Key questions on the evolution and biogeography of New Zealand alpine insects.

New Zealand alpine environments host a diverse assemblage of insect lineages, with virtually every major insect group represented. The modern mountain ranges of New Zealand are relatively young and large areas of habitat above the tree line have only been in continual existence for the past one million years. We discuss the geological history and physical characteristics of New Zealand alpine environments and the resulting selective pressures placed on insect species. Some notable alpine taxa and previous faunistic research is highlighted. We discuss examples where single lineages have colonised the alpine zone and contrast these with larger radiations of alpine species which in some cases are the result of multiple colonisation events. The age of most alpine lineages is consistent with the young geological age of the mountains, nevertheless there are some much older alpine lineages of uncertain evolutionary history. We show that alpine species have employed a very broad range of morphological, physiological, and behavioural adaptations to survive in the alpine zone, and new studies are starting to unpick their genomic basis. Finally, we look to the future and assess threats to the unique New Zealand alpine insect fauna.

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.

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.

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.

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.

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.

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.

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.

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.

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.

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 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.

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.

Generic Review of New Zealand Chrysomelinae (Coleoptera: Chrysomelidae).

The leaf beetle subfamily Chrysomelinae is reviewed for New Zealand. The native fauna consists of six genera, three new, all of which are described: Aphilon Sharp, 1876; Caccomolpus Sharp, 1886; Chalcolampra Blanchard, 1853; Mauroda gen. nov.; Nanomela gen. nov.; Zeaphilon gen. nov.. Chalcolampra is the senior synonym of Cyrtonogetus Broun, 1915 (comb. nov.). These genera include 51 species, nine newly described and eight in new combinations, as follows: Caccomolpus laticollis (Broun, 1893) comb. nov., from Aphilon; C. pretiosus (Broun, 1880) comb. nov., from Aphilon; Chalcolampra crassa (Broun, 1915) comb. nov., from Cyrtonogetus; Maurodus arcus sp. nov.; M. cinctiger (Broun, 1921) comb. nov., from Caccomolpus; M. impressus (Broun, 1914) comb. nov., from Aphilon; M. lepidus sp. nov.; M. maculatus (Broun, 1893) comb. nov., from Caccomolpus; M. nunni sp. nov.; M. occiduus sp. nov.; M. ornatus (Broun, 1910) comb. nov., from Caccomolpus; M. owenensis sp. nov.; M. plagiatus (Sharp, 1886) comb. nov., from Caccomolpus; M. supernus sp. nov.; Nanomela tiniheke sp. nov.; Zeaphilon marskeae sp. nov.; Z. mirandum sp. nov.. All 11 species in the genus Maurodus are described and a key given for their identification. Type material of the New Zealand species of Aphilon (10 species), Caccomolpus (14 species) and Chalcolampra (13 species) is reviewed and lectotypes designated for 16 species, as well as M. cinctiger. A type species is designated for Caccomolpus: C. globosus Sharp, 1886. Holotypes are confirmed for 26 species. Seven genera and 13 species of exotic Chrysomelinae also occur in New Zealand and their presence is briefly reviewed. One of these exotics has been misnamed as Paropsisterna variicollis (Chapuis, 1877), a junior synonym of P. cloelia (Stål, 1860) (syn. nov.). A key to all genera of Chrysomelinae in New Zealand is provided.

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.

Catalogue and type designations for New Zealand Goniaceritae (Coleoptera: Staphylinidae: Pselaphinae).

A catalogue for the eight Goniaceritae genera that occur in New Zealand is presented, treating a total of 62 species. Seventeen holotypes are confirmed, 38 lectotypes and 99 paralectotypes are designated herein for all valid species but Eupines acceptus Broun, for which type material was not located. One synonymy is proposed: Anabaxis electrica (King, 1863) (= Rybaxis brevis Oke, 1928). Four species of Eupines known only by females are valid species, but without subgeneric assignment. Images of primary type specimens and their original labels are provided.

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.