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Bolitoglossa tamaense, sp. nov. Tama salamander, Salamandra del Tama (Figure 3). Holotype. MCNUP 50, an adult male from La Asiria de Bel��n, Parque Nacional Natural Tam�� (PNNT) (7.319278, - 72.374778) at 2,700 m elevation, Departamento de Norte de Santander, Colombia. The specimen was collected by Aldemar A. Acevedo, Karen Silva and Rosmery Franco on August 2010. Paratypes. MCNUP 51���53, adult males with same data as holotype, MCNUP 54���55, adult females with same data as holotype. MCNUP 56���57, adult males and MCNUP 58���61, adult females, collected 2 km away from Los Remansos, Provincia Toledo, (7.330888,- 72.475472), at 2,000 m elevation, Departamento de Norte de Santander, Colombia. All collected by Aldemar A. Acevedo, Karen Silva and Rosmery Franco on October 2010. Diagnosis. A member of the genus Bolitoglossa because of the absence of a sublingual fold and presence of extensive digital webbing and 13 costal grooves between the limbs. A species from the high Andean forests of the eastern flank of the Eastern Colombian Andes (Cordillera Oriental). Distinguished from all other members of South American Bolitoglossa by a combination of coloration, morphological and molecular characters (Fig. 2). Adult individuals exhibit highly distinctive coloration: dorsal and caudal region with orange shades in females and yellow to brown in males. Moderate size (maximum SVL 52.7) with a short snout, short limbs and moderate to extensively webbed hands and feet with longest digits broadly triangular at tip and bluntly pointed (Fig. 4). Males with rounded hedonic mental gland, not prominent. This species is apparently sexually dimorphic in size, mean SVL= 37.2 mm in males and 47 mm in females. Bolitoglossa tamaense can be further distinguished from other South American species of Bolitoglossa (Eladinea) as follows. It is much smaller than B. capitana, which also has more extensively webbed feet and is uniformly black. It is smaller and more extensively webbed than B. adspersa, B. guaramacalensis, B. hypacra, B. tatamae and B. vallecula. It is larger and more extensively webbed than B. orestes and B. spongai. It is larger and has less extensively webbed feet than B. altamazonica, B. biseriata, B. chica, B. digitigrada, B. paraensis, B. peruviana. B. medemi. B. hiemalis and B. walker i. It is a little larger (females) than B. guaneae, has less webbed feet with longest digits more rounded and less pointed at tips, and a tail longer rather than shorter than SVL. It is smaller and has less webbed feet than B. borburata, B. ramosi, B. pandi, B. nicefori and B. lozanoi. Lack of dermal subterminal pads on the ventral surface of the digits differentiates B. tamaense from the highland species B. adspersa, B. hiemalis, B. hypacra, B. savagei and B. vallecula. It has more maxillary teeth (males mean= 35; females mean= 40) and more vomerine teeth (males mean = 18; females mean= 21) than B. hiemalis, B. orestes, B. spongai, B. palmata, B. pandi, B. altamazonica, B. sima, B. peruviana, B. chica, B. lozanoi, and B. leandrae sp. nov. It has fewer maxillary teeth and fewer vomerine teeth that B. vallecula, B. guaramacalensis, B. savage i, B. tatamae, B. ramosi, B. capitana, B. nicefori, B. borburata, B. medemi, B. silverstonei and B. biseriata. While B. asdpersa, B. walkeri and B. equatoriana have fewer maxillary teeth (> 35), they have more vomerine teeth (Description. A medium-sized species, SVL ranges from 39.2 to 52.7 mm (mean= 47 mm) for six females, and from 36.2 to 40.3 mm (mean= 37.2 mm) for six males. Maxillary teeth of moderate size, range from 38 to 39 (mean= 39) in males, and from 39 to 42 (mean= 40) in females. Vomerine teeth range from 17 to 19 (mean= 18) in males, and from 19 to 23 (mean= 21) in females. The trunk is relatively long, ranging from 23.8 to 34.7 mm (mean= 29.3 mm) in females, and from 22.1 to 24.5 mm (mean= 23.5 mm) in males. Distance across the shoulders is 6.0 to 7.5 mm (mean= 6.6 mm) in females, and 4.4 to 5.7 mm (mean= 5.1 mm) in males. Tails are long and generally slender, usually exceeding standard length (from 1.0 to 1.1 mm, mean= 1.1 mm, in males and from 0.9 to 1.1 mm, mean=1.0 mm, in females. The head is moderately broad, from 7.0 to 7.7 mm (mean= 7.6 mm) in females, and 6.9 to 7.2 mm (mean= 7.1 mm) in males. Hind limbs are relatively long (8.4���9.8 mm, mean= 9.1 mm in males; 9.5���11.2 mm, mean= 10.1 mm in females). Moderately webbed feet bearing subterminal pads on digits 2���3 ��� 4. Fingers, in order of decreasing length, are 3���4 ��� 2 ��� 1; toes are 3���4 ��� 5 ��� 2 ��� 1 (Fig. 4). Measurements and morphology of holotype. Male (MCNUP 50) SVL 40.3 mm; head width 7.5 mm; snout to gular fold 10.5 mm; eyelid width 1.6 mm; eyelid length 3.4 mm; anterior rim of orbit to tip of snout 5.6 mm; horizontal orbit diameter 2.4 mm; interorbital distance between angle of eyes 3.1 mm; distance between nuchal groove and gular fold 3.9 mm; snout to forelimb 16.2 mm; distance separating external nares 2.1 mm; distance separating internal nares 2.0 mm; hedonic mental gland 1.9 mm; axilla to groin 25.7 mm; shoulder width 2.1 mm; tail length 43.2 mm; tail width at base 4.6 mm; forelimb length 7.0 mm; hind limb length 10.4 mm; appressed limbs are separated by 3 1 / 2 costal folds; hand width 3.2 mm; foot width 3.8 mm; 32 maxillary teeth; 20 vomerine teeth. Tail and limbs yellow color, dorsum dark brown with yellow spots, gray abdomen, and head darker than the rest of the body (Fig. 3). Coloration in life. We examined 35 adults of Bolitoglossa tamaense, which showed eight patterns of variation in color: A) males with tail and limbs yellow, dorsum dark brown with small yellow spots; B) females with the tail, dorsum and limbs orange, with a dark brown head; C) males and females dark brown with orange feet; D) females orange from the tail to the head, with limbs dark brown. E) Males with the tail, head and feet slightly reddish; F) males with dorsum partially orange with dark spots, with dark brown limbs; G) Males with the tail orange, with yellow feet, head slightly stained orange; H) Males and females with a cream-colored dorsum. All juveniles exhibit dark gray skin coloration. In all individuals, regardless of dorsal color patterns, ventral surface is gray with small brown spots (Fig. 3). Coloration in preservative. All yellow individuals and individuals with gray and brown retain the original color. The orange individual lost all color and turned gray. Variation. The type series consists of 12 specimens (six males and six females), with a SVL range from 36.2- 52.7 mm. Size is sexually dimorphic: mean SVL of six males= 37.2 mm, six females= 47 mm. The largest individual was a gravid female (SVL= 52.7 mm), and the smallest male had a SVL= 36.2 mm. The number of maxillary teeth also varies between the sexes (male mean= 35; females mean= 40), as well as the vomerine teeth (males mean= 18; females mean= 21). Etymology. The specific epithet refers to the Tama National Natural Park; B. tamaense is the first species of salamander recorded from this part of Colombia. Geographic distribution and natural history. Bolitoglossa tamaense is known only from high Andean forests in the eastern flank of the Cordillera Oriental, municipality of Toledo, Departamento de Norte de Santander, Colombia (Fig. 5). Only two localities have been recorded for this species, both in small forest patches, one at 2,000 m elevation and the other population at 2,700 m elevation. Field trips in the area, between 1,800 to 3,300 m elevation, revealed no additional localities. Individuals were active during nighttime, mostly between 18:00 and 19:00 hours, when they are found perched on ferns, bromeliads and low vegetation. During daytime they were found under leaf litter. The population of B. tamaense from Remansos (2,000 m) inhabits a small patch associated with a stream. Whereas several individuals were found in riparian vegetation at Remansos, the salamanders at Asiria inhabit a forest patch without flowing water. After dissecting three females from Remansos we found five pairs of oviductal eggs in each of them. In both localities B. tamaense shares the habitat with at least two species of frogs of the genus Pristimantis., Published as part of Acevedo, Aldemar A., Wake, David B., M��rquez, Roberto, Silva, Karen, Franco, Rosmery & Am��zquita, Adolfo, 2013, Two New Species of Salamanders, Genus Bolitoglossa (Amphibia: Plethodontidae), from the Eastern Colombian Andes, pp. 69-84 in Zootaxa 3609 (1) on pages 75-79, DOI: 10.11646/zootaxa.3609.1.5, http://zenodo.org/record/283297

Testing for Batrachochytrium dendrobatidis (Bd) Samples used to test for the presence of Bd were collected by swabbing at least 25 times on the ventral surfaces and rear foot following Brem et al. (2007). The swabs were stored in sterile Eppendorf vials, which were refrigerated at - 15 ��C. Bd detection was carried out by using the PCR approach described by Annis et al. (2004). DNA was extracted from each swab using Prepman Ultra (Applied Biosystems). All sampled were handled with sterile latex gloves maintaining biosecurity measures to avoid Bd spread between sites and individuals (Aguirre & Lampo 2006)., Published as part of Acevedo, Aldemar A., Wake, David B., M��rquez, Roberto, Silva, Karen, Franco, Rosmery & Am��zquita, Adolfo, 2013, Two New Species of Salamanders, Genus Bolitoglossa (Amphibia: Plethodontidae), from the Eastern Colombian Andes, pp. 69-84 in Zootaxa 3609 (1) on page 72, DOI: 10.11646/zootaxa.3609.1.5, http://zenodo.org/record/283297

Acevedo, Aldemar A., Wake, David B., Márquez, Roberto, Silva, Karen, Franco, Rosmery, Amézquita, Adolfo (2013): Two New Species of Salamanders, Genus Bolitoglossa (Amphibia: Plethodontidae), from the Eastern Colombian Andes. Zootaxa 3609 (1): 69-84, DOI: http://dx.doi.org/10.11646/zootaxa.3609.1.5

Bolitoglossa leandrae, sp. nov. Leandra salamander, Salamandra de Leandra (Figure 6) Holotype. MCNUP 62, male from San Antonio, Parque National Natural Tam�� (PNNT) (7.153092, - 72.227306), 600 m elevation, Departamento de Norte de Santander, Colombia. Collected by Aldemar A. Acevedo, Karen Silva and Rosmery Franco on September 2010. Paratypes. MCNUP 63-64, males, MCNUP 65 female, same data as holotype. Collected by Aldemar A. Acevedo, Karen Silva and Rosmery Franco on September 2010. Diagnosis. A member of the genus Bolitoglossa because of the absence of a sublingual fold and presence of extensive digital webbing and 13 costal grooves between the limbs. Inhabitant of lowland tropical humid forests from the eastern flank of the Eastern Colombian Andes (Cordillera Oriental). Distinguished from all other South American members of Bolitoglossa by a combination of morphological and molecular traits (Fig. 2). A small species (maximum SVL 39.2 mm), snout rounded to truncate in dorsal view, moderate nostrils, eyes are well developed, large, rounded and protruding from the outline of the head, short limbs and extensively webbed digits in hands and feet, without subterminal dermal pad on digits (Fig. 6). This is apparently a sexually dimorphic species in size, with males (mean SVL= 30.3 mm) smaller than females (SVL= 39.2 mm), and males with two-lobed and unpigmented testes, but without an evident hedonic mental gland. Bolitoglossa leandrae can be distinguished from other South American species of Bolitoglossa (Eladinea) as follows. It is smaller than all South American Bolitoglossa (Eladinea) species (Table 2). It has more extensive webbing that highland species B. hypacra, B. adspersa, and B. guaramacalensis. It has moderately extensive webbing, more than in mid-elevation species B. tatamae, B. vallecula, B. spongai, B. orestes, B. palmata, B. savagei, B. ramosi, B. capitana, B. nicefori, B. phalarosoma, B. pandi, B. borburata, B. sima, B. peruviana, B. equatoriana. It is less extensively webbed than lowland species B. biseriata, B. lozanoi, B. altamazonica, B. chica and B. medemi. It is more extensively webbed, smaller, and has fewer maxillary teeth than B. guaneae. It has fewer maxillary teeth (males mean 23; females mean 29) than most South American Bolitoglossa (Eladinea) species (Table 2). It has more maxillary teeth that B. adspera, B. orestes, B. altamazonica, B. chica, and B. equatoriana. It has more vomerine teeth (males mean 19; females mean 20) that B. hiemalis, B. orestes, B. spongai, B. altamazonica, B. sima, B. peruviana and B. lozanoi. Description. A relatively small species, three males SVL range from 29.19 to 31.1 mm (mean= 30.3 mm) one female (39.2 mm). Maxillary teeth of moderate size ranging from 23 to 24 (mean= 23) in males and 29 in the female. Vomerine teeth range from 18 to 19 (mean= 19) in males and 20 in the female. The trunk ranges from 19.6 to 29.31 mm (mean= 21.8 mm) in males; and 29,31 mm in the female. Distance across the shoulders is 4.7 to 6.3 mm (mean= 4.9 mm) in males; 6.3 mm in the female. Tails are long and generally slender, 22.1 to 28.8 mm (mean= 23.8 mm) in males; 28.3 mm in the female. The head is narrow in relation to other species, ranging from 5.4 to 6.7 mm (mean= 5.6 mm in males; 6.7 mm in the female. Hind limbs are relatively short, ranging from 6.4���8.2 mm (mean= 6.8 mm) in males; 8.2 mm in the female. Fingers, in order of decreasing length, are 3 - 2-4 - 1; toes are 3 - 2-4 - 5 - 1. Measurements and morphology of holotype. Male (MCNUP 62) SVL 31.1 mm; head width 5.6 mm; snout to gular fold 7.1 mm; eyelid width 1.0 mm; eyelid length 1.7 mm; anterior rim of orbit to tip of snout 5.3 mm; horizontal orbit diameter 1.1 mm; interorbital distance between eyes 2.8 mm; distance between nuchal groove and gular fold 2.9 mm; snout to forelimb 11.7 mm; distance between external nares 1.9 mm; distance between internal nares 1.4 mm; axilla to groin 19.5 mm; shoulder width 1.8 mm; tail length 25.3 mm; tail width at base 3.8 mm; forelimb length 5.2 mm; hind limb length 5.9 mm; appressed limbs are separated by 2 1 / 2 costal folds; hand width 1.2 mm; foot width 1.9 mm; number of teeth maxillary 23; vomerine 19. Fingers, in order of decreasing length, are 3 - 2-4 - 1; toes are 3 - 2-4 - 5 - 1. Tail and dorsum gray with lines light brown, abdomen brown with light brown spots (Fig. 6). Coloration in life. In the three males of Bolitoglossa leandrae the tail and trunk dorsum are dark brown, with fine yellow stripes along the length of the body, the venter is predominantly grey with small brown spots. The dorsal surface of the female is copper-brown to reddish brown (Fig. 6). Coloration in preservative. All individuals retained the original color. Variation. The type series consists four specimens (three males and one female). The number of maxillary teeth varies between the sexes (male mean= 23; female= 29), as well as the vomerine teeth (males mean= 19; female= 20). Etymology. The specific epithet honours Leandra Mojica, a girl living in the rural area of San Antonio, the type locality of B. leandrae. She represents the children of the rural area in the Tama National Park, who show so much enthusiasm for learning about amphibians. Geographic distribution and natural history. Bolitoglossa leandrae is known only from the eastern flank of the Eastern Colombian Andes (Cordillera Oriental), within the PNN Tama area near San Antonio, Departamento de Norte de Santander, Colombia (Fig. 5). Only one population has been registered, and it was sampled from small forest patches, at 600 m. Field trips between 500 to 1000 m elevation revealed no additional localities. The habitat of B. leandrae consists of secondary tropical forests. B. leandrae, is nocturnal, usually found perched on low vegetation, taking refuge during the day under leaf litter., Published as part of Acevedo, Aldemar A., Wake, David B., M��rquez, Roberto, Silva, Karen, Franco, Rosmery & Am��zquita, Adolfo, 2013, Two New Species of Salamanders, Genus Bolitoglossa (Amphibia: Plethodontidae), from the Eastern Colombian Andes, pp. 69-84 in Zootaxa 3609 (1) on pages 80-81, DOI: 10.11646/zootaxa.3609.1.5, http://zenodo.org/record/283297

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 º N, 83.1683 º 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 Chirripó 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 Chirripó, the highest mountain in Costa Rica. The scientific name is a noun in apposition.

Bolitoglossa aureogularis sp. nov. Yellow-throated Web-footed Salamander Figure 3 Holotype. UCR 19893, an adult female from along the Río Coén on the Trans-Talamancan trail near Cerro Arbolado (9.3925 º N, 83.2119 º W) at an elevation of 1680 m approximately 8 km N of the continental divide, Provincia de Limón, Costa Rica, collected by Eduardo Boza-Oviedo on 1 March 2007. Paratypes. UCR 19892, same data as holotype; UCR 19857 – 59 (3 specimens), 9.3416 º N, 83.232 º W, 2102 m, in headwaters area of Río Coén, about 2 km N continental divide, Provincia de Limón, Costa Rica, collected by Eduardo Boza-Oviedo on 22 February 2007. Diagnosis. Assigned to Bolitoglossa because it lacks a sublingual fold (Wake & Elias 1983), and to subgenus Eladinea based on mtDNA sequence data. A medium-sized member of the genus Bolitoglossa (subgenus Eladinea) with moderate webbing of the digits of the hands and feet that differs from all other species in the genus by its unique coloration (Fig. 3) of reddish tan to yellow dorsal coloration with black flanks and a venter marked by bright yellow gular and yellow-brown chest regions, with a pair of dirty white patches on the ventrolateral surfaces of the posterior venter. In comparison to members of the B. robinsoni complex, this species is much smaller and more slender, in addition to the coloration differences. Description. A slender species of moderate size compared to other members of its genus. SL of holotype, the only adult specimen available (48.8 mm), nearly identical to that of its close geographic neighbor Bolitoglossa splendida and close to mean value of such Talamancan species as B. pesrubra, B. subpalmata (García-París et al. 2008), and B. gomezi (Wake et al. 2007). Tail slender but relatively short (tip broken). Head narrow (Wake & Brame 1972); SL/HW = 8.0. Relatively short snout broadly rounded. Small nostrils are typical for this genus. Nasolabial protuberances not pronounced; paler than surroundings and appear to be pigmented with white. Eyes small, do not protrude beyond lateral margins of head, not visible in ventral view. Teeth moderate in size and numerous (57 MT, 6 PMT, 27 VT). Limbs relatively short with LI of 3.5. Hands and feet moderate in size; FW = 5.1 mm. Digits well-differentiated but short and knob-like with distinct subdigital pads on longer digits. Webbing, reaching between first and second phalangeal articulations of longest digits; webbing more extensive in foot than in hand. Fingers, in order of decreasing length, are 3-4 - 2 - 1; toes are 3-4 - 2-5 - 1. Postiliac glands pale and inconspicuous. Measurements (in mm), limb interval and tooth counts of the female holotype (Table 2). HW 6.1, SG 10.1, HD 2.3, EW 1.3, EL 2.7, ES 1.7, ED 2.2, IC 3.5, IO 2.4, SF 12.5, IN 1.5, SP 0.4, SW 4.8, SL 48.8, SAV 45.0, AX 26.2, LI 6.5, FLL 9.3, HLL 10.2, HAW 3.3, FW 4.3, T 5 0.9, T 3 1.4. Numbers of teeth: PMT 6, MT 29 / 28, VT 13 / 14. Coloration of the holotype in life (Fig. 3). Dorsal coloration golden-tan with some bright highlights on dorsolateral regions and with some narrow streaks of dark brown. Golden-tan coloration in form of broad band extending from snout onto tail. Tail becomes increasingly reddish brown posteriorly. Lateral surfaces dark brown with numerous white speckles. Dark coloration forms lateral margin of dorsal band and continues onto tail and forward all the way to eye and is present almost to tip of snout. Dark coloration extends to area above limb insertions so light band does not contact limbs. Dorsal surfaces of limbs similar to dorsal band in color. White pigment present ventrolaterally along trunk and becomes prominent on venter, where pair of lightly colored patches is separated by region of dark pigmentation. Gular area bright yellow, which becomes golden on chest before fading into darker color in midtrunk region. Venter of tail speckled with white and tan spots. Iris golden. Habitat and range. The species is known from two nearby localities on the Caribbean slope of the Cordillera de Talamanca, both with mature cloud forest similar to the habitat at the type locality of Bolitoglossa splendida. The first locality is at ca. 2100 m elevation, 700 m from the nearest stream; the second, the type locality, is located at 1680 m elevation near the river edge. Variation. There are four paratypes but only one approaches maturity in size. All specimens show coloration typical of the holotype, but some are more reddish gold dorsally and they are covered with tiny white speckles. Even the smallest individual (UCR 19858, 9.2 mm SL) displays the characteristic ventral coloration. This is a slen- der species with a small, narrow head and slender trunk and tail. Etymology. The species name is derived from aurea (L.), golden, and gula (L.), throat, in recognition of the unusual bright yellow coloration of the gular and chest regions of this species. Natural history and behavioral observations. The specimens from the first site were inactive in bromeliads (2.8–6.2 m above ground) during daytime, one in a cavity formed by the bromeliad roots in the trunk of the tree and the other two inside the leaves. All were in different plants in two trees, with two found in separate branches of the same tree; the temperature of the retreats was 12 °C with an air temperature of 15 °C. The juvenile and holotype from the second locality were found active at night on low vegetation or in the leaf litter (0–1.45 m), 1.8–3.7 m from the river with an air temperature of 15–18 °C during the observation period. Behavioral observations were made from 20:00–00:03 before collecting both specimens. The juvenile moved within the Araceae leaf where it was found but never went away. The adult female (holotype) climbed to the top of two shrubs and one palm seedling and moved through the leaf litter between plants. On two occasions, it held up a third of its body off surface of the leaf, and sometimes used its tail as a prehensile organ when moving along steams.

Batrachoseps bramei new species Suggested common name: Fairview Slender Salamander Figures 4–5 B. simatus (part) Brame and Murray 1968: 16 Holotype. Museum of Vertebrate Zoology (MVZ) 217944, an adult female from Packsaddle Canyon, adjacent to Kern River, 1137 m elevation, Tulare Co., California, USA (35.945 °N, 118.476 °W), collected 9 March 1991 by Robert W. Hansen. Paratypes. MVZ 168749–168755, same locality as holotype, collected 10 March 1979 by Robert W. Hansen; MVZ 217945, same data as holotype; MVZ 168756–168763 and MVZ 168950 from Brin Canyon at junction Kern River, Tulare Co., California, USA (35.952 °N, 118.471 °W), collected 11 March 1979 by Robert W. Hansen; MVZ 168737–168738 and MVZ 168740 from Fairview, Tulare Co., California, USA (35.927 °N, 118.494 °W), collected 10 March 1979 by Robert W. Hansen. Referred specimens. MVZ 107168–107169 from 3.6 km S Fairview, E side of Kern River, Tulare Co., California, USA (35.892 °N, 118.464 °W), collected 18 February 1973 by Samuel Sweet and Marjorie Reaka; MVZ 107170 (collected 18 February 1973 by Samuel Sweet), MVZ 168739, MVZ 168741–168748 (collected 10 March 1979 by Robert W. Hansen) all from Fairview, E side of Kern River, Tulare Co., California, USA (35.927 °N, 118.494 °W); MVZ 217946–217954 from Cannell Creek, S side, W of Aqueduct, E of Kern River, Kern and Tulare counties, California, USA (35.794 °N, 118.434 °W), collected 10 March 1991 by Robert W. Hansen; MVZ 224858– 224859 from Upper Kern River Canyon at Mountain Highway 99 bridge over Kern River at confluence with South Falls Creek, E side of Kern River, Tulare Co., California, USA (35.968 °N, 118.486 °W), collected 27 March 1994 by Elizabeth L. Jockusch, David B. Wake and others; MVZ 267129–267137 from trail along W side of Kern River, ca. 0.8 –1.0 km N Mountain Highway 99 bridge over Kern River at confluence with South Falls Creek, Tulare Co., California, USA (35.973 °N, 118.488 °W to 35.977 °N, 118.487 °W), collected 28 January 2010 by Elizabeth L. Jockusch, Elizabeth K. Timpe and Chris Evelyn; MVZ 267140–267142 from Tobias Creek drainage, along trail in vicinity of its crossing of Tobias Creek, Tulare Co., California, USA (35.902 °N, 118.516 °W), collected 22 March 2007 by Elizabeth L. Jockusch and Iñigo Martínez-Solano; MVZ 267154 from Whiskey Flat Trail, ca. 1 km N of Bull Run Creek, W side of Kern River, Tulare Co., California, USA (35.793 °N, 118.452 °W), collected 20 March 2008 by Iñigo Martínez-Solano and Elizabeth L. Jockusch; MVZ 267118–267122 from Plater Rd., just N junction Burlando Rd., Kern Co., California, USA (35.736 °N, 118.428 °W), collected 21 March 2008 by Elizabeth L. Jockusch and Iñigo Martínez-Solano; MVZ 251741–251749 from WNW of Wofford Heights, Tillie Creek drainage, Kern Co., California, USA (35.714 °N, 118.476 °W), collected 27 March 2005 by Brad Alexander. Diagnosis. A small, slender species (standard length, SL, of ten adult males 34.3 mm ± 2.4 mm; of eleven adult females 35.5 mm ± 3.8 mm) distinguished from other species of the B. nigriventris group as follows: from B. simatus by smaller adult size, somewhat more robust appearance (including a longer, broader head, longer limbs and larger feet) and fewer trunk vertebrae (mode 18–19 versus 20–21 in B. simatus); from B. relictus by its longer legs and wider feet; from B. gregarius and B. nigriventris by its substantially more robust appearance including a longer and broader head, longer limbs, and larger hands and feet; from B. stebbinsi Brame and Murray 1968 by its much smaller size and less robust morphology. Distinguished from other species of Batrachoseps in the southern Sierra Nevada as follows: from B. campi Marlow, Brode and Wake 1979 and B. robustus, both members of subgenus Plethopsis, by its unpaired premaxillary bones and smaller, less robust habitus and from B. kawia by its more robust, longer-limbed morphology. Description. Batrachoseps bramei is a small (adults generally less than 45 mm standard length), slender species with a relatively broad, dorsoventrally flattened head and long limbs. The facial region (from the eyes to the snout) is relatively large and almost as broad as the posterior portion of the head; the eyes are moderately protuberant and visible projecting beyond the jaw line in ventral view. The nostrils are small and the nasolabial protuberances are slight to moderate. There is no evidence of a mental hedonic gland under the chins of males. Grooving patterns of the head, throat, and neck are typical of the genus. Standard length ranges from 7.2–8.1 (mean = 7.7) times head width in males and 7.5–9.9 (mean = 8.0) times head width in females. Teeth are relatively numerous: 6– 11 (mean = 7.4) premaxillary teeth in males, 8–18 (mean = 13.7) in females; 31–49 (mean = 40.0) maxillary teeth in males, 32–68 (mean = 47.2) in females; 12–17 (mean = 14.8) vomerine teeth in males, 10–25 (mean = 17.9) in females. The vomerine teeth are arranged patchily to semi-patchily. Small maxillary teeth are borne in a long row extending about to the posterior end of the eyes in females. The premaxillary teeth are the same size as maxillary teeth in females; in males the premaxillary teeth are substantially enlarged. Both males and females have 17–18 costal grooves between the limb insertions. The tail is tapering and relatively short, exceeding body length only in the largest animals. The tail is 1.0– 1.1 (mean = 1.0) times SL in males, and 0.9–1.1 (mean = 1.0) times in females in specimens lacking evidence of tail regeneration. There is no basal tail constriction. The postiliac gland is visible as an obscure pale spot. The limbs are relatively long; limb interval ranges from 4.5–6 (mean = 5.2) in males and 5–8 (mean = 5.9) in females. SL ranges from 4.5 –6.0 (mean = 5.1) times hind limb length in males and 4.6–5.7 (mean = 5.0) times in females. The hands and feet are relatively large; foot width ranges from 1.7–2.2 mm (mean 2.0 mm) in both males and females. The digits are well formed and discrete with expanded tips that bear subterminal pads. Webbing occurs between the fingers and toes of some individuals, and extends to approximately the second phalanx. Fingers and toes in order of decreasing length are 3 -2,4- 1. Digit 1 is very reduced. Measurements of the holotype (in millimeters). Maximum head width 4.1; snout to gular fold (head length) 8.3; head depth at posterior angle of jaw 1.9; eyelid length 2.5; eyelid width 1.2; anterior rim of orbit to snout 1.3; horizontal orbital diameter 1.7; interorbital distance 1.4; snout to forelimb 10.7; distance separating external nares 1.7; snout projection beyond mandible 0.5; snout to posterior angle of vent (SL) 40.7; snout to anterior angle of vent 37.3; axilla to groin length 23.4; tail length 33.5; tail width at base 1.9; tail depth at base 1.8; forelimb length 6.7; hind limb length 7.2; limb interval 8; width of right hand 1.7; width of right foot 2.2; foot length 2.9; length of third toe 0.8; body width behind forelimbs 2.2. There are 15 premaxillary, 46 maxillary, and 20 vomerine teeth; the row of maxillary teeth on the right appears short, ending at the posterior border of the internal naris. There are 18 costal grooves between the limb insertions. Coloration of the holotype (in life). Uniformly flat black ventrally with delicate sprinkling of punctate guanophores; ventral guanophores are tiny and rarely connect; most numerous on gular region and at lowest density on tail; guanophores abundant on lateral surfaces, where they become expanded and form irregular patches. On lateral surfaces, guanophores cover more area than ground color. Dorsum metallic silver as a result of numerous expanded guanophores that are especially concentrated dorsolaterally; silvery coloration has metallic brassy-gold highlights; from a distance the silver on black background appears bluish-gray. Guanophores on head more disconnected and less concentrated, but the whole head, including eyelids and snout, is heavily spotted. Pigment most concentrated on nape of neck and front of limbs. Limbs heavily pigmented, especially proximally, but even fingers and toes have extensive spotting. Dark gray iris also has some dorsal guanophores. Only face region has light spotting. Ground color of dorsum slate black to brownish black. Under high magnification, it is clear that the lateral guanophores are more white and superficial, while the dorsal ones are deeper and more metallic. They co-occur on the dorsum, with the superficial cells rare. On the tail, the metallic color becomes gold or brassy and very concentrated. Morphological variation. Males sampled from the northern end of the range have relatively small adult body size compared to populations from the southern end of the range (mean of 8 males from southern Wofford Heights 42.1 ± 3.6 mm vs. 33.8 ± 3.7 mm for 28 males from northern Packsaddle/Brin/Fairview). No significant differences in female body size across populations were detected; however, sample sizes were Coloration is highly variable both within and between populations (Fig. 5). Many specimens have distinct patches of color over the shoulders; these are usually coppery to gold in color, and do not fully fuse into the dorsolateral stripes. More northern individuals generally exhibit fairly subdued dorsal coloration, but individuals from Cannell Creek can be boldly marked and have reddish as well as metallic coloration. Habitat and distribution. Batrachoseps bramei is known only from the Upper Kern River Canyon, and along the west side of the current Lake Isabella; the southern limit of its distribution is near the original junction of the main and South forks of the Kern River (Fig. 1). The southernmost samples are from Wofford Heights on the west side of the river and ca. 2 km S of the Cannell Creek drainage on the east side of the river. The range extends north at least as far as 1 km N of the confluence of South Falls Creek with the Kern River. Thus, known populations range for about 30 km from south to north. Areas farther north, which are accessible only by trail, have not been thoroughly surveyed; it is likely that the range extends farther north in the Upper Kern River Canyon. B. bramei occurs within an elevational range of 860–1280 m, one of the most restricted among all species of Batrachoseps. B. bramei has not been found in sympatry with any other species of Batrachoseps. It closely approaches B. simatus in the Lower Kern River Canyon (13 km between Wofford Heights B. bramei and Erskine Creek B. simatus; 20 km between Cannell Trail B. bramei and Erskine Creek B. simatus); however, these two species are separated by a major drainage (either the main or South Fork Kern River) as well as by xeric, inhospitable terrain around the confluence of the South Fork and Upper Kern River at Lake Isabella. B. bramei also closely approaches B. robustus, which has been collected within 7 km of the river, but at much higher elevation (2214 m), and B. altasierrae sp. nov. (see below). The range of B. bramei on the east side of the Kern River lies entirely within a prominent uplifted ridge of metamorphic rocks paralleling the river. Local populations of salamanders are associated with north-facing slopes and talus. A chaparral plant community consisting of species of the genera Ceanothus, Arctostaphylos, Ribes and Chrysothamnus, as well as Pinus sabiniana and occasionally Quercus chrysolepis, characterizes this area. Most individuals have been found beneath rocks, often on or at the base of talus slopes (Fig. 6). However, specimens have been found in a variety of other habitats and under a variety of cover objects, including under a log in an open sandy flood plain, under logs and rocks in grasslands, in gravel on the river bank, and in leaf litter in protected groves (Fig. 6). Individuals have been observed active at night during winter and early spring and have been collected from beneath surface cover objects under snow. Salamanders have been found under surface cover at body temperatures (inferred from substrate temperatures) of 2.2–16.4 °C (mean = 8.4 °C, N = 101). Etymology. Named in honor of Arden H. Brame, Jr., who, along with Keith Murray, was the first to recognize the distinctiveness of Batrachoseps in the Kern River Canyon. The species name is formed as a noun in the genitive. Comments. In their classic work on Batrachoseps, Brame and Murray (1968) referred animals from the Upper Kern River Canyon to B. simatus, but noted that they were morphologically different. Stebbins (1985, 2003) followed this taxonomy. Jockusch and Wake (2002) treated populations from the Upper Kern River Canyon, all included here in B. bramei, as undescribed lineages based on a phylogeny inferred from mitochondrial DNA sequence data, but the high genetic diversity within the region led them to raise questions about whether more than one species might be present. Jennings (2004) treated populations from the Upper Kern River Canyon as an undescribed species and provided a vernacular name “Fairview slender salamander.” Bartlett and Bartlett (2009) refer to the species as awaiting formal description but provide a short species account, a distribution map, and a color photograph and refer to it as the “Fairview Slender Salamander.” Until 2004, it was believed that Batrachoseps from the canyon proper occurred only on the eastern and southern sides of the river. After specimens were found in the Tillie Creek drainage on the northwest side of the river, exploration intensified on that side, and specimens have now been found at almost all sites north of Tillie Creek that we have sampled with moderate effort (> 4 person-hours of searching under good conditions). Conservation. The range of B. bramei lies entirely within plant communities that naturally experience periodic fire. The use of heavy equipment for fire suppression has the potential to negatively impact salamander habitat. Road maintenance of Mountain Highway 99 (Kernville–Johnsondale) should take into account the intimate proximity of some populations to road-edge habitat. Most known populations occur on public lands administered by Sequoia National Forest. Although B. simatus has become difficult to find and is listed as a Threatened Species by the State of California, B. bramei appears to be more abundant and can be found consistently throughout its range.

Batrachoseps altasierrae new species Suggested common name: Greenhorn Mountains Slender Salamander Figure 7 A���C Batrachoseps attenuatus (part) Grinnell and Camp 1917: 137 Batrachoseps attenuatus attenuatus (part) Dunn 1926: 232 Batrachoseps relictus (part) Brame and Murray 1968: 5 Batrachoseps pacificus relictus (part) Yanev 1980: 535 Batrachoseps relictus (part) Jockusch, Wake and Yanev 1998: 13 Holotype. Museum of Vertebrate Zoology (MVZ) 59909, an adult male from 1.5 mi [2.4 km] SE Alta Sierra [Greenhorn Mountains], Kern Co., California, USA (35.732 ��N, 118.538 ��W), collected on 26 June 1952 by Keith Murray. Paratypes. MVZ 190953 and MVZ 190955, both from Hwy. 155, 0.85 km N (by road) Greenhorn Summit, Kern Co., California, USA (35.744 ��N, 118.557 ��W), collected 28 May 1984 by Robert W. Hansen; MVZ 190961 ��� 190963 and MVZ 190965 ��� 190967, all from Hwy. 155, 1 km ESE (by road) Greenhorn Summit, Kern Co., California, USA (35.735 ��N, 118.554 ��W), collected 24 June 1984 by Robert W. Hansen and D. C. Holland; MVZ 190976 ��� 190979 from Tiger Flat Rd. (= U.S. Forest Service Rd. 23 S 16), 0.7 km N (by road) junction Hwy. 155 at Greenhorn Summit, Kern Co., California, USA (35.744 ��N, 118.554 ��W), collected 24 June 1984 by Robert W. Hansen and D. C. Holland; MVZ 190983 ��� 190985 from Tiger Flat Rd. (= U.S. Forest Service Rd. 23 S 16), 0.85 km N (by road) junction Hwy. 155 at Greenhorn Summit, Kern Co., California, USA (35.745 ��N, 118.553 ��W), collected 24 June 1984 by Robert W. Hansen and D. C. Holland. Referred specimens. Paratypes of Batrachoseps relictus: Los Angeles County Museum (LACM) 33082��� 33090 and MVZ 184906���184908 from 1.2 mi [1.9 km] S White River Camp Grounds, Tulare Co., California, USA (35.840 N, 118.649 ��W), collected 31 March 1957 by A. H. Brame, Jr. (MVZ 184906 ��� 8 are described as ��� 3 cleared & stained specimens in D. B. Wake collection��� by Brame and Murray (1968)); LACM 33091���33093 from 1 mi [1.6 km] above O���Quinn Meadow, Tulare Co., California, USA (35.863 ��N, 118.628 ��W), collected 31 March 1957 by L. Hughes and A. H. Brame, Jr.; LACM 33080���33081 from White River Camp Grounds (Lower Camp), Tulare Co., California, USA (35.845 ��N, 118.636 ��W), collected 31 March 1957 by A. H. Brame, Jr. Additional referred specimens: MVZ 156360���156365 from Quaking Aspen Meadows along Hwy. 190, South Fork of Middle Fork Tule River, Tulare Co., California, USA (36.118 ��N, 118.544 ��W); MVZ 156366���156392 from the upper reaches of Spear Creek along U.S. Forest Service Rd. 24 S06, 6.9 km SE junction of 23 S05 and 3.2 km NE Portuguese Pass, Tulare Co., California, USA (35.821 ��N, 118.583 ��W); MVZ 156393���156399 from White River drainage, at Sugarloaf Peak, U.S. Forest Service Rd. 23 S05, 0.6 km N junction U.S. Forest Service Rd. 24 S06, Tulare Co., California, USA (35.843 ��N, 118.614 ��W), all collected 29 July 1975 by Kay P. Yanev and Samuel S. Sweet; MVZ 224835 from road to Sugarloaf Village, 1.0 km SE Sugarloaf Village, Tulare Co., California, USA (35.820 ��N, 118.641 ��W), collected 26 March 1994 by Elizabeth L. Jockusch, David B. Wake and others; MVZ 158244 and MVZ 158226��� 158228 from 9.5 km E Cherry Hill Road on Sherman Pass Road, Tulare Co., California, USA (35.983 ��N, 118.381 ��W), collected 11 July 1980 and 28 July 1979, respectively, by Robert W. Hansen; MVZ 224810 from Hwy. 155, 9.0 km NW Lake Isabella and ca. 2.2 km SE Alta Sierra, Tulare Co., California, USA (35.724 ��N, 118.527 ��W), collected 27 March 1994 by Elizabeth L. Jockusch, David B. Wake and others. Diagnosis. A small, relatively slender species (SL of 8 adult males 38.0 mm �� 2.4 mm; of 8 adult females 40.1 mm �� 3.5 mm) distinguished from other species of Batrachoseps in the southern Sierra Nevada and vicinity as follows: from B. relictus by a relatively longer trunk, a relatively narrower head, shorter limbs, and smaller feet, and by females having many fewer maxillary teeth; from B. simatus by its smaller adult body size with a relatively narrower head and chest and fewer trunk vertebrae (mode 18���19 versus 20���21 in B. simatus); from B. bramei by a relatively shorter and narrower head, longer trunk, shorter limbs, narrower feet and longer tail, and smaller number of maxillary and vomerine teeth; from B. gregarius by its more robust morphology including a relatively longer and broader head, shorter trunk, longer limbs, and larger hands and feet; from B. stebbinsi by its less robust morphology and much smaller size; from B. campi and B. robustus by its less robust morphology, much smaller size, different color pattern and unpaired premaxillary bone. Distinguished from its close relative B. kawia by having fewer maxillary teeth; from B. regius Jockusch, Wake and Yanev 1998 by having a relatively longer trunk and tail, shorter head, shorter limbs, smaller feet and fewer maxillary teeth; and from B. diabolicus Jockusch, Wake and Yanev 1998 by having fewer trunk vertebrae, a relatively narrower head, shorter limbs and smaller feet. Description. Batrachoseps altasierrae is a small (adults less than 50 mm standard length), slender species with a relatively narrow head and short limbs. The facial region is relatively narrow, and the eyes are not generally protuberant enough to be seen in ventral view. Mental hedonic glands are not visible under the chins of males. Grooving patterns of the head, throat, and neck are typical of the genus. Standard length ranges from 8.4���9.5 (mean = 9.0) times head width in males and 8.2���9.9 (mean = 9.3) times head width in females. There are relatively few teeth, especially on the maxilla: 4���5 (mean = 4.8) premaxillary teeth in males, 4���12 (mean = 6.4) in females; 9���27 (mean = 17.4) maxillary teeth in males, 18���28 (mean = 21.9) in females; 6���13 (mean = 10.9) vomerine teeth in males, 9���16 (mean = 11.8) in females. Vomerine teeth are arranged somewhat patchily. Small maxillary teeth are borne in a long row extending about two thirds of the way through the eye in males and to the posterior end of the eye in females. In females, premaxillary teeth are the same size as maxillary teeth; in males, the premaxillary teeth are enlarged. Males and females both have 18���19 trunk vertebrae and 17���18 costal grooves between the limb insertions. The tail is long and fairly cylindrical, tapering at the tip. The tail is 1.3���1.5 (mean = 1.4) times SL in males and 1.1���1.6 (mean = 1.3) times in females in specimens lacking evidence of tail regeneration. There is no basal tail constriction. The postiliac gland is present. The limbs are relatively short in length, and limb interval ranges from 7���9.5 (mean = 8.3) in males and 7.5���9.5 (mean = 8.5) in females. SL ranges from 6.2���7.5 (mean = 6.8) times hind limb length in males and 6.3���7.8 (mean = 7.1) times in females. The hands and feet are relatively narrow; foot width ranges from 1.4���1.7 mm (mean 1.6 mm) in males and 1.4 ���2.0 mm (mean 1.7 mm) in females. The digits are short, well formed and discrete with expanded tips that bear subterminal pads. Webbing is insignificant. Fingers and toes in order of decreasing length are 3 - 2-4 - 1. Measurements of the holotype (in millimeters). Maximum head width 4.9; snout to gular fold (head length) 7.4; head depth at posterior angle of jaw 2.7; eyelid length 2.2; eyelid width 1.2; anterior rim of orbit to snout 1.4; horizontal orbital diameter 1.8; interorbital distance 1.9; snout to forelimb 9.3; distance separating external nares 1.6; snout projection beyond mandible 0.5; snout to posterior angle of vent (SL) 42.5; snout to anterior angle of vent 36.9; axilla to groin length 25.6; tail tip broken after 33.5; tail width at base 3.3; tail depth at base 2.7; forelimb length 6.3; hind limb length 6.8; limb interval 9; width of right hand 1.2; width of right foot 1.6; foot length 2.1; length of third toe 0.9; body width behind forelimbs 2.9. There are 4 premaxillary, 27 maxillary, and 13 vomerine teeth; vomerine teeth are arranged somewhat patchily. There are 17 costal grooves between the limb insertions. Coloration of the holotype (in alcohol). The ground color is dark blackish brown dorsally and laterally (and in the limbs), and fades to lighter brown ventrally. A prominent dorsal stripe ranging in color from dark brown to reddish brown is present and is well demarcated on its lateral borders. Habitat and distribution. This species is restricted to higher elevations in the southern Sierra Nevada. The elevational range is from 900���2440 m. Populations extend from the higher elevations on the northern side of the Lower Kern River Canyon to the Tule River drainage and upper elevations of the Little Kern River drainage in Kern and Tulare counties, California, USA (Fig. 1). Two populations, separated by just over 1 km, are known from the western margin of the Kern Plateau, Tulare Co., California, USA. Most populations are found in coniferous forest containing a mixture of pine, fir and incense cedar (Fig. 7 F). B. altasierrae has not been found in sympatry with any other species of Batrachoseps. It closely approaches B. gregarius at lower elevations on the western edge of its range. B. gregarius is found within 5 km of B. altasierrae along the Middle and North Forks of the Tule River, where B. altasierrae occurs as low as 900 m. In the Greenhorn Mountains, B. gregarius is known from as high as 1460 m (MVZ 266680���266683; 35.745 ��N, 118.598 ��W) and B. altasierrae from as low as 1460 m (California Academy of Sciences (CAS) 224163, identified by morphology; 36.176 ��N, 118.692 ��W) in the vicinity of Cedar Creek; these populations are separated by less than 1.5 km. On the eastern edge of its range, B. altasierrae closely approaches B. bramei, but the two species appear to be separated by elevation, although the highest-elevation samples of B. bramei are from less than 200 m below the lowest-elevation samples of B. altasierrae in the Greenhorn Mountains. The ranges of these species are separated by less than 10 km between Alta Sierra and Wofford Heights. They may also approach each other along the Upper Kern River Canyon. B. altasierrae on Sugarloaf Peak, Tulare Co., California occurs less than 10 km away from B. bramei along Tobias Creek, Tulare Co., California. B. altasierrae is known from Peppermint Creek, Tulare Co., California; this locality is about 2 km from the Kern River. The nearest known B. bramei is about 10 km south; however, the range of B. bramei likely extends north from there along the Kern River. On the Kern Plateau, east of the Kern River, B. altasierrae has been found 1.6 km from the nearest population of B. robustus in habitat typical of both species (Wake et al. 2002) and less than 10 km from the nearest B. bramei. At higher elevations to the north of the Tule River drainage, B. altasierrae is replaced by its sister species, B. kawia. To date, all robust specimens (that is, not B. gregarius) from the South Fork of the Kaweah River identified with molecular data belong to B. kawia, whereas robust specimens from the Tule River drainage carry mitochondrial DNA from B. altasierrae. The northernmost specimen of B. altasierrae is from Mountain Home State Forest, Tulare Co., California, USA (CAS 214814; 36.269 ��N, 118.668 ��W), which is 12 km southeast of the southernmost B. kawia from entrance to Soldiers Cave near South Fork entrance Sequoia National Park, Tulare Co., California, USA (MVZ 237285; 36.345 ��N, 118.7568 ��W). Very few specimens are available from the higher elevations separating these drainages. Molecular sequence data show that some populations along the North Fork of the Middle Fork of the Tule River have genes from both B. altasierrae and B. kawia, but the contact zone appears to be narrow (Jockusch et al. unpublished). Etymology. Named for the mountain hamlet of Alta Sierra, located at the summit of the Greenhorn Mountains, an area where this species is particularly common. The species name is formed as a noun in the genitive. Comments. Jockusch et al. (1998) included a rediagnosis of B. relictus based on specimens from both the Lower Kern River Canyon and the Greenhorn Mountains. We use their series from the Greenhorn Mountains as the paratypes for B. altasierrae. All molecular data identified in previous publications as coming from B. pacificus relictus (Yanev 1978, 1980) or B. relictus (Jockusch 1996; Jockusch et al. 1998; Jockusch & Wake 2002) are from B. altasierrae or from more northern members of its species group. Thus, the molecular diagnoses provided by Jockusch et al. (1998) to distinguish B. relictus from other species all in fact apply to B. altasierrae rather than to B. relictus. Molecular differentiation within B. altasierrae is limited. Yanev (1978) included six populations of B. altasierrae, ranging from the vicinity of the type locality in the Greenhorn Mountains to Quaking Aspen Meadow, on the South Fork of the Middle Fork of the Tule River near the northern end of the range, in her allozyme study of the genus. These populations were separated by a maximum Nei���s (1972) genetic distance (D) of 0.075. Mitochondrial DNA (cob) data also show a low level of variation (Jockusch & Wake 2002). Conservation. By virtue of their previous inclusion in B. relictus, populations of B. altasierrae have been listed as a Species of Special Concern by the State of California and Sensitive Species by the U.S. Forest Service. Our impression is that populations of B. altasierrae are healthy, both in the Greenhorn Mountains and within the Tule River drainage; individuals have been found in large numbers when surface conditions were appropriate. For example, about 60 individuals were seen along a single stream in the vicinity of Sugarloaf Village, Greenhorn Mountains, Tulare Co., California in a few hours of searching in August 1995. B. altasierrae was also abundant in the Tule River drainage, Tulare Co., California in April 2008: individuals were found at numerous sites, and at Moorehouse Creek, along Hwy. 190 (36.153 ��N, 118.657 ��W), 10 individuals were found in an area of a few square meters of moist pine needle litter in less than 10 min., Published as part of Jockusch, Elizabeth L., Mart��nez-Solano, I��igo, Hansen, Robert W. & Wake, David B., 2012, Morphological and molecular diversification of slender salamanders (Caudata: Plethodontidae: Batrachoseps) in the southern Sierra Nevada of California with descriptions of two new species, pp. 1-30 in Zootaxa 3190 on pages 13-16, DOI: 10.5281/zenodo.215268, {"references":["Grinnell, J. & Camp, C. L. (1917) A distributional list of the amphibians and reptiles of California. University of California Publications in Zoology, 17, 127 - 208.","Dunn, E. R. (1926) The Salamanders of the Family Plethodontidae. Smith College, Northampton, Massachusetts, 456 pp.","Brame, A. H. Jr. & Murray, K. F. (1968) Three new slender salamanders (Batrachoseps) with a discussion of relationships and speciation within the genus. Bulletin of the Natural History Museum of Los Angeles County, 4, 1 - 35.","Yanev, K. P. (1980) Biogeography and distribution of three parapatric salamander species in coastal and borderland California. In: Power, D. M. (Ed.), The California Islands: Proceedings of a Multidisciplinary Symposium. Santa Barbara Museum of Natural History, Santa Barbara, California, pp. 531 - 550.","Jockusch, E. L., Wake, D. B. & Yanev, K. P. (1998) New species of Batrachoseps (Caudata: Plethodontidae) from the Sierra Nevada, California. Los Angeles County Museum Contributions in Science, 472, 1 - 17.","Yanev, K. P. (1978) Evolutionary Studies of the Plethodontid Salamander Genus Batrachoseps. Ph. D. Dissertation, University of California, Berkeley, 242 pp.","Jockusch, E. L. (1996) Evolutionary Studies in Batrachoseps and other Plethodontid Salamanders: Correlated Character Evolution, Molecular Phylogenetics, and Evolution of Reaction Norms. Ph. D. Dissertation, University of California, Berkeley, 220 pp.","Jockusch, E. L. & Wake, D. B. (2002) Falling apart and merging: the diversification of slender salamanders (Plethodontidae: Batrachoseps) in the American West. Biological Journal of the Linnean Society, 76, 361 - 391."]}

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

Bolitoglossa kamuk sp. nov. Kamuk Web-footed Salamander Figure 4 Holotype. UCR 20852, a young adult male from the sub-páramo region atop Cerro Apri in the Kamuk Massif (coordinates 9.2546 º N, 83.059 º W) at an elevation of 3126 m, on the continental divide, Provincia de Limón, Costa Rica, collected 18 December, 2007, by Guillermo Artavia, Gerardo Chaves, Sean Rovito, Guido Saborío and Hugo Solano. Paratypes. UCR 20853, 20854; same data as holotype. Diagnosis. Assigned to Bolitoglossa because it lacks a sublingual fold (Wake & Elias 1983), and to subgenus Eladinea based on mtDNA sequence data. A slender member of the subpalmata group of Bolitoglossa (Eladinea) distinguished from all other members of the group by mitochondrial DNA sequences; further distinguished from B. pesrubra by absence of reddish proximal limb segments and absence of dorsal spots and blotches; from B. gracilis by more robust habitus and absence of yellowish coloration; from B. tica by smaller hands and feet. Description. Known only from juveniles and small adults. Appears to be relatively small, slender compared to other members of genus. SL in two small adult males is 34.8 mm and 33.0 mm. Tails slender, about same length as SL; SL/TL in two males is 0.99 and 0.98. Head narrow; SL/HW is 6.0 and 6.2. Head well demarcated from neck. Snout broadly rounded to truncated, not prominent. Nostrils small and nasolabial protuberances poorly developed but lightly pigmented. Eyes moderately prominent, protrude slightly beyond lateral margins of head and are relatively frontal in orientation. Holotype has 2 PMT, but large paratype (UCR 20853) has a damaged snout and teeth cannot be counted. MT 34 in holotype, and 15 on one side of large paratype. VT number 16 (holotype) and 13. Limbs slender, moderate in length; LI 1.5 (holotype) and 2. Hands and feet narrow (3.7 and 3.6 mm in holotype) with little webbing; fewer than two distal-most phalanges of longest digits are free of webbing. Digital tips truncate to slightly pointed and bear small but distinct subterminal pads. Fingers, in order of decreasing length, are 3-4 - 2 - 1; toes are 3-4 - 2-5 - 1. Postiliac glands not evident. Gonads of the holotype compact and rounded with some spotting of black pigmentation, and they appear to be sexually mature. Measurements (in mm), limb interval and tooth counts of the male holotype (Table 2). HW 5.8, SG 8.5, HD 3.1, EW 1.0, EL 2.5, ES 1.7, ED 1.6, IC 2.6, IO 2.6, length of groove extending posteriorly from eye 2.3, distance between nuchal groove and gular fold 2.6, SF 10.8, IN 1.1, SP 0.4, SL 34.8, SAV 32.0, AX 19.2, LI 1.5, TL 35.2, tail width at base 2.4, tail depth at base 3.0, FLL 9.0, HLL 9.3, HAW 2.5, FW 3.7, T 5 0.7, T 3 1.1, mental gland width 1.3, mental gland length 1.2. Numbers of teeth: PMT 2, MT 17 / 17, VT 8 / 8, arranged in a single row. Coloration of the holotype in life (Fig. 4). Dorsum and dorsal surface of tail and hind limbs black with numerous, uniformly distributed gold flecks. This coloration extends to lateral midline, where gold flecks become much less numerous. Background of fore limbs somewhat lighter. Gold flecks less numerous on top of head. Venter is dark grey, with a few gold specks, particularly towards sides of body. Gular region, underside of limbs, and anterior portion of tail pale grey with a few golden specks, while posterior half of the tail darker grey. Coloration of the holotype in alcohol. Region of head between snout and eyes brown but darkened due to presence of skin glands that are heavily pigmented peripherally; frontal and parietal areas brown with skin glands with lighter edges; orbits completely black, canthus rostralis strongly marked numerous black spots on dark brown background; ventral area cream with numerous black spots that are even more concentrated in anterior region, making gular region lighter. Dorsum of trunk dark brown that darkens posteriorly; flanks marked with numerous black spots on background passing from brown to clear cream from dorsum to venter; venter cream with numerous dots but still noticeably lighter than dorsum. Tail dark; dorsal part of first three post-sacral segments vertebrae and underside of first 11 caudal segments maintains color of trunk, while progressively darkening posteriorly, and becoming completely black. Both hind limbs and forelimbs appear black due to presence of numerous black spots on dark brown background; dorsal part of hands and ventral part of hands and feet lighter than rest of limb. Variation. The juvenile paratype (UCR 20854) is orange in dorsal coloration with both dark grey and lighter colored specks. A darker grey patch is present on the top of the head posterior to the eyes and on the orbits. The dorsal surface of the limbs and tail are bright orange, changing to a darker orange-grey at the tip of the tail. The gular region is a pale yellow-orange color, while the venter is a pale golden color with numerous black specks. The underside of the tail is a uniform orange. Habitat and range. This species is known only from the type locality on Cerro Apri, southwest of Cerro Kamuk just off the continental divide. Habitat consists of sub-páramo vegetation with extensive, deep moss mats, spongy soil, ferns, and small, isolated trees with arboreal bromeliads. The species was found both within these bromeliads and under moss. Etymology. The species is from the Kamuk Massif, named for one of the dominant peaks in the region, Cerro Kamuk. The scientific name is a noun in apposition.

Blackburn, David C., Wake, David B. (2012): Additions and corrections. Zootaxa 3381 (1): 45-46, DOI: 10.11646/zootaxa.3381.1.2, URL: https://biotaxa.org/Zootaxa/article/view/zootaxa.3381.1.2

Pseudoeurycea cafetalera sp. nov. Coffee Grove Salamander, Salamandra de cafetal Figure 2 A, 2 C– 2 H. Pseudoeurycea cephalica: Parra-Olea (2002). Table 1 (voucher: IBH 14343) Holotype. Colección Nacional de Anfibios y Reptiles IBH 14341, an adult female from Rancho dos Puentes, 6.0 km (by road) north of entrance to Huatusco, Veracruz, México (19 ° 11.13 ' N, 96 ° 57.72 ' W, 1322 m elevation) (Fig. 1), collected by Sean M. Rovito on November 17, 2009. Paratypes. Seven specimens. Two males: IBH 14344, same data as holotype; IBH 14349, from Las Cañadas, 5.4 km NW of center of Huatusco, Veracruz, México (19 ° 11.42 ' N, 96 ° 59.30 ' W, 1367 m elevation), collected by J. C. Winfield-Pérez. Five females: IBH 14338 -14340, Zoncolco, Veracruz, México (18 º 46.4 ' N, 97 º 07.61' W, 1600 m elevation); IBH 14343, same locality as holotype, collected by GPO, DBW, M. García-París and J. Hanken; IBH 14346, 3.5 km SW La Raya, road to Trincheras, Veracruz, México. Referred specimens. Three juveniles: IBH 14337, same data as holotype; IBH 14345, Zoncolco, Veracruz, México (18 º 46.4 ' N, 97 º 07.61' W, 1600 m elevation); MVZ 196052, La Joya, Veracruz, México (19 ° 36.65´N, 97 ° 1.63´W, 2125 m elevation). Diagnosis. A medium sized species of Pseudoeurycea; distinguished from all other members of the genus Pseudoeurycea (except P. cephalica group) by the combination of: stout body, long stout legs, shorter digits, more webbed hands and feet with first and fourth fingers barely emerging from membrane. Distinguished from members of P. bellii group by lack of reddish spots on dorsum. Distinguished from its two sister taxa P. quetzalanensis and P. cephalica in the following characters: from P. quetzalanesis by its larger size (mean SL = 45 mm for males and 50 mm for females of P. cafetalera vs. 36 mm for males and 38 mm for females of P. quetzalanensis), longer limbs (adpressed limbs separated by - 1–1.5 costal interspaces in females and they meet or are separated by 0.5 costal interspace in males of P. cafetalera; for P. quetzalanensis, separated by 1– 2.5 for females and 0.5–1 for males), and wider feet (mean FW = 4.8 mm for females and 4.6 mm for males of P. cafetalera vs. 3.0 mm and 3.2 mm for females and males, respectively, of P. quetzalanensis). It differs from the other closely related species P. cephalica by its broader head (mean HW = 8.5 mm and 7.7 mm for females and males, respectively, of P. cafetalera and 7.1 mm and 6.9 mm for females and males, respectively, of P. cephalica), and more maxillary and vomerine teeth in females (mean MT = 62 in P. cafetalera vs. 56 in P. cephalica; mean VT = 32 in P. cafetalera vs. 27 in P. cephalica). It differs from other members of the P. cephalica group (P. galeanae and P. scandens) in being smaller; P. galeanea can reach 70 mm SL and P. scandens can reach 71 mm SL (Taylor, 1941; Walker, 1955). It also differs from these species in coloration; P. galeanae has large whitish spots on the tail, especially near the base, which are not present in P. cafetalera, and P. scandens has a uniform dark belly while P. cafetalera can have light grey flecks on the belly. Finally, P. scandens has longer legs with longer truncate digits (Walker 1955), and P. galeanae has more maxillary teeth (Taylor, 1941). Description. A medium sized species; SL in 2 adult males 42.8–48.1 mm (mean = 45.5 mm), in 6 females 43.1–59.8 mm (mean = 49.9 mm), with robust habitus; head relatively broad, 0.16–0.18 SL in both males and females, snout broadly rounded in females and squared in males; neck region ill-defined, only slightly narrower than head. Parotoid glands not evident. Costal grooves 13, counting one each in axilla and groin. Limbs relatively long; digits relatively short and stout for Pseudoeurycea, bearing small subterminal pads; fifth toe much shorter than fourth. Digits in order of decreasing length: fingers 3 - 2-4 - 1; toes 3-4 - 2-5 - 1; basal webbing present. Tail missing in many specimens; when present, 0.69–0.76 (mean = 0.79) SL in females and 0.75 in males; tail stout, tapering rather abruptly toward tip. Maxillary teeth small, 45–53 (mean 49) in males, 58–73 (mean 62.5) in females; premaxillary teeth 3–4 (mean 3.5) and enlarged in adult males, 6–12 (mean 9.3) and smaller in females; vomerine teeth in long rows, 24 in males, 25–43 (mean 31.8) in females. Measurements (in mm), limb interval and tooth counts of the holotype. HW 9.7; HD 4.8; eyelid length 4.1; eyelid width 2.2; anterior rim of orbit to snout 3.2; interorbital distance 3.2; distance between outer corners of eyes 8.3; snout to forelimb 16.8; ND 0.3; distance between external nares 2.6; projection of snout beyond mandible 0.5; SG 13.1; Sh-W 8.3; SL 59.8; snout to anterior angle of vent 54.4; AG 31.6; TL 45.6; tail depth at base 5.8; tail width at base 5.5; FLL 14.8; width of hand 4.5; HLL 16.4; FW 5.5; length of longest (third) toe 2.0; length of fifth toe 0.8. LI 1.5. Numbers of teeth: premaxillary 12; maxillary 65; vomerine 43. Coloration (in life) of the holotype (Fig. 2 A). Dorsum and dorsal surface of head solid dark chocolate brown. Grey specks on snout, mostly pale grey above eyes. Iris dark brown. Sides of head mostly brown with some grey speckling, with proportion of grey increasing toward back of head and mouth. Dorsal surface of tail same color as dorsum at insertion of hindlimbs, transitioning gradually to medium reddish brown at tip. Sides of body dark brown above midline, with some pale grey flecks, and pale grey mottled with dark brown below midline. Dorsal surface of limbs lighter reddish brown (similar to dorsal surface of tail); dorsal surface of feet brown with grey specks as on lateral surface of body. Gular region, lateral and ventral surfaces of tail, and underside of limbs light grey with brown speckles throughout. Venter dark brown with very small grey flecks. Underside of feet solid brown. Coloration (in alcohol) of the holotype. Dorsum and dorsal surface of head and limbs dark brown. Dorsal surface of tail solid medium red-brown. Small grey flecks above eyes, with larger grey flecks along labial surfaces, on rostrum and on sides of head below jaw. Sides of body mostly dark brown with few grey specks above midline and mostly grey with some dark brown specks below midline. Lateral and ventral surfaces of tail grey mottled with lighter brown compared to dorsum. Gular region and underside of limbs mostly pale grey mottled with some dark grey-brown. Venter slate grey with small specks of light grey throughout. Underside of feet slate grey. Color variation. Paratype IBH 14344 has a mostly grey venter, composed of tiny specks on a dark brown background. Dark brown blotches present on sides and at base of tail, at boundary between lateral and ventral coloration. Several lichen-like pale orange-brown blotches on dorsum. Lighter brown dorsal coloration compared to holotype. Almost no grey color above eyes. Other specimens lack gray mottling on ventrolateral surface and gular region, with underside of feet and ventral surface of tail light brown and light brownish color on rostrum and labial surfaces. Distribution. This species is known from the vicinity of the type locality near Huatusco, Veracruz, north to La Joya where it occurs in sympatry with P. cephalica, and from the northern Sierra de Zongolica near the locality of Zoncolco, 49 km south-southwest of the type locality (Fig. 1). The species is presumed to occur in cloud forest to the south of Huatusco between Huatusco and Zoncolco, but appears not to occur on Cerro Chicahuaxtla and other well-collected sites near the city of Cordoba. Natural history. Pseudoeurycea cafetalera appears to be a terrestrial species, like most members of its genus. It has been found under logs and in leaf litter in cafetal habitat with adjoining forest (Fig. 2 B). Cooccurring species of salamanders at the type locality include Bolitoglossa platydactyla, Chiropterotriton chiropterus, Parvimolge townsendi, and Thorius pennatulus. Etymology. This species is named for the coffee grove (cafetal) habitat where it is found. It is one of only three species of Pseudoeurycea (along with P. quetzalanensis and P. l i n e o l a) that has been found in this habitat so far. Phylogenetic results. The 16 S phylogeny (Fig. 3) provides strong support for the monophyly of Pseudoeurycea cafetalera (BS = 99, PP = 100). The P. cephalica group, which contains P. cafetalera, is also well supported (BS = 84, PP = 98) and is the sister clade to the P. bellii group (BS = 99, PP = 100). The relationship of P. cafetalera as the sister taxon to P. cephalica + P. quetzalanensis has little support (BS = 62, PP = 70). The divergence between P. cafetalera and P. cephalica ranged from 4.5 –6.0% (mean = 5.4 %), and the divergence between the two populations (Huatusco and Zongolica) of P. cafetalera is 2.6 %. The divergence between P. cafetalera and P. quetzalansis ranged from 5.0– 5.8 % (mean = 5.6 %). The divergence between P. cafetalera and P. cephalica + P. quetzalanensis (4.4 –6.0%, mean = 5.4 %) is comparable to that between other species of Pseudoeurycea (Fig. 3). Sequence divergence within the P. cephalica group was as high as 10.1 % (between P. galeanae and P. cafetalera).

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

Cruz, Gabriel, Murrieta-Galindo, Rene, Wake, David B. (2010): A new species of Pseudoeurycea from the cloud forest in Veracruz, México. Zootaxa 2725: 57-68, DOI: 10.5281/zenodo.199975

Bolitoglossa robinsoni sp. nov. Robinson���s Web-footed Salamander Figure 4 Holotype. UCR 11216, an adult male from Cerro Echandi (top of Cerro 3167 m), Prov. Puntarenas, Costa Rica, latitude 9 ��01��� 50 ��� N (9.0306), longitude 82 �� 49 ��� 20 ��� W (- 82.8222) (3.4 km max. error, WGS 84 datum), elevation 3162 m, collected on 6 January 1992 by Eugenia Arguedas. Paratypes. Three individuals: UCR 11217 -11218, same data as holotype; UCR 10921, Cerro Bur��, below Cerro Echandi, Prov. Puntarenas, Costa Rica, latitude 9 ��00��� 45 ��� N (9.0125, longitude 82 �� 49 ��� 50 ��� W (- 82.8306), elevation 2300 m, collected 16 October 1991 by M. Zumbado and E. Arguedas. Referred material. We tentatively assign morphologically divergent populations from nearby peaks to this taxon. See section on variation for detailed comments. Diagnosis. This is a relatively large, stout-bodied species assigned to Bolitoglossa because it lacks a sublingual fold. It resembles B. cerroensis in general form, but differs by having more slender habitus, more maxillary teeth and broader hands and feet, which however are less broad than those of the even larger B. marmorea. Differs from B. sombra by having more maxillary teeth, broader feet, shorter limbs, and lacking a uniform black coloration. Differs from B. compacta in having many more maxillary teeth and lacking extensive red or red-orange blotching or striping. Differs from all other salamanders in the region by its large size, relatively long and moderately webbed digits and extensive yellowish spotting on the body, limbs and tail. Description. This is a relatively large, stout species. Standard length (SL) in two adult males is 45.9 and 63.5. Tails are moderate, shorter than SL; SL divided by tail length in two males is 1.34 and 1.06. The head is broad; SL divided by head width is 5.7 and 6.2. The head is well demarcated from the neck. The snout is broadly rounded and not prominent. Nostrils are small and nasolabial protuberances are poorly developed. Eyes are moderately prominent; they protrude slightly beyond the lateral margins of the head and are relatively frontal in orientation. Teeth are numerous. There are 2 and 6 premaxillary teeth in males. Maxillary teeth are 50 and 65 in the two adult males, and vomerine teeth are 22 in each. Limbs are stout but moderate in length; limb interval is 1.5 and 2 in males. Hands and feet are broad (6.0 and 7.4) with little webbing; fewer than two distal-most phalanges of the longest digits are free of webbing (Fig. 2, B). Digital tips are truncate and bear well-developed subterminal pads. Fingers, in order of decreasing length, are 3-4 - 2 - 1; toes are 3-4 - 2- 5 - 1. Postiliac glands are not evident. Measurements (in mm), limb interval and tooth counts of the male holotype. Head width 10.2; snout to gular fold (head length) 13.9; head depth at posterior angle of jaw 6.6; eyelid width 2.0; eyelid length 4.4; anterior rim of orbit to tip of snout 3.4; horizontal orbit diameter 2.8; interorbital distance between angle of eyes 4.9; interorbital distance between eyelids 4.9; length of groove extending posteriorly from eye 5.8; distance between nuchal groove and gular fold 6.3; snout to forelimb 20.0; distance separating external nares 2.8; snout projection beyond mandible 0.8; snout to posterior angle of vent (standard length) 63.5; snout to anterior angle of vent 58.6; axilla to groin 34.0; limb interval 2.0; tail length 59.7; tail width at base 5.4; tail depth at base 5.6; forelimb length (to tip of longest finger) 16.1; hind limb length 16.6; hand width 6.0; foot width 7.4; length of fifth toe 1.2; length of third toe 2.7. Numbers of teeth: premaxillary 6; maxillary 31-34; vomerine 10-12, arranged in a single row. Coloration of the holotype in alcohol (Fig. 4). Dorsal ground color dark brownish-black but with so much pale spotting that the animal appears light. Patches of light yellow have joined together along the trunk to produce a very complex pattern of light on dark. Costal grooves are mainly dark, and the intercostal folds are light. Limbs and tail are mottled dark brown and yellowish-gray. Venter mottled with dark and light but mainly pale because of spaces in the punctate melanophore background. Throat pale but with punctate melanophores. Tail venter dark with ragged light spots. Variation. We tentatively assign specimens from four localities on or near the continental divide in Panam�� and Costa Rica to this species (Fig. 3). However, these specimens differ in several respects from the type specimens and they may represent as many as four additional undescribed species. The four localities and the specimens associated with them are Valle del Silencio (Silencio in Fig. 3 and Table 1, elevation 2500 m, 9 ��06��� 30 ��� [9.1083], 82 �� 58 ���00��� [- 82.9667]), MVZ 193550���193559 (193550, 193553, 193556, 193558, 153550 used in morphometrical analysis, see below); Cerro Bin�� (3200 m, 9 ��05��� 12 ��� [9.0867], 82 �� 54 ��� 34 ��� [- 82.8928]), MVZ 222487 ��� 89 and UCR 11787 (MVZ specimens used in morphometrical analysis); Cerro Itamut (3000 m, 9 ��05��� 40 ��� [9.0944], 82 �� 53 ���00��� [- 82.8833]), MVZ 222485 ��� 86 and UCR 11783 ���11786 (all used in morphometrical analysis); Cerro F��brega (3335 m, 9 ��07���00��� [9.1167], 82 �� 52 ��� 40 ��� [- 82.8778]); MVZ 222477 ���222483, 222484 and UCR 11775 ���11782 (MVZ 222478 and 222480, and UCR 11775 ���11778 used in morphometrical analysis). Specimens as indicated plus the type series of B. robinsoni were included in a discriminant function analysis. We conducted several slightly different analyses. In the first we used SL, tail length, head width, head length, forelimb and hind limb length, axilla-groin, shoulder width, foot width, and total numbers of maxillary, premaxillary and vomerine teeth as separate characters (Table 1, Fig. 5). A second analysis excluded tails, because we could not always be certain whether a tail had regenerated completely or in part from external morphology. A third analysis included the tail but excluded the teeth (on the grounds that they are counts rather than measurements). A final analysis included only teeth. All specimens in the discriminant function analysis were classified in the appropriate group using all measurements (Wilks��� Lambda= 0.0025, F[60, 69]= 2.9923 p=Habitat and range. Bolitoglossa robinsoni is known from a section of the Cordillera de Talamanca on the continental divide that forms the boundary between Costa Rica and Panam��, and in the high F��brega Massif, which extends northward into Panam�� (Fig. 3). Some specimens were obtained from areas of paramo, but others are from nearby montane forests. Etymology. The species is named in honor of the late Douglas C. Robinson, who did a significant survey of amphibians and reptiles in Costa Rica, consolidated the UCR collection and was the mentor of most herpetologists still present in the country., Published as part of Bola��os, Federico & Wake, David B., 2009, Two new species of montane web-footed salamanders (Plethodontidae: Bolitoglossa) from the Costa Rica-Panam�� border region, pp. 57-68 in Zootaxa 1981 on pages 61-66, DOI: 10.5281/zenodo.185285

Bolitoglossa pygmaea sp. nov. Pygmy Web-footed Salamander Figure 1 Holotype. UCR 11788, an adult female from F��brega Massif (coordinates 9 ��07���00��� N, 82 �� 52 ��� 40 ��� W) at 3100 m elevation, Provincia de Bocas del Toro, Panam��, collected 1-10 March, 1984 by Luis Diego G��mez. Paratypes. Twenty-five individuals with the same collecting and locality data as holotype: UCR 17789 ��� 11799, MVZ 222490���222503. Diagnosis. A very small species of Bolitoglossa (subgenus Eladinea) with a short tail and moderately webbed hands and feet that differs from all other species in the genus in its pale coloration and its blackpigmented stomach and peritoneum, which are apparent through the largely depigmented skin. Assigned to Bolitoglossa because it lacks a sublingual fold. Description. This diminutive species is in the small size-class category for the genus (Wake & Brame 1972). Standard length ranges from 23.6-32.6 (mean 28.1) in fifteen male paratypes; SL is 23.5-36.8 (mean 29.8) in nine adult females. Tails are short and never exceed standard length; SL divided by tail length ranges from 1.25 to 1.54 in males and from 1.23 to 1.85 in females. The head is relatively broad (Wake & Brame, 1972); SL divided by head width equals 5.5���6.6 in males and 5.6���6.5 in females. The snout is broadly rounded and of moderate length. Nostrils are large for this genus, although nasolabial protuberances are slight, better developed in males than in females. Eyes are relatively large and prominent and protrude beyond the lateral margins of the head and are visible in ventral view. Teeth are relatively large and well developed, and moderate in number (but relatively numerous in relation to body size). There are 2 or 3 premaxillary teeth (mean 2.1) in males and 3-6 (mean 4) in females. Maxillary teeth number 27���44 (mean 34.6) in males and 18���60 (mean 39) in females. Vomerine teeth number 14���20 (mean 16.3) in males and 15���25 (18.9) in females. Limbs are relatively short; limb interval ranges from 2.0 to 3.0 in males and from 2.5 to 4.5 in females. Hands and feet are relatively large; foot width ranges from 2.1 to 3 (mean 2.6) in males and from 1.9 to 3.5 (mean 2.5) in females. Digits are well differentiated but broadly pointed and more joined basally than webbed (Fig. 2 A). The first digit of the manus and the first and fifth of the pes are very small. The longer digits of the larger specimens bear small subterminal pads. About one and one-half phalanges of the longest digits are free of the basal digital fusion (which constitutes the webbing in this species). Fingers, in order of decreasing length, are 3-4 - 2 - 1; toes are 3-4 - 2-5 - 1. Postiliac glands are obscure, but collections of unicellular glands are prominent behind the hind limbs on the lateral sides of the body anterior to the vent. Measurements (in mm), limb interval and tooth counts of the female holotype. Head width 5.3; snout to gular fold (head length) 8.2; head depth at posterior angle of jaw 3.7; eyelid width 1.2; eyelid length 2.1; anterior rim of orbit to tip of snout 2.0; horizontal orbit diameter 1.7; interorbital distance between angle of eyes 2.9; interorbital distance between eyelids 3.2; snout to forelimb 10.3; distance separating external nares 1.3; snout projection beyond mandible 0.9; shoulder width 3.7; snout to posterior angle of vent (standard length) 33.7; snout to anterior angle of vent 30.5; axilla to groin 18.2; limb interval (number of costal interspaces between adpressed limbs) 2.5; tail length 18.2; tail width at base 1.6; tail depth at base 2.4; forelimb length (to tip of longest finger) 7.4; hind limb length 7.2; hand width 2.4; foot width 2.9; length of fifth toe 0.8; length of third toe 1.1. Numbers of teeth: premaxillary 3; maxillary 27-28; vomerine 10-12. Coloration of the holotype in alcohol (Fig. 1). Very pale general pigmentation, essentially transparent or translucent. Line of brown pigment from snout along canthus rostralis and through eye (one edge of eyelid) and behind eye in line over each shoulder, where it becomes a diffuse and irregular "line" that extends to the pelvic region. Pigment extends in a pair of weakly defined lines on tail nearly to tip. Scattering of brownish color on top of head, behind eye, and lightly on dorsum, but no evident ground color. Limb insertions, limbs, and hands and feet with lightly scattered brown pigment. Venter almost unpigmented with pale ventrolateral brownish color and some punctate melanophores thinly scattered over ventral surfaces. On the tail venter and behind hind limbs, transparent gland openings stand out as silvery circles. Internal black pigmentation in posterior abdomen and stomach shows through semitransparent ventral and lateral body wall, and heavily pigmented eyeball shows as very dark patch through eyelid. Iris dark brownish-black. Osteology. The following account is based on radiographs of 12 paratypes, one of which (MVZ 222492) was cleared and stained. Although small, this species has a well developed, well articulated skull. The unpaired premaxilla is small but rather robust and is well articulated to the well-formed maxillary bones. An unpaired dorsal process rises from the dorsum of the tooth-bearing portion of the premaxillary between the enlarged, cartilaginous nasal capsules; at the point where the nasal capsules diverge laterally the process splits into a pair that embraces a relatively large internasal fontanelle. These processes, well articulated to the nasals on their lateral borders, are slender and unexpanded distally. They end about half-way along the length of the fontanelle, where they articulate with the enlarged anterior parts of the frontals. The frontals articulate with each other, and the dorsal fontanelle often found in tropical plethodontids between the frontal and parietal bones is virtually absent. While well formed and relatively large, nasals are not protuberant anteriorly. A small lateral evacuation of the nasals expands internally and marks the passage of the nasolacrimal duct. Septomaxillaries and prefrontals are absent and the nasals form the anterior rim of the orbit. The nasals also articulate with the frontals. No otic crests are present. The operculum lacks a columella but has a minute anteroventral projection that contacts the otic capsule. The preorbital processes of the vomer are long and well developed, and bear teeth nearly to their tips. A large fontanelle separates the bodies of the vomers from each other medially. The 14 trunk vertebrae are well formed and bear bicipital ribs on the first 13. There are two caudosacral vertebrae. Tails are short, and several specimens have lost the tail. Six males have from 22 to 25 (mean 23) caudal vertebrae; two females have 22 and 24 caudal vertebrae. The first caudal vertebra has relatively stout, dorsoventrally expanded, rather short transverse processes that arise at the anterior end of the vertebra and are frontally oriented, but not at an acute angle. The limbs of the cleared and stained individual are in poor condition, but eight mesopodial elements are present in both the manus and pes (distal tarsal 3 is separate from a combined distal tarsal 4 + 5). Only digit 5 is complete on one limb and it contains two phalanges. The radiographs do not always display phalanges clearly, but when evident the phalangeal formulas are 1-2 - 3 - 2 and 1-2 - 3 - 3 - 2. All mesopodials are cartilaginous. No tibial spur is present. The hyoid apparatus is well stained and entirely cartilaginous. The basibranchial is relatively stout (diameter at midpoint about 14 % of length) and bears two short cornua at the broad anterior end. No lingual cartilage is present. The first ceratobranchial is extraordinarily slender (diameter 2.5 % of length and apparently discontinuous; it is strongly bowed). The second ceratobranchial, in contrast, is short and stout, with a length of about 50 % that of the basibranchial and about 75 % that of the first ceratobranchial. Its diameter at midpoint is 27 % of its length. The epibranchial is elongated and slender, tapering to a pointed tip; it is 2.3 times the length of the basibranchial, 3.4 times the length of the first ceratobranchial, and 4.5 times the length of the second ceratobranchial. The ceratohyals are relatively short and lack a slender anterior extension, and they are relatively narrow. There is no urohyal. Habitat and range. The species is known only from a small region called the F��brega Massif, a general plateau with a mean elevation above 3000 m. The main peak, Cerro F��brega, is 3335 m high (9 �� 07' N, 82 �� 53 ' W) and 3.5 km north of the Costa Rican border on the continental divide (the peak here is Cerro Bin��, Fig. 3). The plateau is mainly to the north of the peak, and according to the collector the salamanders were found among grass tussocks in an open area. Etymology. This species name is derived from pygmaeus (L.) and pygmaios (Gr.), in allusion to its miniature size, perhaps the smallest of any member of this large genus., Published as part of Bola��os, Federico & Wake, David B., 2009, Two new species of montane web-footed salamanders (Plethodontidae: Bolitoglossa) from the Costa Rica-Panam�� border region, pp. 57-68 in Zootaxa 1981 on pages 58-61, DOI: 10.5281/zenodo.185285, {"references":["Wake, D. B. & Brame, A. H., Jr. (1972) New species of salamanders (genus Bolitoglossa) from Colombia, Ecuador and Panama. Contributions to Science, Natural History Museum, Los Angeles County, 219, 1 - 34."]}

Bolitoglossa robinsoni sp. nov. Robinson’s Web-footed Salamander Figure 4 Holotype. UCR 11216, an adult male from Cerro Echandi (top of Cerro 3167 m), Prov. Puntarenas, Costa Rica, latitude 9 °01’ 50 ” N (9.0306), longitude 82 ° 49 ’ 20 ” W (- 82.8222) (3.4 km max. error, WGS 84 datum), elevation 3162 m, collected on 6 January 1992 by Eugenia Arguedas. Paratypes. Three individuals: UCR 11217 -11218, same data as holotype; UCR 10921, Cerro Burú, below Cerro Echandi, Prov. Puntarenas, Costa Rica, latitude 9 °00’ 45 ” N (9.0125, longitude 82 ° 49 ’ 50 ” W (- 82.8306), elevation 2300 m, collected 16 October 1991 by M. Zumbado and E. Arguedas. Referred material. We tentatively assign morphologically divergent populations from nearby peaks to this taxon. See section on variation for detailed comments. Diagnosis. This is a relatively large, stout-bodied species assigned to Bolitoglossa because it lacks a sublingual fold. It resembles B. cerroensis in general form, but differs by having more slender habitus, more maxillary teeth and broader hands and feet, which however are less broad than those of the even larger B. marmorea. Differs from B. sombra by having more maxillary teeth, broader feet, shorter limbs, and lacking a uniform black coloration. Differs from B. compacta in having many more maxillary teeth and lacking extensive red or red-orange blotching or striping. Differs from all other salamanders in the region by its large size, relatively long and moderately webbed digits and extensive yellowish spotting on the body, limbs and tail. Description. This is a relatively large, stout species. Standard length (SL) in two adult males is 45.9 and 63.5. Tails are moderate, shorter than SL; SL divided by tail length in two males is 1.34 and 1.06. The head is broad; SL divided by head width is 5.7 and 6.2. The head is well demarcated from the neck. The snout is broadly rounded and not prominent. Nostrils are small and nasolabial protuberances are poorly developed. Eyes are moderately prominent; they protrude slightly beyond the lateral margins of the head and are relatively frontal in orientation. Teeth are numerous. There are 2 and 6 premaxillary teeth in males. Maxillary teeth are 50 and 65 in the two adult males, and vomerine teeth are 22 in each. Limbs are stout but moderate in length; limb interval is 1.5 and 2 in males. Hands and feet are broad (6.0 and 7.4) with little webbing; fewer than two distal-most phalanges of the longest digits are free of webbing (Fig. 2, B). Digital tips are truncate and bear well-developed subterminal pads. Fingers, in order of decreasing length, are 3-4 - 2 - 1; toes are 3-4 - 2- 5 - 1. Postiliac glands are not evident. Measurements (in mm), limb interval and tooth counts of the male holotype. Head width 10.2; snout to gular fold (head length) 13.9; head depth at posterior angle of jaw 6.6; eyelid width 2.0; eyelid length 4.4; anterior rim of orbit to tip of snout 3.4; horizontal orbit diameter 2.8; interorbital distance between angle of eyes 4.9; interorbital distance between eyelids 4.9; length of groove extending posteriorly from eye 5.8; distance between nuchal groove and gular fold 6.3; snout to forelimb 20.0; distance separating external nares 2.8; snout projection beyond mandible 0.8; snout to posterior angle of vent (standard length) 63.5; snout to anterior angle of vent 58.6; axilla to groin 34.0; limb interval 2.0; tail length 59.7; tail width at base 5.4; tail depth at base 5.6; forelimb length (to tip of longest finger) 16.1; hind limb length 16.6; hand width 6.0; foot width 7.4; length of fifth toe 1.2; length of third toe 2.7. Numbers of teeth: premaxillary 6; maxillary 31-34; vomerine 10-12, arranged in a single row. Coloration of the holotype in alcohol (Fig. 4). Dorsal ground color dark brownish-black but with so much pale spotting that the animal appears light. Patches of light yellow have joined together along the trunk to produce a very complex pattern of light on dark. Costal grooves are mainly dark, and the intercostal folds are light. Limbs and tail are mottled dark brown and yellowish-gray. Venter mottled with dark and light but mainly pale because of spaces in the punctate melanophore background. Throat pale but with punctate melanophores. Tail venter dark with ragged light spots. Variation. We tentatively assign specimens from four localities on or near the continental divide in Panamá and Costa Rica to this species (Fig. 3). However, these specimens differ in several respects from the type specimens and they may represent as many as four additional undescribed species. The four localities and the specimens associated with them are Valle del Silencio (Silencio in Fig. 3 and Table 1, elevation 2500 m, 9 °06’ 30 ” [9.1083], 82 ° 58 ’00” [- 82.9667]), MVZ 193550–193559 (193550, 193553, 193556, 193558, 153550 used in morphometrical analysis, see below); Cerro Biné (3200 m, 9 °05’ 12 ” [9.0867], 82 ° 54 ’ 34 ” [- 82.8928]), MVZ 222487 – 89 and UCR 11787 (MVZ specimens used in morphometrical analysis); Cerro Itamut (3000 m, 9 °05’ 40 ” [9.0944], 82 ° 53 ’00” [- 82.8833]), MVZ 222485 – 86 and UCR 11783 –11786 (all used in morphometrical analysis); Cerro Fábrega (3335 m, 9 °07’00” [9.1167], 82 ° 52 ’ 40 ” [- 82.8778]); MVZ 222477 –222483, 222484 and UCR 11775 –11782 (MVZ 222478 and 222480, and UCR 11775 –11778 used in morphometrical analysis). Specimens as indicated plus the type series of B. robinsoni were included in a discriminant function analysis. We conducted several slightly different analyses. In the first we used SL, tail length, head width, head length, forelimb and hind limb length, axilla-groin, shoulder width, foot width, and total numbers of maxillary, premaxillary and vomerine teeth as separate characters (Table 1, Fig. 5). A second analysis excluded tails, because we could not always be certain whether a tail had regenerated completely or in part from external morphology. A third analysis included the tail but excluded the teeth (on the grounds that they are counts rather than measurements). A final analysis included only teeth. All specimens in the discriminant function analysis were classified in the appropriate group using all measurements (Wilks’ Lambda= 0.0025, F[60, 69]= 2.9923 p= *Variable not included in discriminant function analysis Despite the results of the discriminant function analysis there is little to distinguish these populations in particular characters. The number of costal folds between adpressed limbs gives an integrated measure (although inexact) that is useful for quick identification of many species of salamanders. In the small Echandi population this value ranges from 1.5-3.5, which is a value larger than for any of the other populations (total range of values from - 1.5 to + 1.5). Habitat and range. Bolitoglossa robinsoni is known from a section of the Cordillera de Talamanca on the continental divide that forms the boundary between Costa Rica and Panamá, and in the high Fábrega Massif, which extends northward into Panamá (Fig. 3). Some specimens were obtained from areas of paramo, but others are from nearby montane forests. Etymology. The species is named in honor of the late Douglas C. Robinson, who did a significant survey of amphibians and reptiles in Costa Rica, consolidated the UCR collection and was the mentor of most herpetologists still present in the country.

Bolitoglossa pygmaea sp. nov. Pygmy Web-footed Salamander Figure 1 Holotype. UCR 11788, an adult female from Fábrega Massif (coordinates 9 °07’00” N, 82 ° 52 ’ 40 ” W) at 3100 m elevation, Provincia de Bocas del Toro, Panamá, collected 1-10 March, 1984 by Luis Diego Gómez. Paratypes. Twenty-five individuals with the same collecting and locality data as holotype: UCR 17789 – 11799, MVZ 222490–222503. Diagnosis. A very small species of Bolitoglossa (subgenus Eladinea) with a short tail and moderately webbed hands and feet that differs from all other species in the genus in its pale coloration and its blackpigmented stomach and peritoneum, which are apparent through the largely depigmented skin. Assigned to Bolitoglossa because it lacks a sublingual fold. Description. This diminutive species is in the small size-class category for the genus (Wake & Brame 1972). Standard length ranges from 23.6-32.6 (mean 28.1) in fifteen male paratypes; SL is 23.5-36.8 (mean 29.8) in nine adult females. Tails are short and never exceed standard length; SL divided by tail length ranges from 1.25 to 1.54 in males and from 1.23 to 1.85 in females. The head is relatively broad (Wake & Brame, 1972); SL divided by head width equals 5.5–6.6 in males and 5.6–6.5 in females. The snout is broadly rounded and of moderate length. Nostrils are large for this genus, although nasolabial protuberances are slight, better developed in males than in females. Eyes are relatively large and prominent and protrude beyond the lateral margins of the head and are visible in ventral view. Teeth are relatively large and well developed, and moderate in number (but relatively numerous in relation to body size). There are 2 or 3 premaxillary teeth (mean 2.1) in males and 3-6 (mean 4) in females. Maxillary teeth number 27–44 (mean 34.6) in males and 18–60 (mean 39) in females. Vomerine teeth number 14–20 (mean 16.3) in males and 15–25 (18.9) in females. Limbs are relatively short; limb interval ranges from 2.0 to 3.0 in males and from 2.5 to 4.5 in females. Hands and feet are relatively large; foot width ranges from 2.1 to 3 (mean 2.6) in males and from 1.9 to 3.5 (mean 2.5) in females. Digits are well differentiated but broadly pointed and more joined basally than webbed (Fig. 2 A). The first digit of the manus and the first and fifth of the pes are very small. The longer digits of the larger specimens bear small subterminal pads. About one and one-half phalanges of the longest digits are free of the basal digital fusion (which constitutes the webbing in this species). Fingers, in order of decreasing length, are 3-4 - 2 - 1; toes are 3-4 - 2-5 - 1. Postiliac glands are obscure, but collections of unicellular glands are prominent behind the hind limbs on the lateral sides of the body anterior to the vent. Measurements (in mm), limb interval and tooth counts of the female holotype. Head width 5.3; snout to gular fold (head length) 8.2; head depth at posterior angle of jaw 3.7; eyelid width 1.2; eyelid length 2.1; anterior rim of orbit to tip of snout 2.0; horizontal orbit diameter 1.7; interorbital distance between angle of eyes 2.9; interorbital distance between eyelids 3.2; snout to forelimb 10.3; distance separating external nares 1.3; snout projection beyond mandible 0.9; shoulder width 3.7; snout to posterior angle of vent (standard length) 33.7; snout to anterior angle of vent 30.5; axilla to groin 18.2; limb interval (number of costal interspaces between adpressed limbs) 2.5; tail length 18.2; tail width at base 1.6; tail depth at base 2.4; forelimb length (to tip of longest finger) 7.4; hind limb length 7.2; hand width 2.4; foot width 2.9; length of fifth toe 0.8; length of third toe 1.1. Numbers of teeth: premaxillary 3; maxillary 27-28; vomerine 10-12. Coloration of the holotype in alcohol (Fig. 1). Very pale general pigmentation, essentially transparent or translucent. Line of brown pigment from snout along canthus rostralis and through eye (one edge of eyelid) and behind eye in line over each shoulder, where it becomes a diffuse and irregular "line" that extends to the pelvic region. Pigment extends in a pair of weakly defined lines on tail nearly to tip. Scattering of brownish color on top of head, behind eye, and lightly on dorsum, but no evident ground color. Limb insertions, limbs, and hands and feet with lightly scattered brown pigment. Venter almost unpigmented with pale ventrolateral brownish color and some punctate melanophores thinly scattered over ventral surfaces. On the tail venter and behind hind limbs, transparent gland openings stand out as silvery circles. Internal black pigmentation in posterior abdomen and stomach shows through semitransparent ventral and lateral body wall, and heavily pigmented eyeball shows as very dark patch through eyelid. Iris dark brownish-black. Osteology. The following account is based on radiographs of 12 paratypes, one of which (MVZ 222492) was cleared and stained. Although small, this species has a well developed, well articulated skull. The unpaired premaxilla is small but rather robust and is well articulated to the well-formed maxillary bones. An unpaired dorsal process rises from the dorsum of the tooth-bearing portion of the premaxillary between the enlarged, cartilaginous nasal capsules; at the point where the nasal capsules diverge laterally the process splits into a pair that embraces a relatively large internasal fontanelle. These processes, well articulated to the nasals on their lateral borders, are slender and unexpanded distally. They end about half-way along the length of the fontanelle, where they articulate with the enlarged anterior parts of the frontals. The frontals articulate with each other, and the dorsal fontanelle often found in tropical plethodontids between the frontal and parietal bones is virtually absent. While well formed and relatively large, nasals are not protuberant anteriorly. A small lateral evacuation of the nasals expands internally and marks the passage of the nasolacrimal duct. Septomaxillaries and prefrontals are absent and the nasals form the anterior rim of the orbit. The nasals also articulate with the frontals. No otic crests are present. The operculum lacks a columella but has a minute anteroventral projection that contacts the otic capsule. The preorbital processes of the vomer are long and well developed, and bear teeth nearly to their tips. A large fontanelle separates the bodies of the vomers from each other medially. The 14 trunk vertebrae are well formed and bear bicipital ribs on the first 13. There are two caudosacral vertebrae. Tails are short, and several specimens have lost the tail. Six males have from 22 to 25 (mean 23) caudal vertebrae; two females have 22 and 24 caudal vertebrae. The first caudal vertebra has relatively stout, dorsoventrally expanded, rather short transverse processes that arise at the anterior end of the vertebra and are frontally oriented, but not at an acute angle. The limbs of the cleared and stained individual are in poor condition, but eight mesopodial elements are present in both the manus and pes (distal tarsal 3 is separate from a combined distal tarsal 4 + 5). Only digit 5 is complete on one limb and it contains two phalanges. The radiographs do not always display phalanges clearly, but when evident the phalangeal formulas are 1-2 - 3 - 2 and 1-2 - 3 - 3 - 2. All mesopodials are cartilaginous. No tibial spur is present. The hyoid apparatus is well stained and entirely cartilaginous. The basibranchial is relatively stout (diameter at midpoint about 14 % of length) and bears two short cornua at the broad anterior end. No lingual cartilage is present. The first ceratobranchial is extraordinarily slender (diameter 2.5 % of length and apparently discontinuous; it is strongly bowed). The second ceratobranchial, in contrast, is short and stout, with a length of about 50 % that of the basibranchial and about 75 % that of the first ceratobranchial. Its diameter at midpoint is 27 % of its length. The epibranchial is elongated and slender, tapering to a pointed tip; it is 2.3 times the length of the basibranchial, 3.4 times the length of the first ceratobranchial, and 4.5 times the length of the second ceratobranchial. The ceratohyals are relatively short and lack a slender anterior extension, and they are relatively narrow. There is no urohyal. Habitat and range. The species is known only from a small region called the Fábrega Massif, a general plateau with a mean elevation above 3000 m. The main peak, Cerro Fábrega, is 3335 m high (9 º 07' N, 82 º 53 ' W) and 3.5 km north of the Costa Rican border on the continental divide (the peak here is Cerro Biné, Fig. 3). The plateau is mainly to the north of the peak, and according to the collector the salamanders were found among grass tussocks in an open area. Etymology. This species name is derived from pygmaeus (L.) and pygmaios (Gr.), in allusion to its miniature size, perhaps the smallest of any member of this large genus.

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

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

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