SALAMANDRA 52(4) 283–292 30 DecemberThree new 2016 PsychrophrynellaISSN 0036–3375 from Bolivia Three new Bolivian species of Psychrophrynella (Anura: Craugastoridae), and comments on the amphibian fauna of the Cordillera de Apolobamba Ignacio De la Riva1 & James Aparicio2 1)Museo Nacional de Ciencias Naturales, CSIC, C/ José Gutiérrez Abascal, 28006 Madrid, Spain 2)Museo de Historia Natural-Colección Boliviana de Fauna, Casilla 8706, La Paz, Bolivia Corresponding author: Ignacio De la Riva, e-mail: [email protected] Manuscript received: 2 November 2015 Accepted: 18 July 2016 by Edgar Lehr Abstract. Three new species ofPsychrophrynella from the Bolivian section of the Cordillera de Apolobamba are described. The new species are distinguished from their closest relatives mainly by characters such as colour pattern, size, and skin texture. With the addition of these three new species, the diversity of the genus Psychrophrynella in Bolivia increases to 21 species. The protected Area Natural de Manejo Integrado Nacional (ANMIN) Apolobamba holds seven endemic species, and it is highly likely that more undescribed forms will be discovered when new surveys are conducted in this region, un- derscoring the need to preserve its rich endemic amphibian fauna. Key words. Amphibia, Andes, endemism, new species, Psychrophrynella. Introduction two more are currently being described (De la Riva et al. unpubl. data). However, the large expanse of suitable ar- Direct-developing frogs from the northern and central An- eas that have remained unexplored, and the known high des that live in high-altitude cloud forests, elfin forests, and endemism within the genus, make it likely that many un- humid highland grasslands share a suite of morphological described species of Psychrophrynella occur in the region. characteristics such as small to moderate adult size, short In this paper, we describe three additional Bolivian spe- legs, robust body, and tips of digits not or scarcely expand- cies of Psychrophrynella found during faunistic surveys in ed. Lynch (1975) grouped most of these forms distributed the protected Area Natural de Manejo Integrado Nacional from northern Colombia to central Bolivia in the genus (ANMIN) Apolobamba. At the same time, we discuss the Phrynopus Peters, 1863. However, Hedges et al. (2008) conservation value of the rich, highly endemic amphibian and Padial et al. (2012, 2014) demonstrated the non- fauna of this region. monophyly of Phrynopus sensu Lynch (1975). These dis- tinct Andean lineages have independently evolved similar morphologies, and their members are currently classified Material and methods in several genera. Phrynopus is now restricted to central Peru, whereas all the species occurring in Bolivia were as- Specimens were fixed in 10% formalin and preserved in signed to the genus Psychrophrynella Hedges, Duellman 70% ethanol. They were deposited at the Museo Nacional & Heinicke, 2008, where they represent an important pro- de Historia Natural, Colección Boliviana de Fauna (CBF, portion of the Andean anuran diversity. Eighteen species La Paz, Bolivia). We considered nine morphometric varia- of Psychrophrynella, all of them endemic, are known thus bles as follows: snout–vent length (SVL), head length (HL, far from Bolivia, and many more await description (De la from rictus to tip of snout), head width (HW, at level of ric- Riva 2007, De la Riva & Burrowes 2014). The diversi- tus), internarial distance (IND), distance from eye to nos- ty of these frogs seems to be especially high in certain re- tril (END), eye diameter (ED), tympanum diameter (TD), gions, among them the Cordillera de Apolobamba, which tibia length (TL), and foot length (FL, from proximal border extends across de Bolivian-Peruvian border. So far, four of inner metatarsal tubercle to tip of fourth toe). Lengths species have been described from the Bolivian section of of digits were compared by adpressing them against each Apolobamba (De la Riva 2007), while only one is known other. For measurements, we used a digital calliper to the from the Peruvian section (P. boettgeri [Lehr, 2006]), and nearest 0.01 mm, but rounded results to only one decimal © 2016 Deutsche Gesellschaft für Herpetologie und Terrarienkunde e.V. (DGHT), Mannheim, Germany All articles available online at http://www.salamandra-journal.com 283 Ignacio De la Riva & James Aparicio to avoid pseudo-precision (Hayek et al. 2001). Character Diagnosis: 1) medium size (maximum SVL 22.2 mm), body definitions follow Duellman & Lehr (2009), and diagno- moderately robust, legs moderately short (TL + FL = 78.1% sis and descriptions the format suggested by De la Riva SVL; range, 74.5–81.9%; N = 8); 2) tympanic membrane and (2007). When sex could not be identified by external char- tympanic annulus present and clearly visible; 3) first finger acters (e.g., size, sexual dimorphism in colour pattern, eggs slightly shorter than second; 4) tips of digits slightly swollen, observable through the ventral skin, presence of vocal sac in not expanded laterally; 5) no webbing of toes or lateral fring- adult males, etc.), gonads were examined. Descriptions of es; 6) two metatarsal tubercles, tarsal fold absent; 7) dorsal colour in life were based on field notes. Drawings were made skin and flanks shagreen with scattered small warts; no dor- under a stereomicroscope with the aid of a camera lucida. solateral folds; ventral skin finely granular; 8) snout round- Species of Psychrophrynella are highly endemic, with ed in dorsal view and in profile; 9) dorsum mostly uniform- their distributions being often extremely restricted; thus, ly brown; 10) venter greyish brown, chest and throat tan. they are well characterized by geography, which always The new species is assigned to the genus Psychrophrynel­ strongly supports species separation (De la Riva 2007). la as defined by Hedges et al. (2008) and Duellman & Two species separated by, say, 200 km, can be remarkably Lehr (2009). Frogs of the allied genus Bryophryne are similar in morphology and coloration, while species from morphologically similar to Psychrophrynella, but Bryo­ adjacent valleys are usually easy to distinguish by external phryne does not occur in Bolivia, being restricted to the characteristics. In taxonomy, diagnoses are provided most- Department of Cusco in southern Peru. Psychrophrynella ly for practical purposes, to facilitate telling apart (in the chaupi is superficially similar to P. katantika (De la Riva field or lab) closely related species, externally similar spe- & Martínez-Solano, 2007), which occurs in the nearby cies, or species living in the same region. For this reason, valley of Pelechuco (air-line distance between the two type and given the highly endemic distributions of species of localities, 11.5 km). The new species is distinguished from Psychrophrynella, we herein emphasize diagnostic compar- P. katantika mainly by being smaller (maximum SVL in isons between the geographically closest species (i.e., those P. chaupi 22.2 mm, 27.7 mm in P. katantika) and having a occurring in the Apolobamba range) and from those par- visible tympanum. Other geographically close species are ticularly similar ones, if any. P. colla sp. n. and P. melanocheira sp. n., described here- in, whose type localities are at air-line distances of 12.2 km and 18.5 km, respectively, to the southeast of the type local- Nomenclatural acts ity of P. chaupi. Psychrophrynella chaupi differs from these two species by lacking dorsolateral folds, having a finely The electronic edition of this article conforms to the require- granular ventral skin (smooth in P. colla and P. melano­ ments of the amended International Code of Zoological cheira), and a uniformly greyish brown ventral colora- Nomenclature, and hence the new names contained herein tion (beige with minute white flecks in P. colla; greenish are available under that Code from the electronic edition grey, sometimes with a black belly, in P. melanocheira). of this article. This published work and the nomenclatural The new species furthermore differs from P. colla by hav- acts it contains have been registered in ZooBank, the online ing slightly swollen tips of digits (slightly pointed in P. col­ registration system for the ICZN. The LSID (Life Science la), and from P. melanocheira by having longer legs. Oth- Identifier) for this publication is: urn:lsid:zoobank.org:pub: er, more geographically distant species of Psychrophrynella 1AADF552-DBAE-4765-A6E9-2B9673970FDA. The elec- that occur in the Cordillera de Apolobamba, are P. boett­ tronic edition of this work was published in a journal with geri (in the Peruvian section of the range), P. guillei (De an ISSN, has been archived and is available from the follow- la Riva, 2007), P. kalla waya (De la Riva & Martínez- ing digital repositories: www.salamandra-journal.com. Solano, 2007), and P. saltator (De la Riva, Reichle & Bosch, 2007). Psychrophrynella boettgeri is smaller (maxi- mum SVL 18.4 mm) than P. chaupi (22.2 mm) and has red- Psychrophrynella chaupi De la Riva & Aparicio, sp. n. dish brown blotches on the dorsum and orange blotches (Figs 1, 4C) on the groins and venter; P. guillei has an olive green dor- sum with dark blotches, and a yellowish white venter with ZooBank LSID: urn:lsid:zoobank.org:act: 0119034A-00FB-4AB7- black mottling; P. kallawaya is much larger (maximum A334-AB92F95C5DCC SVL 30.9 mm); and P. saltator has narrow cream bands on the dorsal faces of its digits and a large vocal sac in males. Holotype: CBF 5694, an adult female from Valle de