Brazilian marsupial frogs are diphyletic (Anura: Hemiphractidae: Gastrotheca)

Molecular phylogenetic analyses based on expanded taxonomic and geographic sampling support the monophyly of the marsupial frog genera (family Hemiphractidae), resolve six geographically circumscribed lineages within Gastrotheca, and, for the first time, reveal that two divergent lineages of Gastrotheca inhabit the Atlantic Coastal Forests of Brazil. Within Gastrotheca, the earliest diverging clade is confined to northeastern Brazil, whereas the three subsequent diverging lineages are restricted to northern Venezuela (G. walkeri), southeastern Brazil, and northwestern South America. All species in these clades inhabit humid forests at low to mid-elevations, and their life histories are characterized by lacking free-living tadpoles (i.e., direct development). Two derived clades inhabit the Andes, and both contain species with either direct development or tadpoles. One Andean clade of Gastrotheca ranges in the high Andes from Colombia to extreme northern Peru, whereas the other clade inhabits high elevations in the Andes of southern Ecuador, Peru, and Bolivia, and lower elevations in the Andes of northwestern Argentina. The presence of two non-sister lineages on each side of the Amazon Basin suggests that vicariance across this central region played an important role in diversification within Gastrotheca.

基于扩大分类群与地理采样范围的分子系统发育分析，支持囊蛙科（Hemiphractidae）囊蛙类群的单系性，解析出壶囊蛙属（Gastrotheca）内6个地理界定的支系，并首次揭示壶囊蛙属内存在两个分化支系，其栖息于巴西大西洋沿岸森林。在壶囊蛙属中，最早分化的支系仅分布于巴西东北部，后续分化的3个支系则分别局限于委内瑞拉北部（沃克壶囊蛙G. walkeri）、巴西东南部以及南美洲西北部。上述支系的所有物种均栖息于低至中海拔的湿润森林中，其生活史以无自由生活的蝌蚪为特征，即直接发育。另有两个衍生支系栖息于安第斯山脉，两类支系均包含具直接发育或蝌蚪阶段的物种。其中一支壶囊蛙支系的分布范围覆盖哥伦比亚至秘鲁极北部的安第斯高海拔区域，另一支则栖息于厄瓜多尔南部、秘鲁及玻利维亚的安第斯高海拔地带，以及阿根廷西北部安第斯山脉的低海拔区域。亚马逊盆地两侧各存在两个非姊妹支系，该现象表明该中部区域的地理分隔事件在壶囊蛙属的物种分化过程中发挥了重要作用。