Michael Weingardt

Understanding macroevolution requires integrating morphological and molecular data from fossil and extant taxa to reconstruct phylogenies, trait evolution and biogeographic history. However, the macroevolutionary process since its deep‐time origin via a total‐evidence approach has been investigated only in a limited number of groups. Here we present the first total‐evidence phylogenetic and biogeographic analyses of Dilaridae, a key transitional group of Neuroptera, integrating high‐resolution 3D morphological reconstructions of Cretaceous stem‐group taxa. We describe a new genus and four new species from the Kachin amber, that is, † Palaeonallachius fangyuani gen. et sp. nov., † Coronidilar jiaxiaoae sp. nov., † Cretodilar zhiqiae sp. nov. and † Cretodilar galaxias sp. nov., which provide valuable morphological data for phylogenetic inference. Comparative analyses using alternative sampling strategies (extant‐only or total‐evidence) demonstrate the advantages of integrating fossil data in phylogenetic inference, divergence‐time estimation and ancestral range reconstruction. Our results support a Gondwanan origin of Dilaridae associated with the West Burma Block, followed by dispersal into Laurasia via northward‐drifting Gondwanan landmasses and possible trans‐Antarctic dispersal since the Jurassic. The absence of extant dilarids in Australasia is likely due to extinction. These findings highlight the importance of fossil integration in resolving deep‐time evolution and the role of Gondwanan dynamics in shaping insect biogeography.

Five specimens of primary larvae of Ripiphoridae (Insecta: Coleoptera) are reported from one piece of Cretaceous Kachin amber. They represent two morphotypes: one conicocephalate and one belonging to the tribe Ripidiini (represented by four specimens). The conicocephalate morphotype is compared with similar larvae known from Kachin, Taimyr and Manitoba Cretaceous amber, and the larvae of Ripidiini are compared with their Cretaceous, Eocene and extant relatives. Phylogenetic analyses were performed to establish a working hypothesis about possible relationships of both lineages. The results, which conform with a recent molecular phylo­geny, indicate that the larvae described here belong to Ripidiini or are closely related, respectively. To allow taxonomic and systematic work with conicocephalate larvae from Kachin amber, a collective group name †“Ripilarva” nov. is proposed here to accommodate these immature stages. Both species of †“Ripilarva” nov. from Kachin amber are described: †“R.” parabolicasp. nov. and †“R.” kachinensissp. nov. Syninclusion of †“Ripilarva” kachinensissp. nov. and four larvae of a species of Ripidiini indicate that females of both ripiphorids chose the same time and microhabitat for oviposition in the Kachin Cretaceous forest. The results are further verified by the use of UV-photography, as the different larval morphotypes occurred in the same resin flow. The presence of larvae of Ripidiini in clusters contrary to the solitary occurrence of †“Ripilarva” nov. in Cretaceous amber of Russia, Canada, USA, and Myanmar is interpreted as a possible result of different oviposition strategies, with different numbers of eggs deposited at one spot. Graphical abstract

As the only direct records of the history of evolution, it is critical to determine the geological source of biota-bearing fossils. Through the application of synchrotron-radiation micro-computed tomography (SR-µ-CT), Fourier-transformed infrared-spectroscopy (FT-IR), visual evaluation of ultraviolet fluorescence (UV-VS), radiocarbon dating (¹⁴C quantification), and historical sleuthing, we were able to identify and sort 161 (83 Baltic amber, 71 Copal and 7 Kauri gum pieces) individually numbered and largely mislabeled pieces of East African Defaunation resin (~145 years old) and copal (~390 years old), as well as Baltic amber (~35 million years old) from the Phyletisches Museum collection. Based on this collection, we define two new species: ‡Amphientomum knorrei Weingardt, Bock & Boudinot, sp. nov. (Psocodea: Amphientomidae, copal) and †Baltistena nigrispinata Batelka, Tröger & Bock, sp. nov. (Coleoptera: Mordellidae, Baltic amber). For selected taxa, we provide systematic reviews of the fossil record, including: Amphientomidae, for which we provide a key to all species of Amphientomum, extant and extinct, and recognize the junior synonymy of Am. ectostriolatum Li, 2002 (an unjustified emendation) under Am. ectostriolate Li, 1999 (syn. nov.); the fossil ant genus †Yantaromyrmex and the clades Dorylinae, Plagiolepidini, Camponotus, Crematogaster, and Pheidole (Formicidae); the Nevrorthidae (Neuroptera); and Doliopygus (Coleoptera: Curculionidae: Platypodinae). We synonymize Palaeoseopsis Enderlein, 1925 with Amphientomum Pictet, 1854, syn. nov. and transfer one species from Amphientomum, forming Lithoseopsis indentatum (Turner, 1975), comb. nov. To prevent the uncritical usage of unidentifiable fossils attributed to Camponotus for macroevolutionary analysis, we transfer 29 species to the form genus †Camponotites Steinbach, 1967, which we consider to be most useful as incertae sedis in the Formicinae. We treat †Ctt. ullrichi (Bachmayer, 1960), comb. nov. as unidentifiable hence invalid stat. nov. We also transfer †Ca. mengei Mayr, 1868 and its junior synonym †Ca. igneus Mayr, 1868 to a new genus, †Eocamponotus Boudinot, gen. nov., which is incertae sedis in the Camponotini. Concluding our revision of Camponotus fossils, we transfer †Ca. palaeopterus (Zhang, 1989) to Liometopum (Dolichoderinae), resulting in †L. palaeopterumcomb. nov. and the junior synonymy of †Shanwangella Zhang, 1989, syn. nov. under Liometopum Mayr, 1861. Because the type specimens of the genera †Palaeosminthurus Pierce & Gibron, 1962, stat. rev. and †Pseudocamponotus Carpenter, 1930 are unidentifiable due to poor preservation, we consider these taxa unidentifiable hence invalid stat. nov. To avoid unsupported use of the available fossils names attributed to Crematogaster for divergence dating calibration points, we transfer three species to a new collective taxon that is incertae sedis in Myrmicinae, †Incertogaster Boudinot, gen. nov., forming †In. aurora (LaPolla & Greenwalt, 2015), †In. praecursor (Emery, 1891), comb. nov., and †In. primitiva (Radchenko & Dlussky, 2019), comb. nov. Finally, we transfer †Ph. cordata (Holl, 1829) back to Pheidole, and designate a neotype from our copal collection based on all available evidence. All new species plus the neotype of ‡Ph. cordata are depicted with 3D cybertypes from our µ-CT scan data. We introduce the convention of a double dagger symbol (‡) to indicate fossils in copal or Defaunation resin, as these may yet be extant. To further contextualize our results, we provide a discussion of amber history and classification, as well as the Kleinkuhren locality, to which multiple specimens were attributed. We conclude with conspecti on key biological problems and increasing potential of µ-CT for phylogenetic paleontology.