ABSTRACT The Angolan Adder, Bitis heraldica (Bocage, 1889), is endemic to the Angolan central plateau and is one of the most poorly known and rarely observed species of African snakes. The phylogenetic placement of B. heraldica within the four subgenera of the genus Bitis (Bitis, Macrocerastes, Calechidna, Keniabitis) has been problematic. The few recent taxonomic and phylogenetic revisions dealing with African viperids and the genus Bitis have not included this species, resulting in the default acceptance of B. heraldica as member of the subgenus Calechidna, based on similar morphology and size. During a survey in Huambo Province, central Angola, we collected the first specimen of this species in the past 65 years, allowing us to examine its phylogenetic relationships to other Bitis using multigene Bayesian and maximum likelihood analyses of all species of Bitis, except B. harenna. Contrary to expectations that B. heraldica is a member of the subgenus Calechidna, our results unambiguously place B. heraldica among members of the subgenus Macrocerastes, which contains the largest-bodied species of the genus. The placement of B. heraldica within Macrocerastes raises interesting questions about body-size evolution in viperids and contributes to a better understanding of biogeographic patterns in south-western Africa, and particularly Angola.

中文翻译：

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巨人中的侏儒：难以捉摸的安哥拉加法鱼（Bitis heraldica）的系统发育位置和安哥拉非洲山地地区的生物地理学亲缘关系

摘要 安哥拉毒蛇，Bitis heraldica (Bocage, 1889)，是安哥拉中部高原的特有种，是最鲜为人知和很少观察到的非洲蛇种之一。B. heraldica 在 Bitis 属的四个亚属（Bitis、Macrocerastes、Calechidna、Keniabitis）中的系统发育位置一直存在问题。最近针对非洲蝰蛇和蝰蛇属的少数分类学和系统发育修订未包括该物种，导致基于相似的形态和大小，默认接受 B. heraldica 作为 Calechidna 亚属的成员。在安哥拉中部万博省的一次调查中，我们收集了过去 65 年来该物种的第一个标本，允许我们使用多基因贝叶斯和所有 Bitis 物种的最大似然分析来检查其与其他 Bitis 的系统发育关系，B. harenna 除外。与 B. heraldica 是 Calechidna 亚属成员的预期相反，我们的结果明确地将 B. heraldica 置于 Macrocerastes 亚属的成员中，后者包含该属中体型最大的物种。B. heraldica 在 Macrocerastes 中的放置提出了有关蝰蛇体型进化的有趣问题，并有助于更好地了解非洲西南部，尤其是安哥拉的生物地理模式。其中包含该属中体型最大的物种。B. heraldica 在 Macrocerastes 中的放置提出了有关蝰蛇体型进化的有趣问题，并有助于更好地了解非洲西南部，尤其是安哥拉的生物地理模式。其中包含该属中体型最大的物种。B. heraldica 在 Macrocerastes 中的放置提出了有关蝰蛇体型进化的有趣问题，并有助于更好地了解非洲西南部，尤其是安哥拉的生物地理模式。