Pentatomidae is the third largest family of true bugs, comprising over 40 tribes. Few tribes have been studied in a phylogenetic context, and none of them have been examined using molecular data. Moreover, little is known about the evolution of key morphological characters widely used in taxonomic and phylogenetic studies at multiple levels. Here, we conduct a phylogenetic study of the tribe Chlorocorini (Pentatominae) combining 69 morphological characters and five DNA loci. We use the inferred phylogeny to reconstruct the evolution of key morphological characters such as the spined humeral angles of the pronotum, a dorsal projection on the apices of the femora and characters of male genitalia. We provide solid evidence that the tribe as currently recognized is not monophyletic based both on DNA and morphological data. The genera Arvelius Spinola and Eludocoris Thomas were consistently placed outside of the Chlorocorini, while the remaining genera were found to form a monophyletic group. We also show that nearly all morphological diagnostic characters for the tribe are homoplastic. The only exception is the development of the hypandrium, which, contrary to expectations for genital traits, showed the slowest evolutionary rates. In contrast, the most rapidly evolving trait is the length of the ostiolar ruga, which may be attributed to selection favoring anti-predatory behavior and other functions of its associated scent glands. Lastly, we also provide a preliminary glimpse of the main phylogenetic relationships within the Pentatomidae, which indicates that most of the included subfamilies and tribes are not monophyletic. Our results suggest that the current subfamily-level classification of Pentatomidae is not adequate to reflect its evolutionary history, and we urge for a more complete phylogeny of the family.

中文翻译：

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基于DNA和形态学数据的臭虫部落绿藻（Cheterocorini）的系统发育：主要表型性状的演变

昆虫科是真正的虫子的第三大家族，包括40多个部落。很少有人在系统发育的背景下研究过部落，而且都没有使用分子数据对其进行过检查。此外，对于在多个层次上的分类学和系统发育研究中广泛使用的关键形态特征的演变所知甚少。在这里，我们对绿藻（Pentatominae）部落进行了系统发育研究，结合了69个形态特征和5个DNA位点。我们使用推断的系统发育来重建关键形态特征的演变，例如前胸膜的肱骨角旋转，股骨顶部的背侧投影和男性生殖器特征。我们提供有力的证据，根据DNA和形态学数据，当前公认的部落不是单系的。Arvelius Spinola和Eludocoris Thomas属始终位于绿藻属之外，而其余属被认为是一个单系群体。我们还表明，该部落几乎所有的形态学诊断特征都是同质的。唯一的例外是玻璃纤维的发育，与对生殖器性状的期望相反，其进化速度最慢。相反，发展最快的性状是骨小窝的长度，这可能归因于有利于抗掠食行为及其相关气味腺的其他功能的选择。最后，我们还初步了解了五翅目科内的主要系统发育关系，这表明所包括的大多数亚科和部落都不是单系统的。