David C. Hawks

The beetle tree of life reveals that Coleoptera survived end‐Permian mass extinction to diversify during the Cretaceous terrestrial revolution

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 provide a uniquely well‐resolved temporal and phylogenetic framework for studying patterns of innovation and diversification in Coleoptera, and a foundation for further sampling and resolution of the beetle tree of life.

Evolution of the hymenopteran megaradiation

The Hymenoptera--ants, bees and wasps--represent one of the most successful but least understood insect radiations. We present the first comprehensive molecular study spanning the entire order Hymenoptera. It is based on approximately 7 kb of DNA sequence from 4 gene regions (18S, 28S, COI and EF-1α) for 116 species representing all superfamilies and 23 outgroup taxa from eight orders of Holometabola. Results are drawn from both parsimony and statistical (Bayesian and likelihood) analyses, and from both by-eye and secondary-structure alignments. Our analyses provide the first firm molecular evidence for monophyly of the Vespina (Orussoidea+Apocrita). Within Vespina, our results indicate a sister-group relationship between Ichneumonoidea and Proctotrupomorpha, while the stinging wasps (Aculeata) are monophyletic and nested inside Evaniomorpha. In Proctotrupomorpha, our results provide evidence for a novel core clade of proctotrupoids, and support for the recently proposed Diaprioidea. An unexpected result is the support for monophyly of a clade of wood-boring sawflies (Xiphydrioidea+Siricoidea). As in previous molecular studies, Orussidae remain difficult to place and are either sister group to a monophyletic Apocrita, or the sister group of Stephanidae within Apocrita. Both results support a single origin of parasitism, but the latter would propose a controversial reversal in the evolution of the wasp-waist. Generally our results support earlier hypotheses, primarily based on morphology, for a basal grade of phytophagous families giving rise to a single clade of parasitic Hymenoptera, the Vespina, from which predatory, pollen-feeding, gall-forming and eusocial forms evolved.

A molecular phylogeny of the Chalcidoidea (Hymenoptera).

Chalcidoidea (Hymenoptera) are extremely diverse with more than 23,000 species described and over 500,000 species estimated to exist. This is the first comprehensive phylogenetic analysis of the superfamily based on a molecular analysis of 18S and 28S ribosomal gene regions for 19 families, 72 subfamilies, 343 genera and 649 species. The 56 outgroups are comprised of Ceraphronoidea and most proctotrupomorph families, including Mymarommatidae. Data alignment and the impact of ambiguous regions are explored using a secondary structure analysis and automated (MAFFT) alignments of the core and pairing regions and regions of ambiguous alignment. Both likelihood and parsimony approaches are used to analyze the data. Overall there is no impact of alignment method, and few but substantial differences between likelihood and parsimony approaches. Monophyly of Chalcidoidea and a sister group relationship between Mymaridae and the remaining Chalcidoidea is strongly supported in all analyses. Either Mymarommatoidea or Diaprioidea are the sister group of Chalcidoidea depending on the analysis. Likelihood analyses place Rotoitidae as the sister group of the remaining Chalcidoidea after Mymaridae, whereas parsimony nests them within Chalcidoidea. Some traditional family groups are supported as monophyletic (Agaonidae, Eucharitidae, Encyrtidae, Eulophidae, Leucospidae, Mymaridae, Ormyridae, Signiphoridae, Tanaostigmatidae and Trichogrammatidae). Several other families are paraphyletic (Perilampidae) or polyphyletic (Aphelinidae, Chalcididae, Eupelmidae, Eurytomidae, Pteromalidae, Tetracampidae and Torymidae). Evolutionary scenarios discussed for Chalcidoidea include the evolution of phytophagy, egg parasitism, sternorrhynchan parasitism, hypermetamorphic development and heteronomy.

Phylogeny and evolution of Staphyliniformia and Scarabaeiformia: forest litter as a stepping stone for diversification of nonphytophagous beetles

The beetle series Staphyliniformia exhibits extraordinary taxonomic, ecological and morphological diversity. To gain further insight into staphyliniform relationships and evolution, we reconstructed the phylogeny of Staphyliniformia using DNA sequences from nuclear 28S rDNA and the nuclear protein‐coding gene CAD for 282 species representing all living families and most subfamilies, a representative sample of Scarabaeiformia serving as a near outgroup, and three additional beetles as more distant outgroups. Under both Bayesian inference (BI) and maximum likelihood inference (MLI), the major taxa within Staphyliniformia are each monophyletic: (i) Staphylinoidea, (ii) Hydrophiloidea s.l., and the contained superfamilies (iii) Hydrophiloidea s.s. and (iv) Histeroidea, although Staphylinoidea and Hydrophiloidea s.l. are not strongly supported by MLI bootstrap. Scarabaeiformia is monophyletic under both methods of phylogenetic inference. However, the relative relationships of Staphylinoidea, Hydrophiloidea s.l. and Scarabaeiformia differ between BI and MLI: under BI, Staphyliniformia and Scarabaeiformia were sister groups; under MLI, Hydrophiloidea s.l. and Scarabaeiformia were sister groups and these together were sister to Staphylinoidea. The internal relationships in Scarabaeiformia were similar under both methods of phylogenetic inference, with Cetoniinae, Dynastinae + Rutelinae, Hybosoridae, Passalidae, Scarabaeidae and Scarabaeinae recovered as monophyla. Histeridae comprised two major clades: (1) Abraeinae, Trypanaeine and Trypeticinae; and (2) Chlamydopsinae, Dendrophilinae, Haeteriinae, Histerinae, Onthophilinae, Saprininae and Tribalinae. The relationships among early‐divergent Hydrophiloidea differed between BI and MLI, and overall were unresolved or received only moderate to low nodal support. The staphylinoid families Agyrtidae, Hydraenidae and Ptiliidae were recovered as monophyletic; the latter two were sister taxa, and Staphylinidae + Silphidae was also monophyletic. Silphidae was placed within Staphylinidae in close relation to a subset of Tachyporinae. Pselaphinae and Scydmaeninae were both recovered within Staphylinidae, in accordance with recent analyses of morphological characters, although not always with recently proposed sister taxa. None of the four major groups of Staphylinidae proposed by Lawrence and Newton (1982) was recovered as monophyletic. Certain highly specialized staphyliniform habits and morphologies, such as abdominal defensive glands and reduced elytra, have arisen in parallel in separate lineages. Further, our analyses support two major transitions to an aquatic lifestyle within Staphyliniformia: once within Staphylinoidea (Hydraenidae), and once within Hydrophiloidea s.l. (Hydrophiloidea s.s.). On a smaller scale, the most common transition is from litter to subcortical or to periaquatic microhabitats and the next most common is from litter to carrion and to fungi. Overall, transitions to periaquatic microhabitats were the most numerous. The broad picture in Staphyliniformia seems to be a high level of evolutionary plasticity, with multiple possible pathways to and from many microhabitat associations, and litter as a major source microhabitat for diversification. In Scarabaeiformia, the most common transitions were from litter to foliage, with flowers to litter, litter to flowers, and litter to dung being next, and then litter to roots, logs or carrion. Litter is again the largest overall source microhabitat. The most common transitions were to foliage and flowers. It thus seems that the litter environment presents ecological and evolutionary opportunities/challenges that facilitate entry of Staphyliniformia and Scarabaeiformia into ‘new’ and different ecological adaptive zones.

AN OVERVIEW OF THE CLASSIFICATION AND EVOLUTION OF THE MAJOR SCARAB BEETLE CLADES (COLEOPTERA: SCARABAEOIDEA) BASED ON PRELIMINARY MOLECULAR ANALYSES

Abstract We present a preliminary overview of our molecular phylogenetics research on the superfamily Scarabaeoidea. The molecular data consists of 28S ribosomal DNA sequences (mainly D2 and D3 expansion regions) for over 600 taxa and 18S ribosomal DNA sequences (mainly E17 to E35 expansion regions) for over 150 representative taxa within the lineages sampled. Based on our preliminary molecular phylogenetic results, Scarabaeoidea includes three major groups: 1) Geotrupidae, Passalidae, and Pleocomidae; 2) Lucanidae, Diphyllostomatidae, Trogidae, and Glaresidae; and 3) Hybosoridae, Ochodaeidae, Glaphyridae, and Scarabaeidae. The broad evolutionary patterns within the Scarabaeoidea are discussed with respect to phylogeny and evolution.

Phylogeny and behaviour of the Gollumiellinae, a new subfamily of the ant-parasitic Eucharitidae (Hymenoptera: Chalcidoidea)

Abstract.  Gollumiellinae is proposed as a new subfamily for the Indo‐Pacific genera Gollumiella Hedqvist and Anorasema Bouc̆ek based on analyses of three ribosomal transcript gene regions (28S‐D2 and ‐D3, and 18S‐E23; 1262 aligned base pairs) sequenced for twenty‐eight genera and fifty‐four species of Eucharitidae, and twelve genera and nineteen species of Pteromalidae (Pteromalinae) and Perilampidae (Chrysolampinae and Perilampinae). Gollumiella and Anorasema have been treated as either a monophyletic or paraphyletic group within the Eucharitini (Hymenoptera: Eucharitidae). The monophyly of the Eucharitidae and a sister‐group relationship between Gollumiellinae and Oraseminae +  Eucharitinae are supported strongly in parsimony and Bayesian analyses. The molecular phylogeny contradicts previous morphological hypotheses, in which Gollumiella and Anorasema are within Eucharitini. The strength of the molecular hypothesis is explored through evaluations of data alignments that are eye optimized, clustal‐x aligned, eye optimized but with gaps coded as a fifth base, and eye optimized and combined with morphological data. The oviposition behaviour and host associations of G. longipetiolata on tree ferns (Cyathea latebrosa, Cyatheaceae), and the morphology of the planidium and pupa are newly described. Eggs are deposited vertically, with the base anchored into the leaf or petiolar surface. Unlike other Eucharitidae, the eggs are not stalked, but rather tipped with a secretion that may act as an attractant for their ant host, Paratrechina sp. (Formicidae: Formicinae). Various morphological character state optimizations and their implications for convergent morphology, behaviour and host associations are discussed. All results using gene regions treated separately or combined with the morphological data reach the same conclusion: Anorasema + Gollumiella form the sister group of Oraseminae + Eucharitinae, and thus deserve subfamily status. This suggests that very distinct traits such as the fusion of the pronotum and prepectus in adults and the fusion of the first two tergites of the planidia are convergent apomorphies. Molecular data, when strongly supported, can provide new information to unravel convergent from synapomorphic changes, resulting in more robust hypotheses of relationship.

Molecular phylogenetics and evolution of the food relocation behaviour of the dung beetle tribe Eucraniini (Coleoptera : Scarabaeidae : Scarabaeinae)

A phylogenetic analysis using 28S and 18S rDNA provides evidence that the tribe Eucraniini is a monophyletic group and the sister-group of the Phanaeini and Dichotomiini. Our molecular phylogeny of the dung beetle tribes provides strong evidence for the monophyly of the subfamily Scarabaeinae. The monophyly of the tribe Eucraniini is well supported and it includes the genera Anomiopsoides Blackwelder, Ennearabdus van Lansberge, Eucranium Brulle and Glyphoderus Westwood. The food-lifting relocation behaviour present in species of Eucranium, Anomiopsoides and Glyphoderus is considered a derived condition and it most probably evolved from tunnelling behaviour. The preference for dry dung or dung pellets by species of Eucraniini genera, and feeding on plant material by species of Anomiopsoides, are considered apomorphic. Our analyses suggest that rolling behaviour in the Scarabaeinae evolved at least twice during their evolution. The incidence of high endemicity of dung beetles in the Monte biogeographic province of Argentina suggests that the area constitutes an independent centre of evolution. Our hypothesis is that a vicariant event was responsible for the divergence of the Eucraniini from a Neotropical lineage ancestral to Eucraniini and Phanaeini. The isolation of the Eucraniini lineage probably occurred after the Andean uplift during the Quechua diastrophism (middle Miocene) that resulted in the creation of xeric plains in austral regions of South America.

Systematics of Moths in the Genus Catocala (Lepidoptera: Noctuidae). V. Neotypification of Names in the Nearctic Fauna

Abstract Nine previously published neotype designations in the noctuid moth genus Catocala are reviewed for compliance with the International Code of Zoological Nomenclature. Eight of the designations are found to have not fulfilled one (75.3.2) of the seven qualifying conditions of Article 75.3 when these were originally published, and are redesignated herein. The eight Catocala names involved are connubialis Guenée, dollii Beutenmüller, grotiana Bailey, irene Behr, micronympha Guenée, stretchii Behr, texanae French and walshii Edwards. The ninth neotype designation, for calphurnia Henry Edwards, is considered valid as originally published.

A review of the primary types of the Hawaiian stag beetle genus Apterocyclus Waterhouse (Coleoptera, Lucanidae, Lucaninae), with the description of a new species

Abstract The species of the Hawaiian stag beetle genus Apterocyclus Waterhouse (Coleoptera: Lucanidae) are reviewed following an examination of all primary types. Although the continued existence of the species is unknown and some possibly are extinct there are five recently extant species, including one species that is described here as new. The holotypes for all available names are pictured, and synonymies discussed and updated. Lectotypes are designated for Apterocyclus honoluluensis Waterhouse and A. munroi Sharp. A key to species and a revised catalog for the genus are provided.