Saturday, August 3, 2019

[Herpetology • 2019] Systematics of Huicundomantis, A New Subgenus of Pristimantis (Anura, Strabomantidae) with Extraordinary Cryptic Diversity and Eleven New Species


Pristimantis (Huicundomantis) spp.


Páez & Ron, 2019

Abstract
Pristimantis is the most diverse genus of tetrapods comprising 532 described species. It contains a large number of morphologically cryptic species that are being discovered with the assistance of genetic evidence. We use molecular, morphological, bioacoustic, and environmental data to assess the phylogenetic relationships and determine the species within an Andean clade of Pristimantis, which is distributed from central Ecuador to northern Peru. We assign to this clade the name Huicundomantis and propose it as a subgenus. Our results show that Huicundomantis is composed of two large clades which we name as the P. phoxocephalus species group and the P. cryptomelas species group. Huicundomantis is composed of 28 species of which 12 have been described and 16 are new. We describe 11 of these undescribed species. The most effective characters to discriminate among species are DNA sequences, qualitative morphology, and advertisement calls. Morphometric and environmental characters are not very useful to define species limits. We clarify the identity of P. riveti and show that populations from southern Ecuador traditionally ascribed to P. riveti are a new species, P. lutzae sp. nov. We also show that P. prometeii is a junior synonym of P. hampatusami. The current diversity and geographic distribution of Huicundomantis are consistent with a model of allopatric speciation. All species have a restricted distribution range (less than 4330 km2) and are assigned to the Red List categories Data Deficient or threatened with extinction. We provide new reasons to increase conservation efforts for these species and their habitat. Taking our results into account, Pristimantis species richness in Ecuador increases from 211 to 221 species, and the number of species endemic to Ecuador from 119 to 129.

Keywords: Andes, cryptic diversity, integrative taxonomy, Neotropics, Pristimantis phoxocephalus species group, P. cryptomelas species group



Figure 4. Morphological variation within Pristimantis, subgenus Huicundomantis.
A Pristimantis philipi. Pristimantis cryptomelas group (*): B Pristimantis gagliardoi C Pristimantis muscosus D Pristimantis cryptomelas. Pristimantis phoxocephalus group (**): E Pristimantis torresi sp. nov. F Pristimantis hampatusami G Pristimantis jimenezi sp. nov. H Pristimantis phoxocephalus I Pristimantis totoroi sp. nov. J Pristimantis atratus K Pristimantis chomskyi sp. nov. L Pristimantis multicolor sp. nov. M Pristimantis sp. (CCS2) N Pristimantis gloria sp. nov. O Pristimantis lutzae sp. nov. P Pristimantis tinguichaca Q Pristimantis verrucolatus sp. nov. R Pristimantis sp. (UCS1) S Pristimantis teslai sp. nov. T Pristimantis versicolor U Pristimantis atillo sp. nov. Red branches are for clades without available photographs of live individuals. Not shown at the same scale.

Huicundomantis subgenus nov.

Type species: Pristimantis phoxocephalus (Lynch, 1979).

Definition: This clade is strongly supported by genetic evidence (Fig. 3). Morphological synapomorphies are unknown. Members of this clade are characterized by: (i) dorsolateral folds absent (except for P. atratus); (ii) cranial crests absent (except for P. atratus, P. percultus, and P. spinosus); (iii) tympanic membrane and tympanic annulus prominent (both absent in P. philipi); (iv) dentigerous processes of vomer present; (v) small to prominent tubercles on heel; (vi) fingers with lateral fringes; (vii) basal webbing between toes (except for P. balionotus, P. hampatusami, and P. philipi which lack webbing); (viii) Toe V longer or much longer than Toe III (Figs 7–9); (ix) in life, groins and concealed surfaces of thighs with distinctive coloration patterns including flash colors and light or bright colored flecks or spots on a darker background; colors, shapes and sizes of these ornaments are variable among species; (x) SVL females 24.9–46.8 mm; SVL males 16.1–34.5 mm. Species of this clade may bear a fleshy keel or a papilla at the tip of the snout, and lateral, middorsal, or postocular folds.

Content: This clade comprises 23 described species (11 of them described below): P. atillo sp. nov., P. atratus, P. balionotus, P. chomskyi sp. nov., P. cryptomelas, P. gagliardoi, P. gloria sp. nov., P. hampatusami, P. jimenezi sp. nov., P. lutzae sp. nov., P. multicolor sp. nov., P. muscosus, P. nangaritza sp. nov., P. percultus, P. philipi, P. phoxocephalus, P. spinosus, P. teslai sp. nov., P. tinguichaca, P. torresi sp. nov., P. totoroi sp. nov., P. versicolor, P. verrucolatus sp. nov. This clade encompasses the P. phoxocephalus and P. cryptomelas species groups.

Distribution: Huicundomantis occurs in Eastern and Western Andean slopes and Inter-Andean valleys of southern and central Ecuador, and Eastern Andean slopes of northern Peru. They inhabit the following Natural Regions: Deciduous Costa Forest, Western Foothill Forest, Western Montane Forest, Paramo, Inter-Andean Shrub, Eastern Montane Forest, and Eastern Foothill Forest, between elevations of 230 and 4200 m a.s.l.

Etymology: We name this clade Huicundomantis because these frogs are frequently found inside bromeliad plants. Huicundo is a word in Quechua, an indigenous South American language, locally used to referring to bromeliads.

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 Nadia B. Páez and Santiago R. Ron. 2019. Systematics of Huicundomantis, A New Subgenus of Pristimantis (Anura, Strabomantidae) with Extraordinary Cryptic Diversity and Eleven New Species. ZooKeys. 868: 1-112. DOI: 10.3897/zookeys.868.26766

Resumen: Pristimantis es el género más diverso de tetrápodos, contando con 532 especies descritas. Contiene un gran número de especies morfológicamente crípticas que están siendo descubiertas con el uso de evidencia genética. En el presente estudio usamos análisis integrativos, incluyendo información molecular, morfológica, bioacústica y ambiental para determinar el contenido de especies de un clado andino de Pristimantis que se distribuye desde el centro del Ecuador hasta el norte de Perú. Asignamos a este clado el nombre de Huicundomantis y lo proponemos con el rango de subgénero. Nuestros resultados indican que Huicundomantis está compuesto por dos grandes clados que nombramos como los grupos de especies P. phoxocephalus y P. cryptomelas. Huicundomantis contiene 28 especies de las cuales 12 están descritas y 16 son nuevas. En este estudio describimos 11 de las especies nuevas. Los caracteres más efectivos para discriminar entre especies de Huicundomantis son secuencias de ADN, morfología cualitativa y cantos de anuncio. Diferencias morfométricas y ambientales entre especies son de poca utilidad para delimitar especies. En este estudio también clarificamos la identidad de P. riveti y determinamos que poblaciones del sur del Ecuador, tradicionalmente consideradas P. riveti, corresponden a la nueva especie P. lutzae sp. nov. Además, reportamos a P. prometeii como sinónimo junior de P. hampatusami. La diversidad y distribución geográfica de Huicundomantis son consistentes con un modelo de especiación alopatrica. Todas las especies tienen un rango de distribución restringido (menos de 4330 km2) y son asignadas a las categorías de Lista Roja de Datos Insuficientes o amenazadas de extinción. Nuestros resultados son un nuevo argumento para aumentar los esfuerzos de conservación de estas especies y su hábitat. Tomando en cuenta nuestros resultados, la riqueza de especies de Pristimantis en el Ecuador aumenta de 211 a 221 especies y su número de especies endémicas, de 119 a 129.

Eleven new species of rain frogs discovered in the tropical Andes phys.org/news/2019-08-eleven-species-frogs-tropical-andes.html via @physorg_com