Systematics of the endangered toad genus Andinophryne (Anura: Bufonidae): phylogenetic position and synonymy under the genus Rhaebo

Abstract

Bufonidae is one of the most diverse amphibian families. Its large-scale phylogenetic relationships are relatively well understood with the exception of few Neotropical genera that may have diverged early in the evolution of the family. One of those genera is Andinophryne, a poorly known group of three toad species distributed in the western slopes of the Andes of northern Ecuador and southern Colombia. Their phylogenetic position is unknown due to lack of genetic data. We estimated a new phylogeny (over 200 species) of the family Bufonidae based on DNA sequences of mitochondrial and nuclear genes to assess the phylogenetic position of Andinophryne based on recently collected specimens of A. colomai and A. olallai from Ecuador and Colombia. We also examined external and internal morphology of Andinophryne to explore its congruence with the new phylogeny. The mtDNA and nuclear phylogenies show that Andinophryne is embedded within Rhaebo, a genus that belongs to a large clade characterized by the presence parotoid glands. Morphological characters confirmed the affinity of Andinophryne to Rhaebo and a close relationship between Andinophryne colomai and Andinophryne olallai. Rhaebo was paraphyletic relative to Andinophryne and to solve this problem we synonymize Andinophryne under Rhaebo. We discuss putative morphological synapomorphies for Rhaebo including Andinophryne. We provide species accounts for R. atelopoides new comb., R. colomai new comb. and R. olallai new comb. including assessments of their conservation status. We suggest that the three species are Critically Endangered. Their altitudinal distribution and association with streams are characteristic of endangered Andean amphibians.Key words: Bufonidae, Colombia, conservation status, Ecuador, Rhaebo, synonymy, systematics ResumenBufonidae es una de las familias de anfibios más diversas. Sus relaciones filogenéticas a gran escala están relativamente bien entendidas, con la excepción de algunos géneros Neotropicales que pueden haber divergido temprano en la evolución de la familia. Uno de esos géneros es Andinophryne, un grupo pobremente conocido de tres especies de sapos distribuidos en la vertiente occidental de los Andes en el norte de Ecuador y el sur de Colombia. Su posición filogenética es desconocida por falta de datos genéticos. Estimamos una nueva filogenia (casi 200 especies) de la familia Bufonidae basada en secuencias de ADN de genes mitocondriales y nucleares para evaluar la posición filogenética de Andinophryne utilizando especímenes recientemente colectados de A. colomai y A. olallai en Ecuador y Colombia. También examinamos la morfología externa e interna de Andinophryne para explorar su congruencia con la nueva filogenia. Las filogenias con los ADN mitocondriales y nucleares mostraron que Andinophryne está anidado dentro de Rhaebo, un género que pertenece a un clado grande caracterizado por la presencia de glándulas parotoideas. Los caracteres morfológicos confirmaron la afini-348 · Zootaxa 3947 (3) © 2015 Magnolia Press dad de Andi...

Rediscovery of the Endangered Carchi Andean Toad, Rhaebo colomai (Hoogmoed, 1985), in Ecuador, with comments on its conservation status and extinction risk

“…Identification. All specimens of R. colomai reported herein match descriptions of this species by Hoogmoed (1985Hoogmoed ( , 1989 and Ron et al (2015): flat head, projected snout, upper eyelid margin with sharp rim, elongated parotoids that produce yellow secretions, flanks with areolate skin and oblique and continuous row of small warts, long and slender limbs, webbed fingers and toes, and absent enlarged cephalic crests. Females have smooth dorsal skin, small tubercles, and a larger body size (snout-vent length 54.9-59.1 mm) while males have spiculated tubercular dorsal skin and a smaller body size (32.6-38.4 mm).…”

“…We evaluated the extinction risk according to the IUCN (2012) guidelines. Area of (Hoogmoed 1985, 1989, Ron et al 2015. Circles represent the new records reported herein.…”

“…Thus, this species was only known from 2 localities at the River Baboso and Chical, both located on the Andean slopes of northern Ecuador, about 30 km apart in a straight line. This species remained known only from Ecuador until Ron et al (2015) reported 6 males and 3 females from a population from the Reserva Natural Río Ñambi, department of Nariño, Colombia. This population had been overlooked because it was previously reported as Rhaebo olallai (Hoogmoed, 1985) by Murillo-Pacheco et al (2005).…”

Characterization of the mitochondrial genomes of two toads, Anaxyrus americanus (Anura: Bufonidae) and Bufotes pewzowi (Anura: Bufonidae), with phylogenetic and selection pressure analyses

“…Frost et al (2006) andVan Bocxlaer et al (2009) corroborated the monophyly of Bufonidae as well as the results of Portik & Papenfuss (2015). Our phylogeny within Bufonidae is generally similar to the results of Pramuk et al (2008), Ron et al (2015) and Dong & Yang (2016). The ML and BI analyses also confirmed the monophyly of Bufo, which has also been demonstrated by other researchers (Graybeal, 1997;Dong & Yang, 2016).…”

Biological Diversity in Headwater Streams

“…The toad family, Bufonidae, has nearly 600 species, most of which occur in the tropics. Many clades in the bufonid phylogeny are derived from the evolutionary radiation of headwater lineages (e.g., Atelopus, Rhaebo and Rhinella) [92], and this has occurred repeatedly on several continents. The caddisfly family Limnephilidae is proposed to have diversified from cool stream ecosystems [78], as have several other lineages of freshwater invertebrates, and likely other taxa.…”