A new species of sAlAmAnder ( Bolitoglossa : plethodontidAe ) from the cordillerA orientAl of the colombiAn Andes

Eight species of salamanders are recognized to Cordillera Oriental of Colombia. Here we describe a new species of the genus Bolitoglossa, named Bolitoglossa guaneae sp. nov. The highest number of species of this genus is found in the cloud forests located in the western flank of the Cordillera Oriental. Key-Words: Caudata; Bolitoglossa; New species; Cordillera Oriental.


Bolitoglossa guaneae sp. nov.
(Fig. 1)    1969) (Fig. 2); ventral surface of digit tips without terminal flattened tubercles (Fig. 2); snout short and rounded in lateral profile, head length 2.0-2.92mm in adult males, 1.88-2.88mm in adult females; snout rounded in dorsal view; protruding eyes on dorsal view; well-defined postcephalic constriction (Fig. 3).Ventral surfaces (in alcohol) brown with numerous tiny cream guanophores with irregular spots; nasolabial grooves cream on preserved specimens; males have white testis; adult males 1 or 2 premaxillary teeth pierce the lip; copper iris; Snout in profile lack numerous tiny cream guanophores; terminal phalanges of digits not expanded; pelvic girdle without posterolateral projections.Bolitoglossa guaneae differs from other Colombian salamanders in morphologic and chromatic characteristics.From species with basal webbing with free digital tips on the fingers and toes (Type A in South American species, sensu Brame & Wake, 1963, and type slightly and moderately webbed sensu Wake & Brame, 1969), as B. adspersa, B. hiemalis, B. hypacra, B. ramosi, B. savagei, B. tatamae, B. walkeri and B. vallecula (Fig. 4), by having extensively interdigital webbing with free digital tips on the fingers and toes (Type D in South American species, sensu Brame & Wake, 1963, and   A very distinct constriction at the base of the tail with a slight lateral compression; tail length versus SVL 67.08-89.96%(mean = 78.50,n = 9) in males, and 60.71-79.53%(mean = 72.04,n = 9) in females.Skin of dorsum with lateral and ventral smooth surfaces.
Color in Life: The dorsal and lateral ground color is dark brown, sometimes mottled or streaked with white, diffuse paled blue and cream guanophores, in some individuals reddish marks are also present.Hindlimbs are dark brown with reddish; Tail is light to dark brown, in occasions mottled with white; ventral surfaces are Brown with dense white cream irregular spots and guanophores blue very faint, sometimes mottled white, tail is ventrally mottled white in some individuals; copper iris without black reticulations (Fig. 1).

Coloration in Alcohol:
The ground color of head, trunk, and tail of the type series is dark brown, with palmar and plantar surfaces light brown.Venter dark brown with a few cream patches and cream guanophores.
Variation: The type series presents in general terms, a uniform color in all adult specimens; the dorsal color pattern with inconspicuous paler brown dorsolateral stripe, extending from posterior edge of eyelid dropping towards the base of thighs.
Osteology: The description is based on two specimens (ICN-MNH 8556, ICN-MNH 4418).Skull is oval and well formed, anterior elements are generally well ossified and articulated.Pars dentalis of the premaxilla are short with four to five teeth.The internasal fontanelle is relatively narrow.Nasal bones are protrusive anteriorly and articulate with maxillaries, frontals, and premaxillaries.Prefrontal bones are nearly rectangular, with irregular edges, slightly longer along the posterior rather than the anterior ridge.Frontal bones occupy nearly half the length of the skull, irregular in shape; they overlap each other and the parietal bones.Irregular parietal The right maxilla bears 13-21 teeth and the left one 16-24.Vomers triangular and rounded, bearing 19-28 teeth transversely positioned on the proximal end of the vomer, and wide internal nares.Antorbitalis cartilage is articulated to the posterior edge of the vomer, laterally to the maxilla and posteriorly to the orbitosphenoid bone.A large triangular parasphenoid bone making part of the cranial vault, extended from the vomer interspace to the ear capsules, bearing two plates covered with teeth arranged in short and oblique rows: the right plate bears 13-14 rows with 70-122 teeth, and the left one supports 12-13 rows with 73-138 teeth.Orbitosphenoid rectangular, with articular processes at anterior and posterior rims, and articulating with antorbitalis and pterygoid cartilages anterior and posterior, respectively.A large optic fenestrum; squamosal rectangular, irregular, and thin, investing the palatoquadrate laterally and articulating with the skull roof.The palatoquadrate is small, distally articulating with the articular by cartilaginous process, while the proximal end articulates to the posterior branch of the pterygoid.The right dentary bears 26-42 teeth and the left one 22-46.Fourteen presacral vertebrae, two caudosacral and 24 caudal.The neural arch of the atlas bears a cartilaginous spine.Transverse process forked, perpendicular to the body axis on the posterior vertebrae.Ribs curved; at the distal end of the second vertebra is a V-shaped cartilaginous plate.There are no carpal or tarsal fusions nor reductions.Metacarpi are robust.The phalangeal formula in hands is 1-2-3-2; in feet is 1-2-3-2-2; wide foot and robust metatarsi.
Hyobranchial apparatus cartilaginous; basibranchial cylindrical, with a rectangular posterior end articulating with the visceral arches while its anterior end is laterally extended; ceratobranchial is anteroposteriorly directed from the medial region of the basibranchial, having a wider and more irregular base; the origin of ceratobranchial II is at the posterior end of the basibranchial.The ceratohyal articulates with the distal end of the hypohyal, having narrow plates at the base, and cylindrical ones on the posterior half.and Boyacá from elevations between 1700-2400 m.Annual rainfall ranges between 1400-2000 mm (Fig. 7).

Measurements of Holotype
Habitat and Behavior: Bolitoglossa guaneae is a rare nocturnal species, restricted to near streams in primary or secondary forests with little anthropic intervention.("low montane humid forest" (bh-mb) sensu Holdridge 1996, see also Gutiérrez-Lamus et al. 2004).This species was found standing on low leaves (0,50-1,30 m) of Araceae, Melastomataceae, and ferns.The Holotype was collected on a leaf 0.5 m above ground at 22:06 h, with temperature of 21.1°C and 91% relative humidity.

dIscussIon
The recognition of Bolitoglossa guaneae as a new species is justified by morphological characters such small size, extensively interdigital webbing with free digital tips on the fingers and toes, triangular toe in third finger, well defined postcephalic constriction and terminal phalanges of digits not expanded.It seems that B. guaneae is more closely related to B. pandi (from southwest of Cordillera Oriental of Colombia) and B. phalarosoma (Cordillera Central of Colombia; Acosta-Galvis, 2007).It is endemic to humid relict forests on the western slopes of the Cordillera Oriental, and its known extent of occurrence is about 176.2 km.This distributional pattern is similar to several species of Bolitoglossa restricted in the cloud forests of the western slopes of Cordillera Oriental region as B. capitana, B. pandi and B. nicefori.Bolitoglossa guaneae is should be considered as Vulnerable (A4ce+B2ab(i,iii)+D1; IUCN, 2001).The population remains stable, but there is a rapid reduction and fragmentation of their habitats, and the presence of the pathogenic fungus Batrachochytrium dendrobatidis in its area of distribution (Ruiz & Rueda-Almonacid, 2008).
For other species associated to the Cordillera Oriental as Bolitoglossa altamazonica, it is was found to be common from July to December, associated to the vegetation next to creeks or occasionally under roots of bushes.Brame & Wake (1963) consider that the geographic distribution of this species spreads out to the Magdalena river valley; nevertheless, it is probable that the specimen observed by them belongs to B. lozanoi, which has been recorded at 1100 m in the Municipality of Yacopí, close to Muzo (1200 m) by the authors mentioned above (Fig. 8).Besides, Crump (1977) studied populations from the Peruvian Amazon and related them to the substrate of the interior of "terra firme" and "varzea" forests in the Amazon basin perching on shrubs.Almendáriz et al. (2004) and Cisneros-Heredia (2006) misidentified the Colombian specimens deposited in ICN-MNH collection as B. ecuatoriana.
A recent review of the specimens of B. capitana deposited in the collection of the Universidad de la Salle reveals that the numbers used by the reference collection for the original description of this species must be reassigned as follows: the holotype which used to be MLaS 1 sensu Brame & Wake (1963) (= MLS 119 Nicéforo-María, 1958) is now MLS 183; both paratypes (one male and one female) recognized by Brame & Wake (1963) as MLaS 1 and MLaS 1b are now MLS 182, and MLS 184 respectively.With reference to its large size it is important to note that other species geographically related to B. capitana, such as B. adspersa and B. nicefori can be almost as large as this species.Bolitoglossa capitana is considered as critically endangered by IUCN and has no records since 1975.A specimen deposited at the Natural Sciences Institute of Bogotá (ICN-MNH 9221) exhibits subtle differences from the type specimens collected in 1945 by Brother Nicéforo-María, in which the ventral surfaces, the throat, chest and belly is colored purplish brown with dark brown guanophores diffuse; palms and soles brown, clear guanophores with cream.Chavés-Portilla et al. (2006) mistakenly identified an undescribed species of this genus as B. capitana; this new species differs from B. capitana by its ventral color pattern, more extensive webbing of hand and feet and its smaller size.
Bolitoglossa lozanoi is a nocturnal and arboreal species related to the interior of the underbrush of primary forest close to streams.Specimens were observed perching on Araceae and shrubs between 0.10-2 m above the ground.This species has a seasonal reproduction pattern and is commonly found between August and November.Considered as vulnerable by the red list of Colombian amphibians (Acosta-Galvis, 2004), it is actually widely distributed in the Magdalena valley (Fig. 9).It was erroneously recorded as B. altamazonica in the Muzo area by Brame & Wake (1963), while this locality is close to the collection place of B. lozanoi by Acosta-Galvis (2004) in the Department of Cundinamarca.
Finally, Bolitoglossa palmata is first reported to Colombia based on five specimens of (Fig. 10

AcknowledgeMents
This contribution constitutes part of the project "Taxonomy and homology character evaluation for salamanders of the genus Bolitoglossa (Caudata: Plethodontidae) from Colombia" by Laboratory of Amphibians of Natural Sciences institute of the National University.Additionally the description of the new species was carried out and the Universidad Industrial de Santander (Group of studies in Biodiversity).We thank to Nancy Rivera director of the Fauna and Flora Sanctuary Guanentá-Alto Rio Fonce.We also thank to the following curators for the loan of material: Fernando Castro from Universidad del Valle, Martha Patricia Ramirez from Universidad Industrial de Santander, John D. Lynch from the Instituto de Ciencias Naturales of the Universidad Nacional de Colombia, Vivian Páez from Universidad de Antioquia, Brother Roque Casallas, and Arturo Rodriguez from the Universidad de La Salle, Julio Mario Hoyos from Javeriana University, finally to Perico Delgado from the Instituto Alexander von Humboldt.We thank, J.D. Lynch, Benjamin Tapley and Lilia Svietlana Torres for comments on the manuscript and who helped us translate.Finally, the anonymous reviewers for valuable comments on this paper.
category D, sensu Wake & Brame, 1969) (Fig. 2).From species with extensively or completely webbing (sensu Wake & Brame, 1969 and Types C, D. E and F sensu Brame & Wake, 1963), Bolitoglossa guaneae can be differentiated from B. altamazonica in having extensively interdigital webbing with a longer and triangular third finger (completely webbing and tips rounded in B. altamazonica) (Fig. 5); differs from B. biseriata and B. silverstonei in having a ventral surface brown with white dots (ventral surface cream intensified by numerous small irregular light brown spots in B. biseriata and B. silverstonei), white testes in males (usually black testes in B. biseriata and B. silverstonei) and extensively interdigital webbing with free digital tips on the fingers and toes (being completely webbing without free digital tips in B. biseriata and B. silverstonei); from B. medemi in the absence of digital depressions (present in B. medemi); B. guaneae can be differentiated from B. lozanoi and B. nicefori

:FIgure 8 :
FIgure 8: Altitudinal distribution of species of the genus Bolitoglossa in the Cordillera Oriental of Colombia.

FIgure 9 :
FIgure 9: Distribution of Bolitoglossa lozanoi from Cordilleras Oriental and Central of Colombia, yellow dots correspond to localities, red dot is the type locality sensu Acosta and Restrepo, 2001; orange dot shows erroneously the distribution proposed by IUCN red List.

FIgure 10 :
FIgure 10: Distribution of Bolitoglossa palmata (southeastern of Cordillera Oriental).Red polygon corresponds the distribution proposed by IUCN red List.
: Adult female, ICN-MHN 54440 (field number ARA 4984), collected by Benjamin Tapley and Andrés Acosta, on 8 December 2008, deposited in the Amphibian collection, Institute of Natural Sciences, Universidad Nacional de Colombia.Paratypes: (9) ICN-MHN 34230, UIS-A 2203, UIS-A 2320, UIS-A 2324, UIS-A 2897 adult males from Municipality of Charalá, km 38 Hacienda La Sierra Santuario de Fauna y Flora Guanentá, Alto Río Fonce, vereda Santa Helena, Departamento of Santander Colombia, elevation 2400 m.Collected by Claudia Vélez, Jeannette Nieto, Doris Gutiérrez, Ruth Estupiñán, and Sandy Arroyo; MUJ 7193-4 adult males, Boyacá, ).It is a nocturnal and arboreal species related to primary or secondary forest, being collected perched on shrubs in open vegetation between 0.20-1 m meters above the ground.It is well known from the Municipality of Florencia in the Department of Caquetá.B. palmata occurs in the eastern flank in the southern part of Colombian Cordillera Oriental; it has been recorded in two subandean forest localities on the border between the Department of Caquetá and Huila, distributed in elevations between 2020-2360 m.Basados en las colecciones nacionales de referencia, ocho especies del género Bolitoglossa son reconocidas y una nueva especie Bolitoglossa guaneae sp.nov.es descrita para la Cordillera oriental de Colombia.En términos relativos a la distribución altitudinal la mayor riqueza en la cordillera Oriental se centra en las tierras medias asociadas a los bosques de niebla del flanco occidental.