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.

新热带区蛙类的角蛙科（Ceratophryidae）由躯体粗壮、口部宽阔的蛙类组成。该科现生类群始终被系统发育分析恢复为单系群，但不同研究在其内部系统发育关系上存在分歧。角蛙科拥有无尾目（Anura）中最为丰富的化石记录之一。然而，同时涵盖现生与灭绝类群的系统发育分析仍较为匮乏，且化石类群的系统发育位置始终存在争议。

有鉴于此，在过去十年间，该科的分类系统发生了显著变化——原归入该科的鲍鲁巴蟾（Baurubatrachus pricei，晚白垩世）、魔鬼蟾（Beelzebufo ampinga，晚白垩世）以及瓦韦尔蛙（Wawelia gerholdi，早中新世）已被移出该科。

本文以基于形态学构建的角蛙科系统发育树为研究框架，涵盖11个现生种与10个化石种，以此探讨该科的演化历史与分类系统。本研究通过系统发育分析将鲍鲁巴蟾、魔鬼蟾与瓦韦尔蛙归入非角蛙科类群。

本研究恢复的角蛙科为单系群：丽口蟾属（Lepidobatrachus）与角蟾属（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蟾属内的C. cornuta类群与C. aurita类群。

在化石类群中，基于自有衍征（autapomorphies）的分析结果显示，澳洲丽口蟾（Lepidobatrachus australis）与萨甘角蟾（Ceratophrys sagani）为有效物种，而鲁斯康尼角蟾（Ceratophrys rusconii）是所有角蟾属物种的姊妹群。恩塞纳达角蟾（Ceratophrys ensenadensis）、阿梅吉诺角蟾（Ceratophrys ameghinorum）、金角蟾（Ceratophrys aurita）标本NHMUK PV OR18895/6以及萨甘角蟾均隶属于C. aurita类群。

此外，本文还探讨了角蛙科内的趋同演化（homoplasies）、分化时间估算，以及该科的倍性演化与背盾演化。