Phylogenetic analysis of Ceratophryidae (Anura: Hyloidea) including extant and extinct species

The Neotropical frog family Ceratophryidae is composed of wide-mouthed frogs with stout bodies. Living species of the family are consistently recovered as a monophyletic group, but with disparities among analyses regarding internal relationships. Ceratophryidae presents one of the richest fossil records in Anura. Nevertheless, phylogenetic analyses including both extant and extinct species are still scarce, and the position of fossils is persistently debated. In this sense, the systematics of the family has changed considerably in the last decade with the exclusion of <i>Baurubatrachus pricei</i> (Late Cretaceous), <i>Beelzebufo ampinga</i> (Late Cretaceous) and <i>Wawelia gerholdi</i> (early Miocene). Herein, a morphologically based phylogeny for Ceratophryidae, including living species (11 spp.) and fossil specimens (10 spp.), is used as a background to discuss the evolutionary history of the family and its classification. We phylogenetically placed <i>Baurubatrachus pricei</i>, <i>Beelzebufo ampinga</i> and <i>Wawelia gerholdi</i> as non-ceratophryids. We recovered a monophyletic Ceratophryidae: <i>Lepidobatrachus</i> and <i>Ceratophrys</i> form a clade, with <i>Chacophrys</i> as its sister group. Our analysis corroborates the <i>C. cornuta</i> and <i>C. aurita</i> groups. Among fossils, <i>L. australis</i> and <i>C. sagani</i> were recovered as valid species based on autapomorphies, and <i>C. rusconii</i> was found to be the sister of all <i>Ceratophrys</i>. <i>Ceratophrys ensenadensis</i>, <i>C. ameghinorum</i>, <i>C. aurita</i> NHMUK PV OR18895/6 and <i>C. sagani</i> belong to the <i>C. aurita</i> group. We also discuss homoplasies in Ceratophryidae, divergence-time estimates, and the evolution of ploidy and a dorsal shield in the family.

新热带区蛙类角花蟾科（Ceratophryidae）是一类口部宽阔、躯体粗壮的蛙类类群。该科的现生物种始终被恢复为单系群，但不同系统发育分析对其内部亲缘关系的推断存在分歧。角花蟾科拥有无尾目（Anura）中最为丰富的化石记录之一。然而，同时涵盖现生与灭绝物种的系统发育分析依然较为匮乏，且化石类群的系统发育位置始终存在争议。正因如此，该科的分类系统在近十年间发生了显著变动：晚白垩世的巴鲁巴蟾（Baurubatrachus pricei）、晚白垩世的魔鬼蟾（Beelzebufo ampinga）以及早中新世的瓦韦蟾（Wawelia gerholdi）均被移出该科。本研究以涵盖11个现生物种与10个化石物种的形态学系统发育框架为基础，探讨角花蟾科的演化历史与分类体系。本研究通过系统发育分析将巴鲁巴蟾、魔鬼蟾与瓦韦蟾恢复为非角花蟾科类群。本研究确认角花蟾科为单系群：细趾蟾属（Lepidobatrachus）与角花蟾属（Ceratophrys）构成一支演化支，查科蟾属（Chacophrys）为其姊妹群。本研究的分析结果支持角花蟾属内的C. cornuta组与C. aurita组两个类群。针对化石类群的分析显示，南方细趾蟾（Lepidobatrachus australis）与萨甘角花蟾（Ceratophrys sagani）基于独有衍征被确定为有效物种，而鲁斯康尼角花蟾（Ceratophrys rusconii）被恢复为所有角花蟾属物种的姊妹群。恩塞纳达角花蟾（Ceratophrys ensenadensis）、阿梅吉诺角花蟾（Ceratophrys ameghinorum）、编号为NHMUK PV OR18895/6的C. aurita标本以及萨甘角花蟾均隶属于C. aurita组。本研究同时探讨了角花蟾科的趋同演化特征、分化时间估算，以及该科的倍性演化与背甲演化。