Calaphidinae is the second-largest subfamily in the family Aphididae. Despite their species diversity and some taxonomic controversy, no phylogenetic studies have been conducted on them thus far. Herein, we report the first molecular phylogeny of Calaphidinae and two related lineages, Phyllaphidinae and Saltusaphidinae, based on five genes (3418 bp) for 126 taxa. Maximum parsimony, maximum-likelihood and Bayesian inference phylogenetic analyses were performed on the multilocus dataset. Divergence time estimation, biogeographical reconstruction, ancestral host plant reconstruction and PhyloType analyses were performed to identify evolutionary trends in Calaphidinae. Our phylogenetic results lead to several conclusions: Phyllaphidinae is a sister group to Calaphidinae s.l.; Calaphidinae is paraphyletic with respect to the former “Saltusaphidinae”; the ingroup clade was subdivided into nine newly recognized lineages; and three subtribes of Calaphidinae (Monaphidina, Calaphdina and Panaphidina) and many genera were not recovered as monophyletic. A new classification is proposed with eight tribal divisions that reflect our phylogenetic results, including three new tribes (Pterocallidini trib.n., Pseudochromaphidini trib.n. and Shivaphidini trib.n.) and three new statuses (Saltusaphidini stat.n., Therioaphidini stat.n. and Myzocallidini stat.n.). The ancestral reconstruction results imply that the ingroup taxa’s common ancestor originated in the Eastern Palaearctic and might have fed on Fagaceae in the Late Cretaceous. Later, multiple host shifts and an expanding geographical distribution led to the current species diversity of Calaphidinae. Our reconstructions suggest that species diversification cannot solely be explained by speciation via host shifts and that geographical isolation probably also played a key role. Our results provide new insight into the natural classification and history of the host plant associations and biogeography of Calaphidinae s.l.

中文翻译：

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Calaphidinae（半翅目：蚜科）的分子系统发育和进化

Calaphidinae是蚜虫科的第二大亚科。尽管它们的物种多样性和一些分类争议，但迄今为止尚未对它们进行系统发育研究。在这里，我们报告了 Calaphidinae 和两个相关谱系 Phyllaphidinae 和 Saltusaphidinae 的第一个分子系统发育，基于 126 个分类群的五个基因 (3418 bp)。对多位点数据集进行最大简约、最大似然和贝叶斯推理系统发育分析。进行发散时间估计、生物地理重建、祖先寄主植物重建和系统型分析，以确定 Calaphidinae 的进化趋势。我们的系统发育结果得出几个结论： Phyllaphidinae 是 Calaphidinae sl 的姐妹群；Calaphidinae 与以前的“Saltusaphidinae”是并系的；内群进化枝被细分为九个新确认的谱系；卡拉菲达亚科的三个亚族（单胞菌属、长臂猿属和松尾藻属）和许多属都没有作为单系恢复。提出了一个新的分类，其中包含八个反映我们系统发育结果的部落划分，包括三个新部落（Pterocallidinitrib.n ., Pseudochromaphidini trib.n。和 Shivaphidini trib.n .) 和三个新状态（Saltusaphidini stat.n .、Therioaphidini stat.n . 和 Myzocallidini stat.n.)。祖先重建结果表明，内群类群的共同祖先起源于古北界东部，可能在白垩纪晚期以镰刀科为食。后来，多次寄主转移和不断扩大的地理分布导致了目前的Calaphidinae物种多样性。我们的重建表明，物种多样化不能仅仅通过宿主转移的物种形成来解释，地理隔离可能也起到了关键作用。我们的研究结果为寄主植物协会的自然分类和历史以及 Calaphidinae sl 的生物地理学提供了新的见解