Insects are the most speciose group of animals, but the phylogenetic relationships of many major lineages remain unresolved. We inferred the phylogeny of insects from 1478 protein-coding genes. Phylogenomic analyses of nucleotide and amino acid sequences, with site-specific nucleotide or domain-specific amino acid substitution models, produced statistically robust and congruent results resolving previously controversial phylogenetic relations hips. We dated the origin of insects to the Early Ordovician [~479 million years ago (Ma)], of insect flight to the Early Devonian (~406 Ma), of major extant lineages to the Mississippian (~345 Ma), and the major diversification of holometabolous insects to the Early Cretaceous. Our phylogenomic study provides a comprehensive reliable scaffold for future comparative analyses of evolutionary innovations among insects.

Abstract.Here we present a phylogeny of beetles (Insecta: Coleoptera) based on DNA sequence data from eight nuclear genes, including six single-copy nuclear protein-coding genes, for 367 species representing 172 of 183 extant families. Our results refine existing knowledge of relationships among major groups of beetles. Strepsiptera was confirmed as sister to Coleoptera and each of the suborders of Coleoptera was recovered as monophyletic. Interrelationships among the suborders, namely Polyphaga (Adephaga (Archostemata, Myxophaga)), in our study differ from previous studies. Adephaga comprised two clades corresponding to Hydradephaga and Geadephaga. The series and superfamilies of Polyphaga were mostly monophyletic. The traditional Cucujoidea were recovered in three distantly related clades. Lymexyloidea was recovered within Tenebrionoidea. Several of the series and superfamilies of Polyphaga received moderate to maximal clade support in most analyses, for example Buprestoidea, Chrysomeloidea, Coccinelloidea, Cucujiformia, Curculionoidea, Dascilloidea, Elateroidea, Histeroidea and Hydrophiloidea. However, many of the relationships within Polyphaga lacked compatible resolution under maximum-likelihood and Bayesian inference, and/or lacked consistently strong nodal support. Overall, we recovered slightly younger estimated divergence times than previous studies for most groups of beetles. The ordinal split between Coleoptera and Strepsiptera was estimated to have occurred in the Early Permian. Crown Coleoptera appeared in the Late Permian, and only one or two lineages survived the end-Permian mass extinction, with stem group representatives of all four suborders appearing by the end of the Triassic. The basal split in Polyphaga was estimated to have occurred in the Triassic, with the stem groups of most series and superfamilies originating during the Triassic or Jurassic. Most extant families of beetles were estimated to have Cretaceous origins. Overall, Coleoptera experienced an increase in diversification rate compared to the rest of Neuropteroidea. Furthermore, 10 family-level clades, all in suborder Polyphaga, were identified as having experienced significant increases in diversification rate. These include most beetle species with phytophagous habits, but also several groups not typically or primarily associated with plants. Most of these groups originated in the Cretaceous, which is also when a majority of the most species-rich beetle families first appeared. An additional 12 clades showed evidence for significant decreases in diversification rate. These clades are species-poor in the Modern fauna, but collectively exhibit diverse trophic habits. The apparent success of beetles, as measured by species numbers, may result from their associations with widespread and diverse substrates -especially plants, but also including fungi, wood and leaf litter -but what facilitated these associations in the first place or has allowed these associations to flourish likely varies within and between lineages. Our results pr...

The order Coleoptera (beetles) is arguably the most speciose group of animals, but the evolutionary history of beetles, including the impacts of plant feeding (herbivory) on beetle diversification, remain poorly understood. We inferred the phylogeny of beetles using 4,818 genes for 146 species, estimated timing and rates of beetle diversification using 89 genes for 521 species representing all major lineages and traced the evolution of beetle genes enabling symbiont-independent digestion of lignocellulose using 154 genomes or transcriptomes. Phylogenomic analyses of these uniquely comprehensive datasets resolved previously controversial beetle relationships, dated the origin of Coleoptera to the Carboniferous, and supported the codiversification of beetles and angiosperms. Moreover, plant cell wall-degrading enzymes (PCWDEs) obtained from bacteria and fungi via horizontal gene transfers may have been key to the Mesozoic diversification of herbivorous beetles—remarkably, both major independent origins of specialized herbivory in beetles coincide with the first appearances of an arsenal of PCWDEs encoded in their genomes. Furthermore, corresponding (Jurassic) diversification rate increases suggest that these novel genes triggered adaptive radiations that resulted in nearly half of all living beetle species. We propose that PCWDEs enabled efficient digestion of plant tissues, including lignocellulose in cell walls, facilitating the evolution of uniquely specialized plant-feeding habits, such as leaf mining and stem and wood boring. Beetle diversity thus appears to have resulted from multiple factors, including low extinction rates over a long evolutionary history, codiversification with angiosperms, and adaptive radiations of specialized herbivorous beetles following convergent horizontal transfers of microbial genes encoding PCWDEs.

One hundred seven external and internal characters ofThis results in a distinctly simplified wing folding mechanism. The monophyly of Cucujiformia (؉ Bostrichoidea) larval and adult representatives of 28 genera of the cois supported by the presence of cryptonephric Malpileopteran suborders were analyzed cladistically. Four ghian tubules. Transformations of fore-and hind-wings, groups of Neuropterida were introduced as outgroup. reinforcement and simplification of the thoracic exoskel-The analysis yielded 18 trees with a minimum of 194 eton, and an efficient use of a distinctly reduced set steps (CI 0.691). All trees support the monophyly of all of thoracic muscles play an important role in the early four suborders and a branching pattern (Archostemata evolution of Coleoptera. Many different larval character ؉ (Adephaga ؉ (Myxophaga ؉ Polyphaga))). The prestransformations take place in the earlier Mesozoic within ence of elytra with meso-and metathoracic locking the suborders. ᭧ 2000 The Willi Hennig Society devices, the specific hind-wing folding, the close connection of exposed sclerites, the absence of the mera, the absence of eight thoracic muscles, the reduced abdominal

Hemipteroid insects (Paraneoptera), with over 10% of all known insect diversity, are a major component of terrestrial and aquatic ecosystems. Previous phylogenetic analyses have not consistently resolved the relationships among major hemipteroid lineages. We provide maximum likelihood-based phylogenomic analyses of a taxonomically comprehensive dataset comprising sequences of 2,395 single-copy, protein-coding genes for 193 samples of hemipteroid insects and outgroups. These analyses yield a well-supported phylogeny for hemipteroid insects. Monophyly of each of the three hemipteroid orders (Psocodea, Thysanoptera, and Hemiptera) is strongly supported, as are most relationships among suborders and families. Thysanoptera (thrips) is strongly supported as sister to Hemiptera. However, as in a recent large-scale analysis sampling all insect orders, trees from our data matrices support Psocodea (bark lice and parasitic lice) as the sister group to the holometabolous insects (those with complete metamorphosis). In contrast, four-cluster likelihood mapping of these data does not support this result. A molecular dating analysis using 23 fossil calibration points suggests hemipteroid insects began diversifying before the Carboniferous, over 365 million years ago. We also explore implications for understanding the timing of diversification, the evolution of morphological traits, and the evolution of mitochondrial genome organization. These results provide a phylogenetic framework for future studies of the group.

Abstract. Characters of the head of adephagan larvae were examined and analysed phylogenetically. A labrum which is completely fused to the clypeofrons and the presence of a closed prepharyngeal tube are autapomorphies of Adephaga. Partial reduction of the fossa maxillaris, cardo and stipes forming a functional unit, the immobilization of the lacinia, attachment of M. craniolacinialis to the lateral stipital wall, and loss of one stipitopalpal muscle, are considered autapomorphies of Adephaga excluding Gyrinidae. Complete reduction of the fossa maxillaris and the presence of M. craniostipitalis medialis are possible autapomorphies of Adephaga excluding Gyrinidae and Haliplidae. The presence of caudal tentorial arms, insertion of the galea on the mesal side of palpomere I, and absence of the lacinia are considered synapomorphies of Trachypachidae and Dytiscoidea (Noteridae, Amphizoidae, Hygrobiidae, Dytiscidae). The presence of a slender, elongated process of the head capsule, which articulates with a corresponding socket of the cardo, is a possible autapomorphy of Dytiscoidea. The sinuate frontal sutures, distinctly protruding prementum, shortened M. craniostipitalis medialis, and absence of M. submentopraementalis are considered autapomorphies of Geadephaga excluding Trachypachidae. The presence of a regular row of hairs along the anterior hypopharyngeal margin is a possible autapomorphy of Geadephaga excluding Trachypachidae and Rhysodidae. Improvement of the hypopharyngeal filter apparatus suggests the monophyly of Anisochaeta. Presence of a penicillum and partial reduction of the lacinia are possible autapomorphies of Anisochaeta excluding Omophronini. Larvae of Cychrini, Carabini, Nebriini and Notiophilini are characterized by a strongly developed, cone‐shaped hypodon. Postocular and cervical ridges, crosswise arrangement of antennal muscles, and a completely flattened hypopharynx are considered autapomorphies of Caraboidea Limbata.

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Abstract. Larvae of Rhipsideigma raffrayi are described in detail and those of Distocupes varians are re-examined. Their morpho logical structures are evaluated with respect to their functional and phylogenetic significance. Larvae of Rhipsideigma are woodborers with a straight body and a wedge-shaped head capsule. Most of their apomorphic features are correlated with their xylobiontic habits. The strong mandibles, the sclerotized ligula and the wedge-shaped head enable the larvae to penetrate rotting wood. The broadened prothorax, prosternal asperities, tergal ampullae, the short legs, and eversible lobes of segment IX play an important role in locomotion in galleries within rotting wood. Leg muscles are weakly developed, whereas the dorsal, pleural and ventral muscula ture is complex. The larval features allow Rhipsideigma to be placed in the clades Archostemata, Cupedidae + Micromalthidae, Cupedidae, Cupedidae excl. Priacma, and Cupedidae excl. Priacma and Distocupes. The monophyly of Cupedidae and Cupedidae, excluding Priacma, so far is only supported by apomorphies of the adults. However, the presence of glabrous patches on the pro sternum and of a medially divided field of asperities may be larval apomorphies of the family. A clade, which comprises Rhip sideigma, Tenomerga and probably other genera of Cupedidae with hitherto unknown larvae, is well supported by larval apomorphies such as the broadened prothorax, the presence of coxal asperities and the presence of a distinct lateral longitudinal bulge. Increased numbers of antennomeres and labial palpomeres are apomorphies only found in larvae of Distocupes.