Phylogenomics resolves timing and patterns in the evolution of Australasian Cerambycinae (Coleoptera: Cerambycidae), and reveals new insights into the subfamily-level classification and historical biogeography of longhorn beetles.

Cerambycinae is the second-largest subfamily of longhorn beetles in the Southern Hemisphere. The phylogeny of Cerambycinae is poorly known, resulting in a highly artificial tribal-level classification and a largely speculative evolutionary history. We reconstructed the phylogenetic relationships of Cerambycinae at the generic level using anchored hybrid enrichment data from hundreds of nuclear genes, with a primary focus on the extraordinarily diverse faunas of Australia and New Zealand. We also estimated divergence times by incorporating fossil calibrations in our analyses. We identified two main clades within Cerambycinae, which can also be separated morphologically by a distinct type of antennal foramen. We recovered a Late Jurassic origin of crown Cerambycinae. Dorcasominae, which was newly found to have representatives in Australia, was notably derived from within Cerambycinae. We recovered two independent origins of Australian Cerambycinae: one clade originated in the Early Cretaceous and is likely endemic to the Southern Hemisphere, while the other clade appears to have immigrated to Australia, perhaps from the Northern Hemisphere. Within the Australian lineages were multiple independent origins of New Zealand taxa, all of which are relative host-plant generalists. Tribal relationships and assignments are discussed, and based on our results, the following major nomenclatural acts were made: Dorcasominae Lacordaire, 1868 is downgraded to a tribe Dorcasomini of Cerambycinae Latreille, 1804; Neostenini Lacordaire, 1868 syn. nov. is treated as a junior synonym of Uracanthini Blanchard, 1851.

Ancient relicts or recent immigrants? Different dating strategies alter diversification scenarios of New Zealand aquatic beetles (Coleoptera: Hydrophilidae: Berosus).

Dated species-level phylogenies are crucial for understanding the origin and evolutionary history of modern faunas, yet difficult to obtain due to the frequent absence of suitable age calibrations at species level. Substitution rates of related or more inclusive clades are often used to overcome this limitation but the accuracy of this approach remains untested. We compared tree dating based on substitution rates with analyses implementing fossil data by direct node-dating and indirect root-age constraints for the New Zealand endemic Berosus water beetles (Coleoptera: Hydrophilidae). The analysis based solely on substitution rates indicated a Miocene colonization of New Zealand and Pleistocene origin of species. By contrast, all analyses that implemented fossil data resulted in significantly older age estimates, indicating an ancient early Cenozoic origin of the New Zealand clade, diversification of species during or after the Oligocene transgression and Miocene-Pliocene origin of within-species population structure. Rate-calibrated time trees were incongruent with recently published Coleoptera time trees, the fossil record of Berosus and the distribution of outgroup species. Strong variation of substitution rates among Coleoptera lineages, as well as among lineages within the family Hydrophilidae, was identified as the principal reason for low accuracy of rate-calibrated analyses, resulting in underestimated node ages in Berosus. We provide evidence that Oligocene to Pliocene events, rather than the Pleistocene Glacial cycles, played an essential role in the formation of the modern New Zealand insect fauna.

Total-evidence backbone phylogeny of Aleocharinae (Coleoptera: Staphylinidae).

Phylogenetic studies of Aleocharinae rove beetles, arguably one of the least known and the largest insect lineages, are compromised by its enormous taxonomic diversity. DNA, a powerful resource for phylogenetics, is not available for numerous extant aleocharine species. We provide a broad comparative morphological study of Aleocharinae to frame molecular datasets for total-evidence analyses. Using full-body dissections and slide-mounting techniques for light microscopy supplemented by scanning electron microscopy, we constructed a morphological matrix across all major taxa focused on non-inquiline tribes of Aleocharinae and outgroups. Phylogenetic analyses of this matrix concatenated with earlier published DNA loci and including exemplar taxa lacking molecular data, resolved outstanding controversies and, among other novelties, showed that: the Habrocerinae + Trichophyinae clade is sister group to Aleocharinae; Hypocyphtini are sister to the rest of the "higher Aleocharinae"; Taxicerini are sister to Aleocharini; Hoplandriini and Placusini are nested within a polyphyletic Oxypodini; Hoplandriini are sister to Meoticina; and Actocharini are nested within Liparocephalini. For the first time, morphological synapomorphies are identified for some large clades of Aleocharinae. In addition, 1252 high-resolution microphotographs of aleocharine structures are made available online with the entire matrix for future research.

Mesozoic sooty mould beetles as living relicts in New Zealand.

New Zealand is an island continent that completed its split from the Gondwanan continent at 52 Ma, harbouring an iconic biota of tuatara, kiwi and weta. The sooty mould community is a distinctive trophic element of New Zealand forest ecosystems that is driven by plant-feeding sternorrhynchan Hemiptera. These produce honeydew, which supports fungal growth, which in turn supports numerous endemic invertebrates, including endemic New Zealand beetle families. Ancient New Zealand insect fossils are rare but a single fossil of a sooty mould cyclaxyrid was recently described from Cretaceous Burmese amber, a family that was previously known from two extant New Zealand species. Well-preserved fossils like this one are recasting Earth history, and, based on a wealth of additional specimens, we re-evaluate the taxonomy of Cretaceous cyclaxyrids and one Eocene species here transferred to Cyclaxyridae. Cyclaxyridae are highly tied to the sooty mould community and have now been discovered to occur in disparate biogeographic realms in deep time. Our discovery indicates that the family, and perhaps the sooty mould community in general, was widespread in Pangaea from at least the Cretaceous and survived as a relict in New Zealand. Persistence of a sooty mould ecosystem in New Zealand and fungal specialization may not necessarily be an evolutionary 'dead-end' for cyclaxyrids and other insects.

Basal polyphagan beetles in mid-Cretaceous amber from Myanmar: biogeographic implications and long-term morphological stasis.

The origin and early evolutionary history of polyphagan beetles have been largely based on evidence from the derived and diverse 'core Polyphaga', whereas little is known about the species-poor basal polyphagan lineages, which include Scirtoidea (Clambidae, Decliniidae, Eucinetidae, and Scirtidae) and Derodontidae. Here, we report two new species Acalyptomerus thayerae sp. nov. and Sphaerothorax uenoi sp. nov., both belonging to extant genera of Clambidae, from mid-Cretaceous Burmese amber. Acalyptomerus thayerae has a close affinity to A. herbertfranzi, a species currently occurring in Mesoamerica and northern South America. Sphaerothorax uenoi is closely related to extant species of Sphaerothorax, which are usually collected in forests of Nothofagus of Australia, Chile, and New Zealand. The discovery of two Cretaceous species from northern Myanmar indicates that both genera had lengthy evolutionary histories, originated at least by the earliest Cenomanian, and were probably more widespread than at present. Remarkable morphological similarities between fossil and living species suggest that both genera changed little over long periods of geological time. The long-term persistence of similar mesic microhabitats such as leaf litter may account for the 99 Myr morphological stasis in Acalyptomerus and Sphaerothorax. Additionally, the extinct staphylinoid family Ptismidae is proposed as a new synonym of Clambidae, and its only included species Ptisma zasukhae is placed as incertae sedis within Clambidae.

Specialized proteinine rove beetles shed light on insect-fungal associations in the Cretaceous.

Insects and fungi have a long history of association in shared habitats. Fungus-feeding, or mycophagy, is remarkably widespread in beetles (Coleoptera) and appears to be a primitive feeding habit that preceded feeding on plant tissues. Numerous Mesozoic beetles belonging to extant fungus-associated families are known, but direct fossil evidence elucidating mycophagy in insects has remained elusive. Here, we report a remarkable genus and species, Vetuproteinus cretaceus gen. et sp. nov., belonging to a new tribe (Vetuproteinini trib. nov.) of the extant rove beetle subfamily Proteininae (Staphylinidae) in Mid-Cretaceous Burmese amber. The mouthparts of this beetle have a markedly enlarged protruding galea bearing an apparent spore brush, a specialized structure we infer was used to scrape spores off surfaces and direct them into the mouth, as in multiple modern spore-feeding beetles. Considering the long evolutionary history of Fungi, the Mid-Cretaceous beetles likely fed on ancient Basidiomycota and/or Ascomycota fungi or spore-producing organisms such as slime moulds (Myxomycetes). The discovery of the first Mesozoic proteinine illustrates the antiquity of the subfamily, and suggests that ancestral Proteininae were already diverse and widespread in Pangaea before the supercontinent broke up.

Antillipeltis, a new genus of Antillean Trogossitidae (Coleoptera: Cleroidea) with a key to the Cleroidea.

Antillipeltis gen. nov. is described based on the following six new extant species and two new fossil species from Hispaniola and Puerto Rico: A. alleni sp. nov. (Dominican Republic, Miocene), A. darlingtoni sp. nov. (Haiti), A. iviei sp. nov. (Dominican Republic, Miocene), A. maculata sp. nov. (Dominican Republic), A. minuta sp. nov. (Dominican Republic), A. nitida sp. nov. (Puerto Rico), A. portoricensis sp. nov. (Puerto Rico), and A. pubescens sp. nov. (Dominican Republic). The genus is placed in Cleroidea, as currently delimited, based on the presence of a distinctive type of aedeagus occurring primarily in this superfamily, plus a combination of features excluding it from other cucujiform superfamilies. Within Cleroidea, the genus is tenatively placed in the family Trogossitidae and subfamily Lophocaterinae, but it differs from all other Trogossitidae in the presence of ventral membranous lobes with adhesive setae on tarsomeres 1-4 and in a combination of 9-segmented antennae, weak 3-segmented antennal club consisting of slightly elongate antenomeres, lack of postcoxal processes on the pronotal hypomera, and unique leg modifications. A key is provided for major groups of Cleroidea and all described genera of Peltinae and Lophocaterinae, with the exception of Rentoniini, based in part on the literature and in part on dissections of adult males. Antillipeltis is one of three endemic West Indian genera of Coleoptera that is both extant and known from Dominican amber.

Illustrated catalogue and type designations of the New Zealand Zopheridae (Coleoptera: Tenebrionoidea).

This paper provides a comprehensive catalogue of the New Zealand members of the family Zopheridae Solier (Coleoptera: Tenebrionoidea) in an effort to stabilize the nomenclature preceding extensive revisionary taxonomy within the group. A checklist of the 17 New Zealand zopherid genera and an account for each of the 189 species (by current combination) is provided. Type material for nearly all species was examined, and type specimens are designated herein (90 confirmed holotypes, 3 confirmed paratypes, 102 lectotypes, 280 paralectotypes). Images of all primary type specimens and labels examined are provided. Pycnomerus sulcatissimus Sharp, 1886 is a junior synonym and secondary homonym of Pycnomerus sulcatissimus (Reitter, 1880). One replacement name is proposed, Chorasus buckleyi new name, for Chorasus subcaecus (Broun), and 23 new combinations are given.

Sternodea arcana, new species (Coleoptera: Cryptophagidae: Caenoscelini): part of the Nearctic fauna?

The leaf litter-inhabiting genus Sternodea Reitter (Cryptophagidae: Cryptophaginae: Caenoscelini) presently contains seven Palearctic species and up to 27 undescribed species from North and Central America. Here we describe Sternodea arcana, new species, ostensibly from Florida but possibly mislabeled, and likely originating from Mexico or Central America. Consequently, the existence of Sternodea among the North American cryptophagid fauna is unknown.

Cedraderus, a new genus for "Xylophilus" constrictus Fall (Coleoptera: Tenebrionoidea: Aderidae).

Cedraderus, new genus is erected for a single species described from California, USA, Cedraderus constrictus (Fall, 1910), new combination, and distinguished from all other aderid genera based on the pronotum having subapical and sub-basal sulci, presence of a supraepipleural groove on the elytron, lack of sexual dimorphism and setal brush on the metafemora, and abdominal ventrites 1 and 2 connate with a complete suture.

Analysis of the diversity and distributional patterns of coleopteran families on a global scale.

Invertebrates make up the majority of all living species on earth. Nevertheless, our understanding of the global distribution of terrestrial biodiversity and regional patterns therein has thus far been almost entirely based on vertebrate and vascular plant patterns. Here we try to provide some information on the global biogeography of the largest invertebrate order, the beetles (Coleoptera). We compile and analyse a database of beetle distributions, containing presence-absence data for 177 coleopteran families across 827 ecoregions. We map family richness and weighted endemism and find the highest values in the Neotropics and Southeast Asia, but also in the temperate northern hemisphere. Important centres of beetle family endemism include the western and southern parts of North America, Central America, temperate South America, Europe, South, Southeast and East Asia, and eastern Australia. A series of UPGMA cluster analyses were used to produce two global regionalisation maps. As analyses on the complete dataset failed to produce contiguous clusters, two sub-datasets were considered. The first one excluded widespread families and family-poor ecoregions, and the second was restricted to endemic-rich ecoregions. The clusters resulting from the first analysis are partly similar to vertebrate- and plant-based regionalisation schemes, with easily discernible and extensive Holarctic and Holotropical regions, but also include numerous smaller regions, mostly in the temperate parts of the southern Hemisphere (Andean, Patagonian, Matorral, New Guinean, New Caledonian, southern African), with Australia split between a tropical and a temperate/subtropical part. New Zealand is not analysed in this reduced dataset, but clusters with New Caledonia in the endemic-rich analysis, where further interesting subdivisions emerge (mainly in East Asia and western North America). We discuss these patterns in the light of coleopteran dispersal and the habitat requirements predominant across the coleopteran phylogeny, while considering the age of beetle families.

Fast-tracking bespoke DNA reference database generation from museum collections for biomonitoring and conservation.

Despite recent advances in high-throughput DNA sequencing technologies, a lack of locally relevant DNA reference databases limits the potential for DNA-based monitoring of biodiversity for conservation and biosecurity applications. Museums and national collections represent a compelling source of authoritatively identified genetic material for DNA database development, yet obtaining DNA barcodes from long-stored specimens may be difficult due to sample degradation. Here we demonstrate a sensitive and efficient laboratory and bioinformatic process for generating DNA barcodes from hundreds of invertebrate specimens simultaneously via the Illumina MiSeq system. Using this process, we recovered full-length (334) or partial (105) COI barcodes from 439 of 450 (98%) national collection-held invertebrate specimens. This included full-length barcodes from 146 specimens which produced low-yield DNA and no visible PCR bands, and which produced as little as a single sequence per specimen, demonstrating high sensitivity of the process. In many cases, the identity of the most abundant sequences per specimen were not the correct barcodes, necessitating the development of a taxonomy-informed process for identifying correct sequences among the sequencing output. The recovery of only partial barcodes for some taxa indicates a need to refine certain PCR primers. Nonetheless, our approach represents a highly sensitive, accurate and efficient method for targeted reference database generation, providing a foundation for DNA-based assessments and monitoring of biodiversity.

Integrated phylogenomics and fossil data illuminate the evolution of beetles.

Beetles constitute the most biodiverse animal order with over 380 000 described species and possibly several million more yet unnamed. Recent phylogenomic studies have arrived at considerably incongruent topologies and widely varying estimates of divergence dates for major beetle clades. Here, we use a dataset of 68 single-copy nuclear protein-coding (NPC) genes sampling 129 out of the 193 recognized extant families as well as the first comprehensive set of fully justified fossil calibrations to recover a refined timescale of beetle evolution. Using phylogenetic methods that counter the effects of compositional and rate heterogeneity, we recover a topology congruent with morphological studies, which we use, combined with other recent phylogenomic studies, to propose several formal changes in the classification of Coleoptera: Scirtiformia and Scirtoidea sensu nov., Clambiformia ser. nov. and Clamboidea sensu nov., Rhinorhipiformia ser. nov., Byrrhoidea sensu nov., Dryopoidea stat. res., Nosodendriformia ser. nov. and Staphyliniformia sensu nov., and Erotyloidea stat. nov., Nitiduloidea stat. nov. and Cucujoidea sensu nov., alongside changes below the superfamily level. Our divergence time analyses recovered a late Carboniferous origin of Coleoptera, a late Palaeozoic origin of all modern beetle suborders and a Triassic-Jurassic origin of most extant families, while fundamental divergences within beetle phylogeny did not coincide with the hypothesis of a Cretaceous Terrestrial Revolution.

Reconciling phylogeography and ecological niche models for New Zealand beetles: Looking beyond glacial refugia.

Mitochondrial DNA (cox1) sequence data and recently developed coalescent phylogeography models were used to construct geo-spatial histories for the New Zealand fungus beetles Epistranus lawsoni and Pristoderus bakewelli (Zopheridae). These methods utilize continuous-time Markov chains and Bayesian stochastic search variable selection incorporated in BEAST to identify historical dispersal patterns via ancestral state reconstruction. Ecological niche models (ENMs) were incorporated to reconstruct the potential geographic distribution of each species during the Last Glacial Maximum (LGM). Coalescent analyses suggest a North Island origin for E. lawsoni, with gene flow predominately north-south between adjacent regions. ENMs for E. lawsoni indicated glacial refugia in coastal regions of both main islands, consistent with phylogenetic patterns but at odds with the coalescent dates, which implicate much older topographic events. Dispersal matrices revealed patterns of gene flow consistent with projected refugia, suggesting long-term South Island survival with population vicariance around the Southern Alps. Phylogeographic relationships are more ambiguous for P. bakewelli, although long-term survival on both main islands is evident. Divergence dates for both species are consistent with the topographic evolution of New Zealand over the last 10Ma, whereas the signature of the LGM is less apparent in the time-scaled phylogeny.

New Australian Erotylinae with notes on Dacnini (Coleoptera: Cucujoidea: Erotylidae).

In preparation for upcoming studies, several new taxa of Australian Erotylinae (Coleoptera: Erotylidae) need to be described to make names available. New genera and species in Dacnini and Megalodacnini are described. Echinothallis banderbearella, new genus and species (Dacnini); Microdacne, new genus, with four new species (Dacnini), M. gloriousa, M. lamingtonia, M. nardia, M. styxia; and Episcaphula (Tropidoscaphula) megalodacnoides, new species (Megalodacnini). Variation of female terminalia and other characters in several genera are discussed regarding the monophyly and generic diversity of Dacnini.

Nomenclatural notes for some Australian Erotylinae (Coleoptera: Erotylidae).

In the subfamily Erotylinae (Coleoptera: Erotylidae), several nomenclatural concerns in the Australian fauna are corrected for upcoming publications. Spelling and attribution of the genus "Aulacochilus" is discussed and is correctly cited as Aulacocheilus Dejean, 1836. The species Episcaphula tetrastica Lea, 1921, becomes Aulacocheilus leai (Mader, 1934), new combination. Through a previous synonymy of Tritoma australiae Lea, 1922 with Hedista tricolor Weise, 1927, and the subsequent transfer of T. australiae into Spondotriplax Crotch, 1876, the genus Hedista Weise, 1927 is recognized as a synonym of Spondotriplax Crotch, 1876, new synonymy. Cosmoscaphula Heller, 1920, a subgenus of Episcaphula Crotch, 1876, is discussed. The type species, E. (C.) tamburinea Heller, 1920, was found to have two new synonyms: Episcaphula (Cosmoscaphula) clatrata Heller, 1920 and Episcaphula rufolineata Wilson, 1921. A lectotype is here designated for E. (C.) tamburinea Heller, 1920. Two genus-species combinations used in museum and internet-based lists have not been formally published. They are discussed, illustrated, and officially proposed here: Hoplepiscapha laticollis (Carter, 1908) and Neothallis bizonata (Macleay, 1887), new combinations.

Sexual Dimorphism of New Zealand Puppet Beetles (Aderidae, Coleoptera, Tenebrionoidea): Systematic Revision, Description of Three New Genera, and Phylogeny for Zenascus, gen. n.

The Aderidae (Coleoptera: Tenebrionoidea) of New Zeland are revised to include four genera and fourteen species. Three genera are described as new: one distributed throughout the Australasian region (Zenascus gen. n.) one endemic to the north and south islands of New Zealand (Transrenus gen. n.), and one that is endemic to the south island of New Zealand (Pseudozena gen. n.). Six species are also newly described (Pseudozena denticulata sp. n., Transrenus thulater sp. n., Zenascus roberti sp. n., Z. incensum sp. n., Z. elenae sp. n., Z. aurum sp. n.). All previously described New Zealand species of aderids contained in the preoccupied genus Xylophilus are transferred to the newly erected genus Zenascus, resulting in six new combinations (Z. antennalis (Broun), comb. n.; Z. coloratus (Broun), comb. n.; Z. luniger (Champion), comb. n.; Z. nitidus (Broun), comb. n.; Z. obscurus (Broun), comb. n.; Z. xenarthrus (Broun, 1910: 54), comb. n.). Holotype and lectotype designations are made or verified for all previously described species. The New Zealand species Xylophilus pictipes Broun is synonymized with Zenascus obscurus, syn. n. and Scraptogetus nigricans is synonymized with Scraptogetus anthracinus, syn. n. The Australian genus Pseudananca Blackburn 1893 is synonymized with the New Zealand genus Scraptogetus Broun, syn. n. Keys to the genera and species are included. Phenotypic characters previously employed in the higher classification of the Aderidae, including secondary sexually dimorphic characters, are discussed and a phylogenetic analysis for the ten New Zealand species of Zenascus is performed to reconstruct trait evolution in males, which display extreme dimorphic antennomere modifications.

Revision of Eupines King of New Zealand (Coleoptera: Staphylinidae: Pselaphinae: Goniaceritae).

The species-rich genus Eupines King of New Zealand is revised to include 48 species, with 22 known species redescribed. Twenty-two new species are described: Eupines (Byraxis) brevis sp. n., E. (B.) caesta sp. n., E. (B.) carinata sp. n., E. (B.) coalita sp. n., E. (B.) complector sp. n., E. (B.) dugdalei sp. n., E. (B.) gigas sp. n., E. (B.) graceae sp. n., E. (B.) huizhenae sp. n., E. (B.) hoarei sp. n., E. (B.) insolita sp. n., E. (B.) mayae sp. n., E. (B.) minuta sp. n., E. (B.) obtusa sp. n., E. (B.) ovalis sp. n., E. (B.) pannicula sp. n., E. (B.) petila sp. n., E. (B.) protibialis sp. n., E. (B.) whirinaki sp. n., E. (B.) waikaremoana sp. n., Eupines (Eupines) undecim sp. n. and Eupines novem sp. n. The following 11 synonymies are proposed: E. (B.) dispar (Sharp) (= E. (B.) munroi Broun syn. n.), E. (B.) hectori Broun (= E. (B.) diversides Newton syn. n.), E. (B.) impar (Sharp) (= E. (B.) sanguineua Broun syn. n., E. (B.) foveatissima Broun syn. n.), E. (B.) monstrosa (Reitter) (= E. (B.) rudicornis Broun syn. n., E. (B.) costata Broun syn. n.), E. (B.) mundula (Schaufuss) (= E. (B.) forficulida Broun syn. n.), E. (B.) paganus (Broun) (= E. (B.) allocera Broun syn. n., E. (B.) sylvicola Broun syn. n., E. (B.) rhyssarthra Broun syn. n.) and Eupinolus altulus (Broun) (= E. (E.) nasuta Broun syn. n.). New combinations are proposed for four species previously described in Eupines: Eupinolus calcaratus (Broun) new comb., Eupinolus nasutus (Broun) new comb., Gastrobothrus ignotus (Broun) new comb. and Gastrobothrus platynotus (Broun) new comb. A key and distribution maps for all New Zealand members of Eupines are provided.

A new genus of Pharaxonothinae from Australia (Coleoptera: Erotylidae).

Australonotha gen. nov. (type species: A. monteithi sp. nov.) is described for two specimens collected from Mount Bellender Ker, Queensland, Australia. Apart from having a monocondylic meso-metaventrital articulation, Australonotha shows similarities with other erotylid subfamilies including Dacninae, Cryptophilinae and some smaller bodied glabrous Languriinae. Australonotha is placed in Pharaxonothinae based on the presence of a transverse occipital ridge and absence of abdominal calli. It can be distinguished from members of all other erotylid taxa by the pronotum lacking a basal transverse groove and having a pair of submedial longitudinal pronotal grooves that are deep and widened posteriorly, have sharp edges laterad, and form part of a distinct rectilinear impression in the basal third.

A new Himalayan species of Alfieriella (Coleoptera: Cryptophagidae).

A new species of Alfieriella Wittmer, 1935 (Coleoptera, Cryptophagidae), Alfieriella senguptai sp. n. from China and India, is described. This is the first formal record of the genus Alfieriella and the tribe Hypocoprini from the Himalayan region. Alfieriella senguptai is the largest member of Alfieriella, and its presence in a cold, high-altitude environment conforms to Bergmann's rule. The distribution of the genus Alfieriella may be associated with the breakup of the Tethys Ocean and the origin of A. senguptai influenced by the Himalayan orogeny. A distribution map and a key to species of Alfieriella are also provided.