyt b) divergent from samples from N. nelsoni, the species with the most geographically proximate population. Description of holotype. An adult female (SVL = 33.7 mm, total length = 73.1 mm) with a slender body and reduced limbs. The head is slightly rounded and slightly broader than the body; nostrils are relatively small (NL/ SVL=0.009), and the snout is rounded and of moderate length. Shallow nasolabial grooves are present, with nasolabial protuberances also apparent but not pronounced. The eyes are relatively large and protuberant. There are 47 maxillary teeth, 4 slightly enlarged premaxillary teeth in continuous line with the maxillaries, and 22 vomerine teeth in two well-defined arches. The limbs are short (CLL/SVL=0.37), with approximately five costal grooves between the adpressed limbs. The hands and feet are narrow with well-defined digits that bear subdigital pads. The relative length of the digits is IMeasurements of holotype (in mm). SL 33.7; AG 19.5; TW 4.9; HL 6.0; HW 4.1; TL 39.4; HLL 6.4; FLL 6.0; CLL 12.4; FFW 1.0; HFW 1.9; NL 0.3; eyelid length 1.7; eye width 1.2; interorbital distance 1.3; anterior rim of orbit to snout 1.5; distance separating internal margins of nares 1.1; distance separating external margins of nares 1.6. Coloration. Color in life of the female paratype (USNM 339712) as described by McCranie & Wilson (2002: 144) is as follows (color names follow Smithe 1975 ���1981): dorsal surface of body Mars Brown (223A), all lateral surfaces Mars Brown (223A) with silver flecks; dorsal surface of tail Verona Brown (223B); ventral and subcaudal surfaces Raw Unber (223) with pale gold flecking. Osteology. Based on examination of digital radiographs for the holotype (Fig. 6 B), Nototriton oreadorum is a typical member of the genus possessing a single cervical vertebra, 14 trunk vertebrae (13 of which bear ribs), and 2 caudosacral vertebrae; a partially fused skull roof formed through contact of the frontal and parietal bones, with the anterior portion of the frontals unfused and creating a narrow suture; frontal processes of premaxilla fused at point of origin and separate immediately dorsoposterior to origin; preorbital vomerine processes well-developed into a pair of symmetrical elongate arches, each bearing a single row of numerous teeth, extending beyond the outer margins of the choanae; columella absent; phalangeal formulae 1-2-3-2 and 1-2-3-3-2; penultimate phalanges reduced, equaled or exceeded in length by terminal phalanges on digits II, III, and IV of the forelimbs and digits II, III, IV, and V of hind limbs; terminal phalanges slightly expanded at distal tips, expansion more pronounced in digit III of forelimbs and digit III of the hindlimbs; mesopodial elements not mineralized. Examination of radiographs of the paratype (USNM 339712) generally agreed with those of the holotype, with the paratype having less complete fusion of the frontal and parietal elements than the holotype. Etymology. The specific epithet means ���belonging to the mountain nymphs���, referring to the Oreads of Greek mythology. The Oreads were female nature deities associated with mountains and valleys, alluding to the steep and challenging terrain at the type locality. Geographic and ecological distribution. This species is known only from the vicinity of Cerro B��falo in Parque Nacional Pico Bonito, Honduras, 1,210���1,540 m elevation. The distribution falls within the Lower Montane Wet Forest formation (Holdridge 1967), with the habitat at the type locality classified as Broadleaf Cloud Forest (Townsend 2014). Natural history. The holotype of N. oreadorum was found during the day under a log on a relatively steep forested slope on the south-southeastern face of Cerro B��falo (McCranie & Townsend 2011). This species occurs in sympatry with another endemic semi-fossorial salamander, Oedipina petiola, a species known only from a single specimen collected from under another log located uphill from the type locality (McCranie & Townsend 2011)., Published as part of Townsend, Josiah H., 2016, Taxonomic revision of the moss salamander Nototriton barbouri (Schmidt) (Caudata: Plethodontidae), with description of two new species from the Cordillera Nombre de Dios, Honduras, pp. 511-528 in Zootaxa 4196 (4) on pages 523-525, DOI: 10.11646/zootaxa.4196.4.3, http://zenodo.org/record/168312, {"references":["McCranie, J. R. (1996) Geographic Distibution. Nototriton barbouri (NCN). Herpetological Review, 27, 28.","McCranie, J. R. & Wilson, L. D. (2002) The Amphibians of Honduras. Vol. 19. Society for the Study of Amphibians and Reptiles, Contributions in Herpetology, Ithaca, New York. i - x, pp. 1 - 625.","McCranie, J. R. & Castaneda, F. E. (2005) The herpetofauna of Parque Nacional Pico Bonito, Honduras. Phyllomedusa, 4, 3 - 16. http: // dx. doi. org / 10.11606 / issn. 2316 - 9079. v 4 i 1 p 03 - 16","Townsend, J. H., Butler, J. M., Wilson, L. D. & Austin, J. D. (2010) A distinctive new species of moss salamander (Caudata: Plethodontidae: Nototriton) from an imperiled Honduran endemism hotspot. Zootaxa, 2434, 1 - 16.","Kohler, G. (2011) Amphibians of Central America. Herpeton Verlag, Offenbach, 379 pp.","Townsend, J. H., Medina-Flores, M., Murillo, J. L. & Austin, J. D. (2011 a) Cryptic diversity in Chortis Highland moss salamanders (Caudata: Plethodontidae: Nototriton) revealed using mtDNA barcodes and phylogenetics, with a new species from eastern Honduras. Systematics and Biodiversity, 9, 275 - 287. http: // dx. doi. org / 10.1080 / 14772000.2011.611188","Rafaelli, J. (2013) Les Urodeles du Monde, 2 nd ed. Penclen Edition, Plumelec, France.","Townsend, J. H., Medina-Flores, M., Reyes-Calderon, O. & Austin, J. D. (2013) A new Nototriton (Caudata: Plethodontidae) from Montana de Botaderos National Park in northeastern Honduras. Zootaxa, 3666 (3), 358 - 368. http: // dx. doi. org / 10.11646 / zootaxa. 3666.3.6","Dubois, A. & Raffaelli, J. (2012) A new ergotaxonomy of the order Urodela Dumeril, 1805 (Amphibia, Batrachia). Alytes 28, 77 - 161.","Smithe, F. B. (1975 - 1981) Naturalist's Color Guide. Part I. Color Guide. American Museum of Natural History, New York, 182 color swatches.","Holdridge, L. R. (1967) Life Zone Ecology. Revised Edition. Tropical Science Center, San Jose, Costa Rica, 206 pp.","Townsend, J. H. (2014) Characterizing the Chortis Block Biogeographic Province: Geological, physiographic, and ecological associations and herpetofaunal diversity. Mesoamerican Herpetology, 1, 204 - 252.","McCranie, J. R & Townsend, J. H. (2011) Description of a new species of worm salamander (Caudata, Plethodontidae, Oedipina) in the subgenus Oedopinola from the central portion of the Cordillera Nombre de Dios, Honduras. Zootaxa, 2990, 59 - 68."]}

Genome sizes (expressed as C-values, or haploid genome sizes) of six species of Honduran plethodontid salamanders (one species of Nototriton and five of Bolitoglossa) vary greatly. Nototriton has a moderate-sized genome (29.2 pg) relative to other species of salamanders. Genome sizes in the species of Bolitoglossa span a range of 24 pg (∼23.4 gigabases) of DNA and include the largest genomes (83.7 pg) reported for the genus and for the family Plethodontidae. A phylogenetic analysis indicates that genome evolution in this group of salamanders featured mainly large increases in the mass of nuclear DNA. We propose that these evolutionary changes in genome size reflect random drift in small, isolated populations in the highlands of Central America.

The systematic study of Central American moss salamanders has been a challenge to researchers due to their cryptic nature and subsequent difficulty in sampling. In an effort to elucidate relationships among moss salamanders from the Chortis Highlands, we quantified intra- and interspecific variation using data from the mitochondrial genes 16S (the amphibian barcoding gene), cytochrome b (cyt b; widely used in phylogenetic studies of neotropical salamanders), and cytochrome oxidase subunit I (COI, the universal metazoan barcoding gene). All three loci exhibit a bimodal ‘barcoding gap’ between ranges of intra- and interspecific variation, demonstrating unambiguous species boundaries with respect to the mitochondrial loci. A barcoding approach and Bayesian and maximum likelihood analyses of the mtDNA dataset revealed that the nominal species N. barbouri represents a species complex, with two of three populations assigned to this taxon being paraphyletic with respect to a sample from the type locality. These ...

Despite several recent studies, the phylogeny of plethodontid salamanders is not yet fully resolved and the phylogenetic positions of several key genera, especially Aneides, Hemidactylium, Hydromantes and Karsenia, are contentious. Here we present a combined dataset of complete mitochondrial genomes and three nuclear loci for 20 species (16 genera) of plethodontids, representing all major clades in the family. The combined dataset without mitochondrial third codon positions provides a fully resolved, statistically well-supported tree. In this topology two major clades are recovered. A northern clade includes Aneides, Desmognathus, Ensatina, Hydromantes, Karsenia, Phaeognathus and Plethodon, with Plethodon being the sister taxon to the rest of the clade. Hydromantes and Karsenia are sister taxa, and Aneides is recovered as the sister taxon to Ensatina. Desmognathus + Phaeognathus form the sister taxon to Aneides + Ensatina. An eastern/southern clade comprises two subclades. One subclade, the spelerpines (Eurycea, Gyrinophilus, Pseudotriton, Stereochilus, Urspelerpes) is the sister taxon to a subclade comprising Hemidactylium, Batrachoseps and the tropical plethodontids (represented by Bolitoglossa, Nototriton and Thorius). In this topology Hemidactylium is well-supported as the sister taxon to Batrachoseps. Only when mitochondrial third codon positions are included using maximum likelihood analysis is Hemidactylium recovered as the sister taxon to Batrachoseps + tropical genera. Hypothesis testing of alternative topologies supports these conclusions. On the basis of these results we propose a conservative taxonomy for Plethodontidae.

I describe a new species of salamander of the genus Nototriton from Cerro Saslaya, eastern Nicaragua. It differs from all other species of the genus Nototriton by unique characteristics of coloration, morphometrics, and osteology. Resumen Una nueva especie de salamandra del genero Nototriton es descrita como proveniente de Cerro Saslaya, al este de Nicaragua. Se diferencia de todas las otras especies del genero Nototriton por caracteristicas unicas de coloracion, morfometria y osteologia.

Nototriton mime sp. nov. Figures 3 A–D Holotype. USNM 579870 (Fig. 3 A), an adult male from Cerro Ulloa (15.3833 °N, 86.0399 °W), Parque Nacional Montaña de Botaderos, 1,705 m, Departamento de Olancho, Honduras, collected 16 April 2011 by M. Medina- Flores, O. A. Reyes-Calderón, and J. H. Townsend; original field number JHT 3400; GenBank accession numbers KC 905090 (16 S) and KC 905094 (cyt b). Paratopotypes. Three (same collectors as holotype); a female (Fig. 3 B; MVZ 269306) and a juvenile (Fig. 3 D; USNM 579872), collected 16 April 2011, 1,710 m, 15.3843 °N, 86.0396 °W; and a male (Fig. 3 C; USNM 579871), 17 April 2011, 1,720 m elevation, 15.3874 °N, 86.0467 °W. Diagnosis. A member of the genus Nototriton diagnosed by possessing 13 costal grooves (> 16 costal grooves in Oedipina), hands and feet longer than broad (hands and feet broader than long in Bolitoglossa), and small nares (0.007–0.010 NL/SL; 0.017–0.029 NL/SL in Cryptotriton and Dendrotriton). Distinctiveness of the new species and its assignment to the Nototriton barbouri species group is supported by analysis of sequence fragments from the mitochondrial genes 16 S and cyt b (Fig. 2). Nototriton mime is unique among described congeners in the Chortís Highlands by having a yellow-brown dorsal coloration in adult males, as well as relatively longer front and hind limbs (FLL/SL 0.195–0.246, HLL/SL 0.224–0.254; versus 0.142–0.174 and 0.153 –0.200 in N. barbouri, 0.148–0.151 and 0.166–0.180 in N. brodiei, 0.137–0.160 and 0.158–0.181 in N. lignicola, 0.156–0.183 and 0.164– 0.211 in N. limnospectator, 0.179–0.191 and 0.204–0.218 in N. picucha, 0.172 and 0.178 in N. stuarti, and 0.160 and 0.197 in N. tomamorum). The new species may be further differentiated from the other species of Nototriton found in the Chortís Highlands as follows: from N. brodiei by having a broader head (HW/SL 0.121–0.138, versus 0.120), larger nares (NL/SL 0.007–0.010, versus 0.004–0.005), a shorter tail (TL/SL 0.698 –1.117, versus 1.420– 1.440), and fewer maxillary teeth (27–46, versus 60–62); from N. lignicola by having a broader head (HW/SL 0.121–0.138, versus 0.103–0.118); from N. limnospectator by having a broader head (HW/SL 0.121–0.138, versus 0.095–0.118) and larger nares (NL/SL 0.007–0.010, versus 0.003); from N. picucha by having a narrower head (HW/SL 0.121–0.138, versus 0.140–0.148); from N. saslaya by having narrower hind feet (HFW/SL 0.044 – 0.050, versus 0.075–0.091), larger nares (NL/SL 0.007–0.010, versus 0.002–0.003), and a higher number of maxillary (27–46, versus 17–22) and vomerine (12–24, versus 3–11) teeth; from N. stuarti by having a slightly narrower head HW/SL 0.121–0.138, versus 0.138), a shorter tail (TL/SL 0.698 –1.117, versus 1.264), and smaller nares (NL/SL 0.007–0.010, versus 0.012); and from N. tomamorum by having well-developed digits with subdigital pads (versus hands and feet syndactylous and lacking subdigital pads), a narrower head (HW/SL 0.121– 0.138, versus 0.145), and smaller nares (NL/SL 0.007–0.010, versus 0.018). Description of holotype. An adult male (SL = 31.8 mm, total length = 67.3 mm) with a slender body and reduced limbs. The head is rounded, slightly broader than the body; nostrils are relatively small (NL/SL= 0.010), and the snout is acutely rounded and of moderate length. Nasolabial grooves are visible but slight, and nasolabial protuberances are not apparent. The eyes are relatively large and protuberant, and the elongate parotoid glands are relatively flat and not well-defined. There are 46 maxillary teeth, 3 slightly enlarged premaxillary teeth in line with the maxillaries, and 24 vomerine teeth. The limbs are short (CLL/SL= 0.40), with the adpressed limb interval approximately 4.5 costal grooves. The hands and feet are narrow with well-developed digits that bear poorly developed subdigital pads. The relative length of the digits is I Measurements of holotype (mm). SL 31.8; AG 16.9; TW 3.3; HL 5.8; HW 3.6; TL 35.5; HLL 6.7; FLL 5.8; CLL 12.5; FFW 0.8; HFW 0.9; NL 0.3; EL 1.8; EW 0.9; IOD 0.7; anterior rim of orbit to snout 1.4; distance separating internal nares 1.1; distance separating external nares 1.6; snout projection beyond mandible 0.3; tip of snout to axilla 9.0; snout to anterior edge of vent 27.8; tail depth at basal constriction 2.7; tail width at basal constriction 2.6. Color in life of adult male holotype. Description of the coloration of the holotype based on a series of color photographs taken in life: dorsal surfaces of paratoid region of head, body, and tail Chamois (84); lateral surfaces mottled Ground Cinnamon (270), with Dark Grayish Brown (284) spots beginning on the centrolateral portion of costal groove 4 and becoming larger posteriorly, with one large blotch present centrolaterally from costal groove 12 to the hind limb; narrow middorsal stripe Pale Pinkish Buff (3), with faint herringbone pattern evident dorsally; dorsal and lateral surfaces of head mottled Vandyke Brown (282) with some Smoky White (261) and Pearl Gray (262) stippling above the mouth; dorsal surface of limbs Sepia (286) with Cinnamon-Rufous (31), Yellow-Ocher (14), and Chamois (84) stippling, becoming more profuse towards body; dorsal surface of tail with Pale Mauve (204) spots beginning about halfway down the length of the tail, with tail becoming almost completely Pale Mauve (204) towards the tip; ventral surface of body with a Dark Mauve (208) to Cyanine Blue (189) cast, with scattered Smoky White (261) and Light Lavender (201) stippling; ventral ground color of tail Fuscous (283), darker than body, with a series of three pairs of Salmon Color (58) blotches behind the vent with spacing equivalent to that of the costal grooves, distal portion of ventral surface of tail with Smoky White (261) and Pale Mauve (204) stippling; color of iris Gem Ruby (65). Variation. In all individuals except the holotype, the tail is shorter than the body (TL/SVL= 0.780 in USNM 579871, a male, 0.817 in MVZ 269306, a female, and 0.698 in USNM 579872, a juvenile) In each case, the tail does not appear to be incomplete, and thus the significance of the degree of variation seen in tail length is not clear. The male paratype (USNM 579871) has 27 maxillary, 5 premaxillary, and 12 vomerine teeth; the female paratype (MVZ 269306) has 37 maxillary, 7 premaxillary, and 17 vomerine teeth. The female (MVZ 269306) and juvenile (USNM 579872) paratypes both exhibit a different color pattern than that seen in the male holotype (USNM 579870) and paratype (USNM 579871). The adult female paratype MVZ 269306 has the following coloration in life: dorsal surface of paratoid region of head and body Hazel (26) with darker Brick Red (36) and paler Robin Rufous (29) blotches forming a herringbone pattern, with a narrow Robin Rufous (29) middorsal stripe; irregular Dark Buff Yellow (54) dorsolateral stripes originate approximately halfway down the body and become better defined and continue onto the tail, bordered laterally by a series of small Vandyke Brown (281) blotches positioned on costal grooves 4, 5, 8, 9, and 10, and becoming a continuous blotch over costal grooves 11, 12, and 13 that extends onto the proximal half of the tail; lateral surfaces of body mottled Dark Buff Yellow (54) and Robin Rufous (29) with Pale Bluish Gray (287) stippling; dorsal surface of head with Raw Umber (280) mottling on the parietal region and between the eyes, lateral surfaces of head Russet (44) with Pale Bluish Gray (287) stippling, ventral surface Glaucous (291) with scattered small Pale Bluish Gray (287) spots; ventral surface of body Vandyke Brown (282) with scattered Smoky White (261) and Pale Bluish Gray (287) stippling most prominent near the head and along the centerline of the venter; dorsal surface of tail mottled Hazel (26) and Robin Rufous (29), with Raw Umber (280) to Vandyke Brown (282) lateral and ventral surfaces with Smoky White (261) and Pale Bluish Gray (287) stippling; iris Mahogany Red (34). Etymology. The specific epithet “ mime ” is a noun in apposition to the generic name, given to honor our late friend Arquimides Gabriel Rosales Martinez, or “ Mime ” (pronounced me-may), a young Honduran biologist passionate about amphibians. Mime and his sister, Novy Hortensia Rosales Martinez, were killed by a drunken driver on 17 December 2010 in Tegucigalpa. Geographic distribution. This species is known only from the immediate vicinity of the type locality, 1,705– 1,720 m on Cerro Ulloa, the highest ridge in the Sierra de Botaderos, forming the border between the departments of Colón and Olancho (Figs. 1, 3). Natural history. The habitat at the type locality is tropical montane cloud forest, in the Lower Montane Wet Forest formation. The male holotype (USNM 579870; Fig. 3 A) was active at night among the leaves of an orchid growing on a small stump on the ground along a wet ridge at 1,705 m elevation. The male paratype (USNM 579871; Fig. 3 C) was collected from underneath a small log embedded in the ground at 1,720 m elevation. The female paratype (MVZ 269306; Fig. 3 B) was found in a bromeliad on the ground at 1,705 m elevation, near the holotype. The juvenile paratype (USNM 579872; Fig. 3 D) was collected during the late afternoon from within a small bromeliad approximately 2 m above the ground on a large fallen branch at 1,710 m elevation. Conservation status. Based on the extremely limited extent of intact cloud forest around the only known locality (approximately 300 hectares), and the continuing deforestation surrounding this locality, we recommend Nototriton mime be immediately considered Critically Endangered (CR B 1 ab[i, ii, iii]+ 2 ab[i, ii, iii]), following the IUCN Red List Criteria (IUCN 2012). The broadleaf forests of the Sierra de Botaderos are approximately 33,000 hectares in total extent, and ranges from 100 to 1,724 m elevation. However, less than 1,000 total hectares of forest above 1,500 m elevation exists, and is fragmented into approximately a dozen discrete patches along the spine of the cordillera. The type locality, Cerro Ulloa, is one of these patches, and is the highest elevation forest in the mountain range. Cerro Ulloa is already deforested up to and above 1,500 m elevation on the southern slopes (Fig. 4 A), and preexisting trails made by farmers cross the summit of the peak (Fig. 4 B) and access inholdings deep with the forest on the Colón side of the ridge. The approximately 300 hectares of intact cloud forest habitat surrounding the type locality are already under direct threat from migratory agriculture and cattle ranching, and, most recently, are indirectly threatened by the dramatic increase in international narcotrafficking in adjacent portions of Colón and Olancho. The recent declaration of the remaining premontane and lower montane rainforests as a national park is a positive development; however, the remote location of the park and high-risk nature of conflicts related to enforcing protection of its remaining forests make conservation here, and throughout eastern Honduras, a resourceintensive challenge. Remarks. With the description of Nototriton mime, there are 17 named species of Nototriton distributed from Guatemala to Costa Rica, six of which are endemic to Honduras (Townsend et al. 2011; Boza-Oviedo et al. 2012). Each of these six endemic species have distributions restricted to cloud forest habitat in the upper reaches of isolated mountain ranges, with only one species (N. limnospectator) known to occur at premontane elevations as low as 800 m. In addition to these six species, there are two as yet undescribed species currently referred to the taxon N. barbouri, one from Parque Nacional Pico Bonito (N. sp A in Townsend et al. 2011) and one from Refugio de Vida Silvestre Texíguat (N. sp. B, op. cit.). The coloration seen in Nototriton mime appears to be the first documented case of sexual dichromatism in the genus Nototriton, and one of few documented cases in tropical salamanders. Another endemic Honduras salamander, Bolitoglossa diaphora, from the Sierra de Omoa in northwestern Honduras, also exhibits marked dichromatism, with males being bluish-gray and females being a mottled red-orange coloration (McCranie & Wilson 1995).

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...

Nototriton matama sp. nov. Matama Moss Salamander Figure 5 Holotype. UCR 20215, an apparently mature female from the southeastern end of the Fila Matama (coordinates: 9.8071 �� N, 83.1683 �� W) at an elevation of 1300 m, collected on 30 October 2007, by Eduardo Boza-Oviedo and Ruth Delgado. Paratypes. UCR20168, 20169, 20171, same data as holotype. Diagnosis. Assigned to Nototriton because it has a well-developed sublingual fold and has fewer than 17 vertebrae in the trunk, and to the picadoi group based on mtDNA sequence data and on the basis of having rounded digital tips rather than pointed ones (as in the richardi group). A small member of the Nototriton picadoi group distinguished from all other members of the group by relatively enlarged and elongated nostrils and small, very narrow hands and feet and narrow heads; distinguished from N. picadoi by smaller size (holotype 23.6 mm SL, N. picadoi reaches 32 mm SL), relatively larger nostril (0.02 times SL, vs. 0.010���0.016 in N. picadoi), and very narrow feet (0.05 times SL, vs. 0.06���0.07 in N. picadoi) with pointed outer toe tips (vs. rounded toe tips); distinguished from members of the N. richardi group by discrete columnar digits not fused together and by rounded rather than pointed tips of the longest digits. Description. A diminutive, slender species compared to other members of its genus. Sole adult specimen (based on size and proportions), the holotype, has a SL of 23.6 mm. Because the holotype is a female, typically the larger sex, maximum size of this species is expected to be not much larger than this specimen. Head small and narrow (SG/SL= 0.17; HW/SL= 0.14) with broadly truncated snout. Nostrils enlarged and elongated, with maximal dimension of 0.5 mm. One can see into nostrils from a dorsal view; nostrils oriented mainly frontally. Eyes relatively large and protuberant, visible protruding from side of head when viewed ventrally. Head only moderately differentiated from trunk, mainly by somewhat enlarged parotoid glands located in temporal region of head. Parotoid glands clearly evident as swollen, lightly pigmented oval structures. Teeth moderately abundant; PMT 4, MT 31, VT 11 in holotype. Trunk slender, limbs relatively short (0.18 SL, LI 5). Slender tail only slightly tapered before a break. Hands and feet bear well-formed, columnar digits that are only slightly webbed basally. Longer digits terminate in rounded tips, but shorter digits have bluntly pointed tips. Fingers, in order of decreasing length, are 3 - 2-4 - 1; toes are 3-4 - 2-5 - 1. Measurements (in mm), limb interval and tooth counts of the female holotype (Table 2). HW 3.2, SG 4.2, HD 1.8, EW 0.6, EL 1.3, ES 0.8, ED 1.0, IC 1.8, IO 0.9, length of groove extending posteriorly from eye 1.0, distance between nuchal groove and gular fold 0.9, SF 6.8, IN 0.4, SP 0.2, SL 23.6, SAV 21.8, AX 13.4, LI 5, tail broken at 11.0, tail width at base 1.6, tail depth at base 1.8, FLL 3.7, HLL 4.2, HAW 1.0, FW 1.2, T 5 0.2, T 3 0.6, parotoid width 0.5, parotoid length 1.3, nostril diameter 0.5. Number of teeth: PMT 4, MT 15 / 16, VT 5 / 6. Coloration of the holotype in life. A colorful individual with a generally light golden brown dorsal coloration. Light dorsal band bordered by short streaks of white and tan that constitute an irregular border, especially in pelvic area. Tail more uniformly golden in coloration and a little lighter than trunk. Parotoid region at back of head pale golden. Along generally darker flanks are some dark brown speckles. Broad band of light coloration under dark flanks. Coloration of the holotype in alcohol. Colorful specimen more sharply differentiated into light and dark areas than in life. Specimen grey-brown to tan. Two pale temporal/paratoid patches. Distinct pale stripe extends from shoulder to tail, which is bright yellow and brown. Some suffusion of melanin present on trunk. Ventrolateral parts of trunk cream-colored. Dark interrupted dorsolateral line of pigment extends from shoulder to pelvis. Venter dark with whitish streaks in two ragged rows. Yellow patch in temporal areas descends to gular area in front of gular fold but not on midgular area. Gular area blackish but lighter than midventer region. Small speckles of white on ventral surfaces. Hint of herringbone pattern of dark chevrons present in dorsal stripe. Habitat and range. The species is known only from the type locality along the Matama ridge of the Caribbean slope of the Chirrip�� Massif. The locality has mature cloud forest that includes members of: Araceae (e.g. Philodendron, Anthurium, Monstera), Begonia (Begoniaceae), palms (Arecaceae), Ericaceae, Melastomataceae, Marantaceae, Urticaceae (e.g. Pilea), Acanthaceae, Cyclanthaceae (e.g. Cyclanthus, Carludovica), Rubiaceae, Heliconia (Heliconiaceae), Piperaceae (e.g. Piper), bromeliads, ferns (e.g. Cyatheaceae and no tree ferns), and bryophytes. Humidity was at or near 100 % during the time spent at the site. The specimens were found during daytime in moss mats at 0.95���2.1 m above ground and 400���600 m from the nearest stream. One specimen was found between a plant stem and the moss, while the others were within the moss (35���90 mm wide), some in moss in vertical parts of the trunk and others in moss hanging off the branches. Plagiochila spp was the most frequent bryophyte in the moss mats. The temperature ranged from 16���17 ��C within moss mats and 17���18 ��C in the air. Three specimens were found in the same tree. Etymology. The species was discovered near the terminus of the Fila de Matama, a large mountain ridge that arises as a part of Cerro Chirrip��, the highest mountain in Costa Rica. The scientific name is a noun in apposition., 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 48-49, DOI: 10.5281/zenodo.211943

Nototriton tomamorum new species (Figs. 2, 3) Holotype: A female (UF 155377) from 2.5 km NNE of La Fortuna (15 �� 25.965 ���N, 87 �� 18.556 ���W), 1550 m, Refugio de Vida Silvestre Texiguat, Departamento de Yoro, Honduras, collected 9 April 2008 by J. Slapcinsky and the field team of J. M. Butler, L. P. Ketzler, N. M. Stewart, J. H. Townsend, and L. D. Wilson. Original field number JHT 2437. Diagnosis. A small member of the genus Nototriton (SVL= 26.9 mm; Table 2) based on having 13 costal grooves (> 16 costal grooves in Oedipina), hands and feet longer than broad (hands and feet broader than long in Bolitoglossa), and nares that are smaller than most Cryptotriton and Dendrotriton (Fig. 3 A, B; 0.018 NL/ SVL; 0.020���0.029 NL/SVL in Cryptotriton [except some individuals of C. veraepacis] and Dendrotriton). Cryptotriton veraepacis has nares ranging from 0.017���0.027 NL/SVL (mean 0.022), and can be differentiated from N. tomamorum by having a uniformly dark gray ventral surface (ventral surface pale with gray flecks in N. tomamorum; Fig. 2). Generic placement in Nototriton is also strongly supported by sequence data from the mitochondrial genes 16 S and cob (Table 3; Fig. 4). This new species is distinguished from all other Nototriton, except N. richardi and N. tapanti, by having syndactylous hands and feet (Fig. 3 C, D; hands and feet with free, differentiated toes in all other species) and relatively large nares (Fig. 3 A, B; 0.018 NL/SVL versus 0.010���0.016 in N. picadoi, 0.003���0.014 in N. abscondens, 0.012 in N. stuarti, 0.005���0.011 in N. barbouri, 0.006���0.009 in N. lignicola, 0.004���0.009 in N. guanacaste, 0.004���0.005 in N. brodiei, 0.003 in N. limnospectator, 0.003 in N. major, and 0.002���0.003 in N. saslaya). Nototriton tomamorum can be further differentiated from members of the N. barbouri group by having a broader head (0.145 HW/SVL versus 0.138 in N. stuarti, 0.104���0.132 in N. barbouri, 0.120 in N. brodiei, 0.103���0.118 in N. lignicola, and 0.095���0.118 in N. limnospectator) and fewer maxillary teeth (26, versus 36 in N. stuarti, 41���54 in N. barbouri, 42���55 in N. limnospectator, 46���54 in N. lignicola, and 60���62 in N. brodiei), from members of the N. picadoi group by having a relatively shorter tail (0.911 TL/SVL, versus 1.441 in N. major, 1.123���1.344 in N. picadoi, 1.013��� 1.365 in N. abscondens, 1.210���1.337 in N. guanacaste, and 1.10���1.30 in N. gamezi) and narrower feet (0.037 HFW /SVL, versus 0.058���0.071 in N. abscondens, 0.059 in N. major, 0.060���0.070 in N. picadoi, and 0.066��� 0.072 in N. guanacaste), and from N. saslaya by having shorter forelimbs (0.160 FLL/SVL, versus 0.194��� 0.210 in N. saslaya) and hind limbs (0.197 HLL /SVL, versus 0.217���0.244 in N. saslaya) and narrower feet (0.037 HFW /SVL, versus 0.075���0.091 in N. saslaya). The new species also differs from N. richardi and N. tapanti in having a pale ventral surface mottled with gray chromatophores (ventral surface brown with dark flecks in N. richardi and dark brown in N. tapanti), by having a tail that is shorter than the snout-vent length (0.91 TL/SVL, versus 1.072���1.482 in N. richardi and 1.205 in N. tapanti), longer forelimbs (0.160 FLL/SVL, versus 0.140���0.146 in N. richardi and 0.147 in N. tapanti), longer hind limbs (0.197 HLL /SVL, versus 0.174��� 0.187 in N. richardi and 0.174 in N. tapanti), and narrower feet (0.037 HFW /SVL, versus 0.044���0.050 in N. richardi and 0.041 in N. tapanti). This new species is also well differentiated from all other species of Nototriton based on mitochondrial sequence data (Table 3), and is 3.6 ���6.0% divergent on 16 S and 10.2��� 15.1 % divergent on cob from all other congeners. Description of holotype. Nototriton tomamorum is known only from a single, presumably female (mental gland and cloacal papillae absent) specimen, preserved with its mouth open and tongue extended, and is a relatively small member of the genus (SVL= 26.9 mm, total length= 51.4 mm) with a slender body and reduced limbs. Its head is rounded and slightly broader than the body, and the nostrils are relatively large (NL/ SVL= 0.018), and the snout is rounded and of moderate length (Fig. 3 A, B). The nasolabial protuberances are apparent but not well developed, and barely extend below the upper lip line. The eyes are relatively large and protuberant, and the parotoid glands appear large but not well defined. The teeth are exceedingly small; there are approximately 26 maxillaries, 4 premaxillaries set slightly forward from the maxillary teeth, and 11 vomerines; vomerine teeth are arranged in two short medially-positioned arches. The limbs are short (CLL/ SVL= 0.36), with the adpressed limbs being separated by approximately 5.5 costal grooves. The hands and feet are narrow and have poorly-developed, poorly-differentiated digits that are fused and lack subdigital pads (Fig. 3 C, D). The free tips of digits III on the hands and III and IV on the feet are pointed, and digits I, II and IV on the hands and I, II, and V on the feet are very short and essentially completely fused, being demarcated by shallow grooves on the dorsal side of the feet. The relative length of the digits is IMeasurements of holotype (in mm). SVL 26.9; AG 15.2; TW 3.6; HL 4.8; HW 3.9; TL 24.5; HLL 5.3; FLL 4.3; CLL 9.6; HFL 1.7; HFW 1.2; NL 0.5; eye length 1.6; eye width 1.2; interorbital distance 1.4; anterior rim of orbit to snout 1.1; distance separating internal nares 0.8; distance separating external nares 1.8; snout projection beyond mandible 0.6; tip of snout to axilla 7.7; distance from axilla to groin 15.2; snout to anterior edge of vent 24.9; tail depth at basal constriction 2.8; tail width at basal constriction 2.4. Coloration of holotype. Dorsal surfaces of head, body, and tail medium grayish brown, with profuse pale chromatophores laterally, becoming less abundant dorsolaterally. The head has some pale mottling on the top of the snout, and two irregular lines of pale chromatophores extending from the lateral region above the forelimbs onto the posterior portion of the head and parotoid glands. There is a very thin, pale middorsal stripe, with a herringbone pattern with lines extending from the middorsal stripe posteriorly. There is an indistinct dark dorsolateral stripe starting about one-third the way down the trunk and extending onto the proximal one-third of the tail. Ventral surface of head, body, and tail cream, mottled with dark gray chromatophores, becoming somewhat more profuse toward the distal end of the tail. Etymology. The specific name ��� tomamorum ��� means ���belonging to the Tomams.��� Tomams are the highest level of deities recognized in the belief system of the indigenous Tolupan of Honduras, of which there are four: Tomam Pones Popawai (Grandfather Tomam), his wife Tomam Pones Namawai (mother of all that exists), and their children Tomam the Elder and Tomam the Younger (Chapman 1992). This name is given in recognition that the Tolupan are the traditional inhabitants of this area and that this new species is known only from the Cordillera Nombre de Dios, or ���Name of God Mountains,��� a name which, ironically, was given by 15 th century Spanish explorers. Natural history. The single known specimen of Nototriton tomamorum was collected during the daytime from leaf litter packed onto a rock ledge alongside a small creek at about 1550 m elevation in the Lower Montane Wet Forest formation (Fig. 5). This species is presumably endemic to the vicinity of the type locality, and is likely restricted to the Lower Montane Wet Forest around the neighboring peaks of Cerro San Francisco and Cerro Texiguat. The locality can be characterized as a narrow, steep-sided canyon with riparian forest, and more xeric upland pine-oak forest dominating the terrain above the canyon. The pine-oak forest transitions to mesic cloud forest (or remnant cloud forest) uphill from this locality. This species is sympatric with a congener, N. barbouri, and three other plethodontids: Bolitoglossa dofleini, Bolitoglossa porrasorum and Oedipina gephyra. The canyon where the holotype of N. tomamorum was collected is also the type locality of the hylid frog Isthmohyla insolita, and a known locality for the anurans Lithobates maculatus, Plectrohyla guatemalensis, and Ptychohyla spinipollex. Conservation status. Based on the criteria of the IUCN (2001), Nototriton tomamorum should be classified as Critically Endangered (B 1 ab[iii]+ 2 ab[iii]) due to the extremely limited extent of its known distribution and the direct threat to the remaining habitat around that locality. The primary threat to the type and only known locality for this species is from illegal timber extraction and wholesale slash-and-burn clearing of forest for bean cultivation once it has been depleted of valuable hardwoods, such as mahogany (Fig. 6). Systematic relationships. Nototriton tomamorum bears the strongest morphological resemblance to members of the N. richardi group (Table 2), which are endemic to the central highlands of Costa Rica. The two species, N. richardi and N. tapanti, are the most morphologically distinctive members of the genus, differing from all other Nototriton by having syndactylous feet, relatively large nostrils, and frontal processes of the premaxillary arising separately from the maxillary (Good & Wake 1993; Papenfuss & Wake 1987; Savage 2002). According to Savage (2002), N. richardi is distributed from 1370 to 1800 m elevation along the Atlantic slope of the Cordillera Central, and N. tapanti at a single locality at 1300 m elevation in the northern portion of the Cordillera de Talamanca. These localities are situated in the central montane spine of Costa Rica in Lower Central America and are separated widely from the type locality of N. tomamorum by over 670 airline km and much of the central highlands of Honduras and the lowland Nicaraguan Depression. Despite the morphological similarities between N. tomamorum and members of the N. richardi group, our phylogenetic analysis (Fig. 4) does not support the inclusion of N. tomamorum in the N. richardi group. Preliminary phylogenetic analysis of fragments from two mitochondrial genes (16 S and cob) weakly supports (posterior probability = 0.55, bootstrap support = 58 %) N. tomamorum as sister taxon to the rest of the N. barbouri group (Fig. 4). We also recovered N. richardi as the weakly supported sister (posterior probability = 0.58, bootstrap support = 59 %) to the N. picadoi group, a finding consistent with previous phylogenetic hypotheses for Nototriton and N. richardi (Garc��a-Par��s & Wake 2000; Wiens et al. 2007; Adams et al. 2009). Sequence data from the holotype of Nototriton tomamorum are at least 12.5 % (16 S) and 20.6 % (cob) divergent from representatives of morphologically-similar genera of Central American plethodontids (Cryptotriton, Dendrotriton, and Oedipina), but less than 6.0% (16 S) and 15.1 % (cob) divergent from other species of Nototriton, supporting placement of N. tomamorum in this genus (Table 3). Within the genus Nototriton, the new species demonstrates differing patterns of sequence divergence on the two genes sampled. For 16 S, N. tomamorum is closest to N. limnospectator (3.6���3.8 %), a biogeographically-logical situation given the relative geographical proximity of RVS Texiguat to the type locality of N. limnospectator, approximately 100 airline km to the WSW in Parque Nacional Monta��a de Santa B��rbara; however N. tomamorum is 12.4 % divergent from N. limnospectator for cob, and for that gene is closer to the Costa Rican species N. abscondens (10.2 %) and N. gamezi (11.5 %) than to any other member of the N. barbouri group (Table 3). Given the morphological differentiation and lack of strong phylogenetic support, we refrain from assigning N. tomamorum to any recognized species group pending acquisition and analysis of additional molecular data from mitochondrial and nuclear genes. Our phylogenetic analyses included taxa and populations of Nototriton not available in previous studies (Garc��a-Par��s & Wake 2000; Wiens et al. 2007), including a sample from the vicinity of the type locality of N. barbouri (���Portillo Grande, Yoro, Honduras ��� [Schmidt 1936]). Inclusion of these data reveals the taxon N. barbouri to be paraphyletic (Fig. 4), with strong support for N. barbouri sensu stricto as the sister species of N. limnospectator and for the remaining populations assigned to N. barbouri (N. sp. in Fig 4) from the Cordillera Nombre de Dios to be sister to N. brodiei from NE Guatemala and NW Honduras. We are currently preparing an intergrative systematic revision of the N. barbouri group that will use molecular and morphological analyses to resolve the taxonomic status of populations assigned to N. barbouri. The herpetofauna of the RVS Texiguat is characterized by a remarkable degree of endemism (McCranie & Casta��eda 2004 a; Wilson & McCranie 2004 b). Currently, 39 species are known from 1550 m in elevation and above in this refuge. Of this number, 27 are considered to be conservation priority species (Table 4), defined as those species judged to occupy one of three IUCN categories Critically Endangered, Endangered, or Vulnerable. Nine of the 27 are considered Critically Endangered, 12 are Endangered, and six are Vulnerable; 19 (70.4 %) of these species (four salamanders, seven anurans, four lizards, and four snakes) are endemic to Honduras (Table 4). Of the remainder, six are endemic to Nuclear Central America and two have widespread distributions. As indicated by Townsend & Wilson (In press), 91 species of amphibians and reptiles are endemic to Honduras, thus the Texiguat figure is 20.9 % of the total. Therefore, RVS Texiguat qualifies as a major center of endemism in Honduras, as well as in the whole of eastern Nuclear Central America (as defined by Campbell, 1999). The Environmental Vulnerability Scores, as tabulated by Townsend and Wilson (in press) and indicated in Table 4, indicate that the values for the Texiguat conservation priority species range from 9 to 17. Most of these species involved (16) are considered high vulnerability species (59.3 %), nine (33.3 %) are medium vulnerability species, and only two (7.4 %) have scores lying at the upper end of the range for low vulnerability species (Table 4). In summary, of the 27 species judged to be of conservation priority, 19 are endemic to Honduras and 16 are of high vulnerability. Based on the analysis above, Refugio de Vida Silvestre Texiguat is an area of immense importance for the conservation of the Honduran herpetofauna, especially its endemic component. Unfortunately, little attention has been paid to RVS Texiguat by conservation authorities past its initial designation as a protected area in 1987 (Wilson et al. 2001). Since 1991, ten species (including the present one) with type localities in the area have been described: Anolis kreutzi, A. purpurgularis, A. yoroensis, Celestus scansorius, Geophis damiani, Isthmohyla insolita, Nototriton tomamorum, Oedipina gephyra, Omoadiphas texiguatensis, and Rhadinaea tolpanorum (McCranie et al. 1993 a, b; Wilson et al. 1998; McCranie & Casta��eda 2004 a). Given the pattern of these finds, we expect that additional new taxa remain to be described from RVS Texiguat. During this period, significant environmental damage has been observed, most of it apparently as a consequence of the illegal extraction of mahogany and the subsequent conversion of the remaining forest to cropland for use in subsistence agriculture (Fig. 6). Most recently during our visit in April 2008, we observed that a large stretch of forest upstream from the type locality of Nototriton tomamorum and Isthmohyla insolita had been clear-cut down to the streamside and planted to beans (Fig. 6 C). Damage to the forests in this refuge also has occurred by natural means. The stream that constitutes the type locality of Nototriton tomamorum and Isthmohyla insolita (Fig. 5) was severely damaged by Hurricane Mitch in 1998. At that time, the habitat supporting I. insolita was scoured by high waters in the narrow canyon. The moss masses clinging to the branches of small trees overhanging the stream, within which the adults laid their eggs, were stripped away. Our trip in 2008 evidenced that the moss has yet to be reestablished as we found it in 1991 when I. insolita was discovered. Whether the reestablishment of these conditions will occur is doubtful. Isthmohyla insolita has been located in one or two nearby streams (McCranie & Casta��eda 2007), but the same anthropogenic activities detailed above also are occurring there. It is apparent that the natural habitat of this hotspot for herpetofaunal endemism will continue to succumb to the forces of habitat destruction unless steps are taken to protect it. Therefore, we use this opportunity not only to describe yet another endemic species from this reserve, but to attempt to raise RVS Texiguat���s international profile and issue an urgent call for its conservation. We offer the following recommendations, and emphasize the need for immediate and sustained action to prevent further loss of habitat: Carry out intensive reconnaissance of RVS Texiguat on the Caribbean side of Cerro Texiguat and Cerro San Francisco, to search for and identify previously unknown areas supporting populations of conservation priority species. The majority of work in RVS Texiguat has taken place in the vicinity of La Fortuna, which is accessed by means of a logging road established sometime before 1991. The Caribbean slope of RVS Texiguat remains virtually unexplored and has no known access routes, and so exploratory work is needed immediately to determine if this areas supports target species. Establish a habitat and species monitoring program led by Honduran biologists and local guardarecursos. This program has been initiated in cooperation with PROLANSATE (the organization tasked with managing RVS Texiguat). Initial plans are to hire six guardarecursos in early 2010 and begin training and capacity building for dev, Published as part of Townsend, Josiah H., Butler, Michael, Wilson, Larry David & Austin, James D., 2010, A distinctive new species of moss salamander (Caudata: Plethodontidae: Nototriton) from an imperiled Honduran endemism hotspot, pp. 1-16 in Zootaxa 2434 on pages 4-14, DOI: 10.5281/zenodo.194769, {"references":["Chapman, A. M. (1992). Masters of Animals: Oral Traditions of the Tolupan Indians, Honduras. Gordon and Breach Scientific Publishers, Philadelphia, Pennsylvania. xxi + 259 pp.","Good, D. A. & Wake, D. B. (1993) Systematic studies of the Costa Rican moss salamanders, genus Nototriton, with descriptions of three new species. Herpetological Monographs, 7, 131 - 159.","Papenfuss, T. J. & Wake, D. B. (1987) Two new species of plethodontid salamanders (genus Nototriton) from Mexico. Acta Zoologica Mexicana, 21, 1 - 16.","Savage, J. M. (2002) The Amphibians and Reptiles of Costa Rica: A Herpetofauna Between Two Continents, Between Two Seas. University of Chicago Press. Chicago, Illinois. xx + 934 pp.","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.","Wiens, J. J., Parra-Olea, G., Garcia-Paris, M. & Wake, D. B. (2007) Phylogenetic history underlies elevational biodiversity patterns in tropical salamanders. 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A new species of Nototriton is described from Refugio de Vida Silvestre Texiguat in northern Honduras, and is presented in a new phylogenetic hypothesis for the genus Nototriton. This species differs from all other congeners by a number of morphological characteristics, including but not limited to large nares, syndactylous hands and feet with pointed toe tips, and a pale ventral surface with light mottling. Mitochondrial sequence divergence distinguishes this species from all other Nototriton (3.6–6.0% at 16S and 10.2–15.1% at cytochrome b). Refugio de Vida Silvestre Texiguat harbors a remarkable herpetofauna, including 20% of all endemic amphibian and reptile species in Honduras, among which are five species (one anuran, one lizard, and three snakes) known only from the immediate vicinity of the type locality of the new species. Despite the evolutionary and conservation importance of this reserve, it remains one of the most imperiled cloud forests in Honduras due to persistent illegal logging and forest clearing.

The highlands of northeastern Honduras remain under-characterized in terms of biological diversity, as exemplified by the regularity of new amphibian and reptile taxa discoveries. Following the recent description of a new species of Nototriton from the Sierra de Agalta in northeastern Honduras, we report the discovery of a second new species of Nototriton from the nearby Parque Nacional Montana de Botaderos. This new taxon, Nototriton mime sp. nov., is distinguished from other Nototriton by its distinctive pale brown dorsal coloration in adult males, relatively large nares, a relatively broad head, mitochondrial sequence divergence, and phylogenetic relationships, and is geographically isolated from other populations of Nototriton.

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

Study of allozyme variation, external morphology, and osteology reveals that there are more species of moss salamanders (genus Nototriton) in Costa Rica than the two currently recognized. The three species for which names are available are valid, and new diagnoses are presented for them; three additional species are described. The phylogenetic relationships and biogeography of the six species are investigated. The radiation of Nototriton in present-day Costa Rica has involved miniaturization accompanied by both morphological and ecological specialization. Costa Rican species inhabit moss-mats and leaf-litter; most of the remaining species in the genus are bromeliad-dwellers. The revised genus Nototriton includes two Mexican, one Guatemalan (another, detected in the present study, remains undescribed), two Honduran, and six Costa Rican species. The six Costa Rican species appear to form a monophyletic group, but the phylogenetic relationships of the two northern species groups to each other and to the southern group remain uncertain.

Wake y Elias (1983) encontraron que el género de sala- mandra Chiropterotriton (Plethodontidae) es polifilético. Dos nuevos géneros, Dendrotriton y Nototriton, fueron establecidos para aquellas especies presentes al sur y al oriente del Istmo de Tehuantepec, previamente incluidas en Chiropterotriton. En este trabajo se informa del descubrimiento del género Nototriton en México y de la definición de cuatro grupos de especies. Se describen dos nuevas especies: Nototriton alvarezdeltoroi, habitante del declive caribeño de la Mesa Central de Chiapas, y N. adelos, único miembro del grupo adelos, habitante de la ladera caribeña de la Sierra de Juárez, Oaxaca. El grupo nasa//s también se presenta en Guatemala y Honduras, mientras que los gruposp/cado/y r/chardise conocen sólo de Costa Rica. N. ade/os es de especial interés ya que se encuentra en simpatría con el género norteño Chiropterotriton.

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.