A new salamander of the genus Bolitoglossa (Caudata: Plethodontidae) from the highlands of western Panama Author Ponce, Marcos 0000-0002-6850-178X Bioconsultant, David, Provincia de Chiriquí, Panamá & https: // orcid. org / 0000 - 0002 - 6850 - 178 X Author Navarro, Deivy 0000-0002-7179-283X Protecma Panamá S. A. Santiago, Veraguas, Panamá & https: // orcid. org / 0000 - 0002 - 7179 - 283 X Author Morales, Roger 0000-0001-5511-2964 Bioconsultant, David, Provincia de Chiriquí, Panamá & Fundación Los Naturalistas, P. O. Box 0426 - 01459 David, Provincia de Chiriquí, Panamá & https: // orcid. org / 0000 - 0001 - 5511 - 2964 Author Batista, Abel Fundación Los Naturalistas, P. O. Box 0426 - 01459 David, Provincia de Chiriquí, Panamá & Vicerrectoría de Investigación y Postgrado, Universidad Autónoma de Chiriquí, Ciudad Universitaria El Cabrero David, Chiriquí, 427, Panamá & Asociación ADOPTA el Bosque Panamá, 119 x Gamboa, Panamá text Zootaxa 2022 2022-04-29 5129 4 543 556 journal article 55760 10.11646/zootaxa.5129.4.4 2e2fb97f-7d86-4289-8e27-bf81bfacc4db 1175-5326 6504104 B0368310-A732-4191-8F41-B213BC89BDCF Bolitoglossa cathyledecae sp. nov. Chiriqui fire salamander, Salamandra de fuego chiricana ( Figures 3–6 ) Holotype . MHCH 3240 (original field number MP 12 ), an adult male from Panama : Chiriquí province district Boquete , La Amistad International Park ( PILA ) ( 8.8635°N , 82.4864°W , 1,969 m ); collected by Deivy Navarro along with Carlos Castillo , Erick Nuñez and Marcos Ponce on 20 May 2017 at 19:30 hrs. Paratypes . MHCH 3241–42 , both are juveniles ( MP 175 , 185 ) collected 0.9 km ca. NE of the type locality ( 8.8682°N , 82.4844°W , 1983 m ) collected by Marcos Ponce and Roger Morales on 31 September 2017 , at 23:00 hrs. Diagnosis: Assigned to genus Bolitoglossa due to having fewer than 14 costal grooves and lacking a sublingual fold, and to subgenus Eladinea based on mtDNA sequence data. Bolitoglossa cathyledecae is differentiated from all known species of the subgenus Eladinea by the combination of the following characters: (1) Unique coloration consisting of pinkish-flesh with flame-scarlet speckles, ventral salmon color; (2) Moderately small size with wide head HW/SVL=0.21; (3) moderate -sized extremities; (4) digits with moderate to extensive webbing on the hands and feet; (5) prehensile and considerably longer tail than body TL/SVL=1.5; (6) high tooth counts: premaxillary (7), maxillary teeth (78). Similar species: Bolitoglossa cathyledecae is well differentiated from all other described species of the subgenus Eladinea by 16S mtDNA distances ( Fig. 2 ; Table 1 ). It is further distinguished morphologically by the following characteristics (condition for B. cathyledecae in parentheses): It differs from B. alvaradoi Taylor , B. biseriata Tanner , B. chucantiensis Batista, Köhler, Mebert and Vesely , B. colonnea Dunn , B. cuna , Wake , Brame, and Duellman B. lignicolor , B.medemi Brame and Wake , B. minutula Wake , Brame and Duellman, B. schizodactyla Wake and Brame, B. taylori Wake , Brame and Myers, by its less extensively webbed hands and feet. All these species have fully webbed hands and feet (well-developed digital webbing; only portions of distal-most phalanges of longest digits free of webbing). Bolitoglossa cathyledecae is easily distinguishable by its unique coloration (pinkish flesh with flame scarlet speckles and ventral salmon color) from the large black salamanders found between Costa Rica and Panamá , including the almost entirely black species B. anthracina Brame, Savage , Wake and Hanken; B. compacta Wake , Brame and Duellman; B. copia Wake , Hanken and Ibañez; B. magnifica Hanken , Wake and Savage; B. nigrescens Taylor ; B. robusta Cope ; and B. sombra Hanken , Wake and Savage. This new species is differentiated from other salamanders found in the Talamanca Mountain range between Costa Rica and Panama as follows: Bolitoglossa subpalmata Boulenger a moderate-sized species with SVL of 60 mm , ( 46 mm in adult male), slate-gray color with brownish flecking evident on the back; grayish venter, the median area darker than outer portions (pinkish flesh with flame scarlet speckles and ventral salmon color). Bolitoglossa pesrubra Taylor , is a moderate-sized salamander with an SVL = 67 mm , ( 46 mm ) and has fewer premaxillary and maxillary teeth, with 3 PMT (7), MT and 66 MT (78), has a shorter tail in proportion to body size, TL/SVL = 1, (1.5). Bolitoglossa marmorea a moderated-sized species with an SVL of 60.1 mm , ( 46 mm ) having longer legs, in which the toes of the adpressed legs are usually in contact (limb interval between toes of: 3, not in contact when adpressed). TABLE 1. Estimates of uncorrected p-distances between sequences of 16S mtDNA (in %), between Bolitoglossa subpalmata , B. epimela and B. robinsoni species group and B. cathyledecae sp. nov.

Species		1	2	3	4	5	6	7	8	9	10	11	12	13	14	15	16	17
1.	B. pesrubra
2.	B. kamuk	1.3
3.	B. anthracina	3.5	3.0
4.	B. splendida	3.2	2.6	3.7
5.	B. subpalmata	2.4	1.3	3.4	2.6
6.	B. tica	3.4	3.0	4.3	4.5	1.9
7.	B. gracilis	4.3	3.6	4.5	4.7	3.0	4.1
8.	B. bramei	3.2	2.4	3.9	3.9	2.6	3.9	3.0
9.	B. gomezi	3.6	2.6	4.1	3.9	3.0	4.1	4.7	3.0
10.	B. sooyorum	4.1	3.3	5.0	5.2	3.5	4.3	5.7	4.1	3.9
11.	B. minutula	3.5	2.6	5.0	4.6	3.3	4.8	5.4	3.9	3.7	1.7
12.	B. marmorea	3.9	3.0	5.3	5.0	3.3	4.3	5.0	3.9	3.7	2.0	2.0
13.	B. cerroensis	5.2	4.3	6.3	5.4	4.6	5.2	6.1	4.8	4.6	3.5	3.1	2.8
14.	B. epimela	6.3	5.5	5.9	6.1	5.2	5.9	6.8	5.0	5.2	5.0	5.3	4.8	4.2
15.	B. jugivagans	6.6	6.3	8.1	6.6	6.6	6.9	8.6	6.3	7.2	5.8	6.1	6.1	5.2	5.3
16.	B. robinsoni	5.6	4.9	6.5	6.0	5.2	6.0	6.9	5.4	5.6	4.8	4.8	5.0	4.8	6.1	4.0
17.	B. aureogularis	4.9	4.3	5.8	5.4	4.5	5.4	6.2	4.7	4.9	4.1	4.1	4.3	4.1	5.5	3.4	0.6
18.	B. cathyledecae sp. nov.	6.1	5.7	7.2	6.8	6.5	7.2	7.4	5.4	6.1	6.4	6.2	6.2	5.9	7.3	6.7	6.1	5.9

FIGURE 2. Maximum likelihood trees based on 16S mtDNA sequence of Bolitoglossa spp. of the Eladinea subgenus. SHaLRT (Shimodaira and Hasegawa approximate likelihood ratio test) support (%) / ultrafast bootstrap support (%) are shown on the internodes. Outgroups not shown. Bolitoglossa pygmaea Bolaños and Wake , B. gracilis Bolaños, Robinson and Wake and B. diminuta Robinson are small species with SVL ≤ 43 mm (46.0 mm), also, their hands and feet have less webbing, digital webbing well developed). Bolitoglossa robinsoni is a relatively large SVL in adult males with 45.9‒63.5 mm ( 46 mm ), tails are moderate, shorter in proportion to body size TL/SVL = 1.06‒1.34 (1.5); hands and feet with little webbing; fewer than two distal-most phalanges of the longest digits are free of webbing (digital webbing well developed; only portions of distal-most phalanges of longest digits free). Bolitoglossa gomezi Wake , Savage and Hanken has a shorter tail in proportion to body size TL/SVL: 1.15–1.19 (TL/SVL = 1.5) and a lower tooth count in PMT: 2 or 3 (7), MT: 29–52 in males (78), and VT: 17–30 (33). Bolitoglossa bramei Wake , Savage and Hanken as well has fewer premaxillary, maxillary and vomerine teeth PMT: 3‒4 (7), MT: 49–66 (78), VT: 24–28 (33), limbs long; limb interval 0–0.5 (3). Bolitoglossa splendida Boza-Oviedo, Rovito, Chaves, García-Rodríguez, Artavia, Bolaños and Wake presents a coloration with bright-red dorsal band and bright-yellow spots laterally and ventro-laterally (pinkish flesh with flame scarlet speckles and ventral salmon color). Bolitoglossa kamuk Boza-Oviedo, Rovito, Chaves, García-Rodríguez, Artavia, Bolaños and Wake has a dark ground coloration with golden flecking on the posterior part of the body and the tail (pinkish -flesh with flame scarlet speckles, ventral salmon color), as well as fewer maxillary and vomerine teeth MT: 34 (78), VT: 16 (33), and a shorter tail in proportion to body size TL/SVL ≈ 1 (TL/SVL = 1.5). Bolitoglossa aureogularis Boza-Oviedo, Rovito, Chaves, García-Rodríguez, Artavia, Bolaños and Wake with dorsal coloration golden-tan with some bright highlights on dorsolateral regions and with some narrow streaks of dark brown, gular area bright yellow, which becomes golden on chest before fading into darker color in the midtrunk region (pinkish flesh with flame scarlet speckles and ventral salmon color), Head is narrow in proportion to body size HW/SVL = 0.13 (0.21). Bolitoglossa jugivagans , Hertz, Lotzkat and Köhler is a small thin Bolitoglossa with an SVL of 31.2 mm ( 46 mm ) with a narrow head in proportion to body size HW/SVL = 0.14 (0.21), relatively shorter front limbs HLL about 20 % of SVL (HLL = 25.7 % of SVL). The last Bolitoglossa described in the Talamanca mountain range Bolitoglossa aurae Kubicki and Arias has a significantly different coloration, dorsum uniform light yellow with a thin dark brown middorsal stripe on the trunk, flanks light yellow to light brown, ventrum uniform translucent yellow and lacking any evident or contrasting light or dark dermal pigmentation or midventral stripe (pinkish flesh with flame scarlet speckles and ventral salmon color), relatively longer trunk in female AX = 58 % of SVL (51.3% in male holotype ), with 4.5 costal folds between adpressed limbs in female ( 3 in male holotype ). Description of the holotype . A moderately small species (SVL = 46.0 mm), head relatively broad compared to other members of genus; head long (HL about 29 % of SVL) and broad (HW = 21 % of SVL); tail long and prehensile, TL/SVL is 1.5; hands and feet broad (HAW = 10 %, HFW = 14% of SVL), and limbs moderate (HLL about 19 % of SVL, limb interval 3 costal folds between adpressed limbs); digital webbing well developed, only portions of distal-most phalanges of longest digits free ( Fig. 3 ). Fingers in order of decreasing length: 3, 4, 2, 1; toes in order of decreasing length: 3, 4, 5, 2, 1; subterminal pads present on longest digits; eyes relatively small; do not protrude beyond lateral margins of head and are not visible in ventral view; labial protuberances weakly developed; longer snout, ES = 43.75 % of HW. Teeth moderate in size and numerous (PMT 7, MT 78, VT 33). Digits of hands and feet easily distinguishable but extensively webbed; digital tips broadly rounded, hands and feet flattened; digits flattened, not cylindrical ( Fig. 3 ). Measurements (in mm), limb interval and tooth count of the male holotype ( Table 2 ). SVL: 46.0, TL: 68.8, HW: 9.6, SG: 13.2, HD: 5, EW: 1.9, EL: 3, ES: 4.2, ED: 2.9, IC: 4.9, IO: 3.5, SF: 17.6, IN: 2.7, SP: 0.8, SW: 8.2, SAV: 18.5, AX: 26.9, LI: 3.5, FLL: 10.8, HLL: 8.6, HAW: 4.8, FW: 6.4, T3: 1.4, T5: 0.9, Numbers of teeth: PMT: 7, MT: 40/38, VT: 17/16. FIGURE 3. Details of hand and foot of preserved holotype of Bolitoglossa cathyledecae sp. nov. (MHCH 3240); A) Right hand, left: dorsal view, right: ventral view; B) Right foot, left: dorsal view, right: ventral view. Scale bars: 5 mm. Photographs taken by MP. TABLE 2. Morphometric characters of Bolitoglossa cathyledecae sp. nov. holotype and paratypes. see methods for abbreviation explanations of abbreviations.

Measurements (mm)	Holotype	Paratype	Paratype
MP-12 (MHCH 3240)	MP-175 (MHCH 3241)	MP-185 (MHCH 3242)
Sex	Male	Male (subadult)	Female (subadult)
Length total (LT)	115.9	60.7	57.7
Snout-vent length (SVL)	46	30.9	25
Tail length (TL)	68.8	27.8	32.6
Axilla-groin (AG)	26.9	18.9	14.7
Head width (HW)	9.6	6.7	5.4
Hind limb length (HLL)	11.8	6.1	4.6
Maxillary teeth (MT)	78 (40/38)	36 (17/19)	25 (13/12)
Vomerine teeth (VT)	33 (17/16)	19 (10/11))	17 (8/9)

Coloration of the holotype in life ( Fig. 4 ). Dorsal ground of the head, back color Pinkish Flesh (253), with Flame Scarlet (73) speckles, upper part of the extremities similar in color to dorsal surface. Lateral and ventral ground coloration Salmon Color (251), with Pinkish Flesh (253) speckles, in the gular and ventral region, lower surface of the hands, and feet Light Pinkish Rose (217). Most of the tail similar in color to the background dorsal color with the tip of the tail Spectrum Yellow (78). Iris Pale Sulphur Yellow (92) with reticulum Flame Scarlet (73). Coloration in ethanol. After three years in ethanol (70%), the dorsal coloration of the holotype has darkened, the head and the anterior part of the trunk are Burnt Umber (48) while the rest of the trunk and tail are Amber (51). The ventral coloration of the specimen is mainly Clay Color (18) speckled with Drab (19) ( Fig. 5 ). FIGURE 4. Holotype of Bolitoglossa cathyledecae sp. nov. (MHCH 3240) in life; A–B) Details of head; C–D) Body coloration in life. Photograph by MP. FIGURE 5. Holotype after three years in 70 % ethanol; A ) Dorsal view; B ) ventral view. Scale bars= 2 cm. Photographs taken by MP. Variation ( Fig. 6 ). There are two paratypes (a male and a female) both juveniles, these specimens present a coloration like the holotype , but with the dorsal coloration Chrome Orange (74) and the smallest paratype presents the last third of the tail Spectrum Yellow (78). Etymology. The specific epithet of this beautiful new species honors Cathy Ledec, a passionate conservationist and long-time supporter of conservational organizations working to preserve the habitat of salamanders in the Neotropics. Habitat and natural history observations. Bolitoglossa cathyledecae , was found in a montane rain forest, characterized by a short dry season (one to three months, sensu Holdridge 1967 ). Annual precipitation oscillates between 3600 and 7500 mm and the annual temperature between 12ºC and 17ºC. At this site trees are covered with epiphytes and bryophytes, with a dense undergrowth vegetation ( Fig. 7 ). The holotype was found on 20 May 2017 at 19:30 h. active on a feather bamboo of the genus Chusquea sp. at 1 m above the ground, there was a light rain that day. Paratypes were found on September 31, 2017 between 20:00‒23:30 h active on ferns approximately 2m above the ground. The night was humid due to rain between 16:00 and 18:00 h. Other amphibian species collected in the vicinity of the type series include Bolitoglossa compacta , B. minutula , Craugastor blairi , Diasporus aff. hylaeformis , Incilius epioticus , Pristimantis caryophyllaceus , and Pristimantis museosus . Conservation status. This species is restricted to a small area of a high elevation habitat in the Talamanca Mountain range, suggesting it be classified as Critically Endangered (A3c, B2a [ii, iii, iv], according to the IUCN Red List categories and criteria (IUCN Standards and Petitions Committee. 2019). Anthropogenic pressure around the area will probably lead to further declines of populations through habitat deterioration and reduction of the area of occupancy. This species will also be allocated to “High” level of environmental vulnerability using the EVS of Johnson et al. (2015) . FIGURE 6. Color variation in the paratypes. A) Dorsum in daytime coloration (MHCH 3242); B-C) Details of head and dorsum in daytime coloration (MHCH 3241); D) Paratype at the moment of encounter at night (MHCH 3242). Photographs taken by MP. Distribution. Bolitoglossa cathyledecae is known only from one site within the montane rain forest life zone ( sensu Holdridge 1967 ) along the northeastern mid-elevation slopes of the Cordillera de Talamanca, in the vicinity of the continental divide between Chiriquí and Bocas del Toro , within the La Amistad National Park, district Boquete, at about 1900 m above sea level ( Fig. 1 ). Remarks. The striking color pattern and the differences in molecular makeup, support the recognition of Bolitoglossa cathyledecae as a species new to science in lower Central America, placed within the subgenus Eladinea (Parra et al. 2004) . Bolitoglossa cathyledecae sp. nov. , is the 23rd Bolitoglossa species present in Panama . The new species is phylogenetically related to the epimela , and subpalmata species group ( sensu Garcia-Paris et al. 2008 ) and shows the lowest genetic distance to B. bramei , a species of the subpalmata group, with distribution in southwestern Costa Rica and western Panama . Considering that the estimated average evolutionary divergence over sequence pairs within the groups mentioned above is as high as 3.5% (3.5% for epimela , and 3.3% for subpalmata ), contrary to the adspersa species group where the genetic distances between some species are higher (up to 11.9%, Batista et al . 2014 ; Jaramillo et al. 2020 ), along with the morphological distinctions found in B. cathyledecae compared to other species groups in the subgenus Eladinea ; we refrain from assigning B. cathyledecae to a species group at this time. Although the Talamancan clade in the phylogenetic analysis, with the epimela , subpalmata species group and cathyledecae was not supported by the ML analysis, it got strong support with the Bayesian analysis (in part). In both phylogenetic hypothesis, there is a paraphyly, having species of the epimela species group nested within species of the subpalmata species group ( Fig. 2 ), although there is one evident clade, with the northern subpalmata species from Costa Rica ( B. subpalmata , B. tica , B. gracilis , B. splendida , B. pesrubra , B. kamuk )). The evidence of the paraphyly found among the species in the subpalmata species group found in this study could be an artifact of the use of a single genetic marker, given that, the epimela and subpalmata clades have been well supported in other studies when using more than one molecular marker ( Kubicki and Arias 2016 , Jaramillo et al. 2020 ), although in those studies B. anthracina has not been included which could give a different framework of the relationship between groups. Taxonomic work is still needed in the highlands between Costa Rica and Panamá ( Garcia-Paris et al. 2008 ; Bolaños and Wake 2009 ; Boza-Oviedo et al. 2012; Kubicki and Arias 2016 ), with vast remote areas still unexplored, mainly along the Serranía de Talamanca. Cloud forests are vulnerable to climate change due to the low range of mobility of the upper highland endemic species ( Davies et al. 2004 ; Paaijmans et al. 2013 ). Consequently, species or populations restricted to such habitats are enormously susceptible to slight changes in the environment and thus face an elevated risk of extinction (Batista et al. 2020). An urgent conservation plan is required to protect the surrounding cloudforest and the entire La Amistad International Park to preserve its unique pool of endemic species.