Chalcidoidea are a megadiverse group of mostly parasitoid wasps of major ecological and economical importance that are omnipresent in almost all extant terrestrial habitats. The timing and pattern of chalcidoid diversification is so far poorly understood and has left many important questions on the evolutionary history of Chalcidoidea unanswered. In this study, we infer the early divergence events within Chalcidoidea and address the question of whether or not ancestral chalcidoids were small egg parasitoids. We also trace the evolution of some key traits: jumping ability, development of enlarged hind femora, and associations with figs. Our phylogenetic inference is based on the analysis of 3,239 single-copy genes across 48 chalcidoid wasps and outgroups representatives. We applied an innovative a posteriori evaluation approach to molecular clock-dating based on nine carefully validated fossils, resulting in the first molecular clock-based estimation of deep Chalcidoidea divergence times. Our results suggest a late Jurassic origin of Chalcidoidea, with a first divergence of morphologically and biologically distinct groups in the early to mid Cretaceous, between 129 and 81 million years ago (mya). Diversification of most extant lineages happened rapidly after the Cretaceous in the early Paleogene, between 75 and 53 mya. The inferred Chalcidoidea tree suggests a transition from ancestral minute egg parasitoids to larger-bodied parasitoids of other host stages during the early history of chalcidoid evolution. The ability to jump evolved independently at least three times, namely in Eupelmidae, Encyrtidae, and Tanaostigmatidae. Furthermore, the large-bodied strongly sclerotized species with enlarged hind femora in Chalcididae and Leucospidae are not closely related. Finally, the close association of some chalcidoid wasps with figs, either as pollinators, or as inquilines/gallers or as parasitoids, likely evolved at least twice independently: in the Eocene, giving rise to fig pollinators, and in the Oligocene or Miocene, resulting in non-pollinating fig-wasps, including gallers and parasitoids. The origins of very speciose lineages (e.g., Mymaridae, Eulophidae, Pteromalinae) are evenly spread across the period of chalcidoid evolution from early Cretaceous to the late Eocene. Several shifts in biology and morphology (e.g., in host exploitation, body shape and size, life history), each followed by rapid radiations, have likely enabled the evolutionary success of Chalcidoidea.

金缕梅属是一个具有多种多样的类群，主要是具有重要的生态和经济意义的寄生类黄蜂，几乎在所有现存的陆地生境中都普遍存在。迄今为止，人们对类葫芦科植物多样化的时机和方式了解甚少，这使得对类藻科动物进化史的许多重要问题没有得到解答。在这项研究中，我们推断金缕石属中的早期趋异事件，并解决祖先的金缕石类是否为小卵类寄生虫的问题。我们还追踪了一些关键特征的演变：跳跃能力，后股骨增大的发展以及与无花果的关联。我们的系统发育推论是基于对48个类拟南芥黄蜂和外群代表的3,239个单拷贝基因的分析得出的。我们应用了创新的后验一种基于九种经过仔细验证的化石的分子钟约会的评估方法，从而导致了首次基于分子钟的深金缕梅发散时间估算。我们的结果表明，晚侏罗世起源于金缕梅属，在129至8100万年前（mya）之间，在白垩纪的早期至中期，形态和生物学上的独特群体首次出现差异。在现今的白垩纪晚期，大多数现存世系的多样化迅速发生在75至53的Mya之间。推断的金缕梅树暗示了在金缕梅演化的早期历史中，从祖先的微小卵类寄生虫过渡到其他寄主阶段的大体寄生性寄生虫。跳跃的能力至少在Eupelmidae，Encyrtidae和Tanaostigmatidae中独立进化了至少三次。此外，大型的强硬核种，在后肢科和隐鳞科中具有较大的后股骨，并没有密切的关系。最后，一些类蜂类黄蜂与无花果的紧密联系，无论是作为传粉者，还是作为询问/居群动物或作为寄生类动物，都可能至少独立地进化了两次：在始新世，产生了无花果传粉者，在渐新世或中新世中，在无授粉的无花果黄蜂中，包括gall虫和寄生虫。在从白垩纪早期到始新世晚期的类鲨鱼类进化期间，非常特殊的谱系（如Mymaridae，Eulophidae，Pteromalinae）的起源均匀分布。生物学和形态上的几次变化（例如，宿主的发育，身体形状和大小，生活史）发生变化，每一次都伴随着快速的辐射，这很可能促使了金龟子的进化成功。