The latest advancements in DNA sequencing technologies have facilitated the resolution of the phylogeny of insects, yet parts of the tree of Holometabola remain unresolved. The phylogeny of Neuropterida has been extensively studied, but no strong consensus exists concerning the phylogenetic relationships within the order Neuroptera. Here, we assembled a novel transcriptomic dataset to address previously unresolved issues in the phylogeny of Neuropterida and to infer divergence times within the group. We tested the robustness of our phylogenetic estimates by comparing summary coalescent and concatenation-based phylogenetic approaches and by employing different quartet-based measures of phylogenomic incongruence, combined with data permutations. Our results suggest that the order Raphidioptera is sister to Neuroptera + Megaloptera. Coniopterygidae is inferred as sister to all remaining neuropteran families suggesting that larval cryptonephry could be a ground plan feature of Neuroptera. A clade that includes Nevrorthidae, Osmylidae, and Sisyridae (i.e. Osmyloidea) is inferred as sister to all other Neuroptera except Coniopterygidae, and Dilaridae is placed as sister to all remaining neuropteran families. Ithonidae is inferred as the sister group of monophyletic Myrmeleontiformia. The phylogenetic affinities of Chrysopidae and Hemerobiidae were dependent on the data type analyzed, and quartet-based analyses showed only weak support for the placement of Hemerobiidae as sister to Ithonidae + Myrmeleontiformia. Our molecular dating analyses suggest that most families of Neuropterida started to diversify in the Jurassic and our ancestral character state reconstructions suggest a primarily terrestrial environment of the larvae of Neuropterida and Neuroptera. Our extensive phylogenomic analyses consolidate several key aspects in the backbone phylogeny of Neuropterida, such as the basal placement of Coniopterygidae within Neuroptera and the monophyly of Osmyloidea. Furthermore, they provide new insights into the timing of diversification of Neuropterida. Despite the vast amount of analyzed molecular data, we found that certain nodes in the tree of Neuroptera are not robustly resolved. Therefore, we emphasize the importance of integrating the results of morphological analyses with those of sequence-based phylogenomics. We also suggest that comparative analyses of genomic meta-characters should be incorporated into future phylogenomic studies of Neuropterida.

中文翻译：

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一种综合系统发育学方法，用于阐明神经翅目（昆虫纲：Holometabola）的进化历史和分化时间。


DNA测序技术的最新进展促进了昆虫系统发育的解决，但全变态树的部分内容仍未解决。脉翅目的系统发育已被广泛研究，但对于脉翅目内的系统发育关系还没有达成强有力的共识。在这里，我们组装了一个新的转录组数据集，以解决神经翅目系统发育中以前未解决的问题，并推断该群体内的分歧时间。我们通过比较基于合并和串联的系统发育方法，并采用不同的基于四重奏的系统发育不一致性测量方法，并结合数据排列，测试了系统发育估计的稳健性。我们的结果表明，Raphidioptera 目是脉翅目 + 大翅目的姐妹。 Coniopterygidae 被推断为所有剩余神经翅目家族的姐妹，这表明幼虫隐肾可能是神经翅目的地面特征。包括 Nevrorthidae、Osmylidae 和 Sisyridae（即 Osmyloidea）的进化枝被推断为除 Coniopterygidae 之外的所有其他脉翅目的姐妹，而 Dilaridae 则被视为所有剩余脉翅目科的姐妹。 Ithonidae 被推断为单系 Myrmeleontiformia 的姐妹群。 Chrysopidae 和 Hemerobiidae 的系统发育亲缘关系取决于分析的数据类型，基于四重奏的分析显示，对于将 Hemerobiidae 视为 Ithonidae + Myrmeleontiformia 的姐妹的支持微弱。我们的分子测年分析表明，大多数脉翅目科在侏罗纪开始多样化，我们的祖先特征状态重建表明脉翅目和脉翅目幼虫的主要陆地环境。 我们广泛的系统发育分析巩固了脉翅目主干系统发育的几个关键方面，例如脉翅目中锥翅目的基本位置和螟总科的单系。此外，它们还为神经翅目多样化的时机提供了新的见解。尽管分析了大量的分子数据，我们发现神经翅目树中的某些节点尚未得到可靠解析。因此，我们强调将形态学分析结果与基于序列的系统发育组学结果相结合的重要性。我们还建议将基因组元特征的比较分析纳入神经翅目未来的系统发育学研究中。