John F. Lawrence

Family-group names in Coleoptera (Insecta)

Abstract We synthesize data on all known extant and fossil Coleoptera family-group names for the first time. A catalogue of 4887 family-group names (124 fossil, 4763 extant) based on 4707 distinct genera in Coleoptera is given. A total of 4492 names are available, 183 of which are permanently invalid because they are based on a preoccupied or a suppressed type genus. Names are listed in a classification framework. We recognize as valid 24 superfamilies, 211 families, 541 subfamilies, 1663 tribes and 740 subtribes. For each name, the original spelling, author, year of publication, page number, correct stem and type genus are included. The original spelling and availability of each name were checked from primary literature. A list of necessary changes due to Priority and Homonymy problems, and actions taken, is given. Current usage of names was conserved, whenever possible, to promote stability of the classification. New synonymies (family-group names followed by genus-group names): Agronomina Gistel, 1848 syn. nov. of Amarina Zimmermann, 1832 (Carabidae), Hylepnigalioini Gistel, 1856 syn. nov. of Melandryini Leach, 1815 (Melandryidae), Polycystophoridae Gistel, 1856 syn. nov. of Malachiinae Fleming, 1821 (Melyridae), Sclerasteinae Gistel, 1856 syn. nov. of Ptilininae Shuckard, 1839 (Ptinidae), Phloeonomini Ádám, 2001 syn. nov. of Omaliini MacLeay, 1825 (Staphylinidae), Sepedophilini Ádám, 2001 syn. nov. of Tachyporini MacLeay, 1825 (Staphylinidae), Phibalini Gistel, 1856 syn. nov. of Cteniopodini Solier, 1835 (Tenebrionidae); Agronoma Gistel 1848 (type species Carabus familiaris Duftschmid, 1812, designated herein) syn. nov. of Amara Bonelli, 1810 (Carabidae), Hylepnigalio Gistel, 1856 (type species Chrysomela caraboides Linnaeus, 1760, by monotypy) syn. nov. of Melandrya Fabricius, 1801 (Melandryidae), Polycystophorus Gistel, 1856 (type species Cantharis aeneus Linnaeus, 1758, designated herein) syn. nov. of Malachius Fabricius, 1775 (Melyridae), Sclerastes Gistel, 1856 (type species Ptilinus costatus Gyllenhal, 1827, designated herein) syn. nov. of Ptilinus Geoffroy, 1762 (Ptinidae), Paniscus Gistel, 1848 (type species Scarabaeus fasciatus Linnaeus, 1758, designated herein) syn. nov. of Trichius Fabricius, 1775 (Scarabaeidae), Phibalus Gistel, 1856 (type species Chrysomela pubescens Linnaeus, 1758, by monotypy) syn. nov. of Omophlus Dejean, 1834 (Tenebrionidae). The following new replacement name is proposed: Gompeliina Bouchard, 2011 nom. nov. for Olotelina Báguena Corella, 1948 (Aderidae). Reversal of Precedence (Article 23.9) is used to conserve usage of the following names (family-group names followed by genus-group names): Perigonini Horn, 1881 nom. protectum over Trechicini Bates, 1873 nom. oblitum (Carabidae), Anisodactylina Lacordaire, 1854 nom. protectum over Eurytrichina LeConte, 1848 nom. oblitum (Carabidae), Smicronychini Seidlitz, 1891 nom. protectum over Desmorini LeConte, 1876 nom. oblitum (Curculionidae), Bagoinae Thomson, 1859 nom. protectum over Lyprinae Gistel 1848 nom. oblitum (Curculionidae), Aterpina Lacordaire, 1863 nom. protectum over Heliomenina Gistel, 1848 nom. oblitum (Curculionidae), Naupactini Gistel, 1848 nom. protectum over Iphiini Schönherr, 1823 nom. oblitum (Curculionidae), Cleonini Schönherr, 1826 nom. protectum over Geomorini Schönherr, 1823 nom. oblitum (Curculionidae), Magdalidini Pascoe, 1870 nom. protectum over Scardamyctini Gistel, 1848 nom. oblitum (Curculionidae), Agrypninae/-ini Candèze, 1857 nom. protecta over Adelocerinae/-ini Gistel, 1848 nom. oblita and Pangaurinae/-ini Gistel, 1856 nom. oblita (Elateridae), Prosternini Gistel, 1856 nom. protectum over Diacanthini Gistel, 1848 nom. oblitum (Elateridae), Calopodinae Costa, 1852 nom. protectum over Sparedrinae Gistel, 1848 nom. oblitum (Oedemeridae), Adesmiini Lacordaire, 1859 nom. protectum over Macropodini Agassiz, 1846 nom. oblitum (Tenebrionidae), Bolitophagini Kirby, 1837 nom. protectum over Eledonini Billberg, 1820 nom. oblitum (Tenebrionidae), Throscidae Laporte, 1840 nom. protectum over Stereolidae Rafinesque, 1815 nom. oblitum (Throscidae) and Lophocaterini Crowson, 1964 over Lycoptini Casey, 1890 nom. oblitum (Trogossitidae); Monotoma Herbst, 1799 nom. protectum over Monotoma Panzer, 1792 nom. oblitum (Monotomidae); Pediacus Shuckard, 1839 nom. protectum over Biophloeus Dejean, 1835 nom. oblitum (Cucujidae), Pachypus Dejean, 1821 nom. protectum over Pachypus Billberg, 1820 nom. oblitum (Scarabaeidae), Sparrmannia Laporte, 1840 nom. protectum over Leocaeta Dejean, 1833 nom. oblitum and Cephalotrichia Hope, 1837 nom. oblitum (Scarabaeidae).

Phylogeny of the Coleoptera Based on Morphological Characters of Adults and Larvae

Abstract. In order to infer phylogenetic relationships within the extraordinarily speciesrich order Coleoptera, a cladistic analysis is performed, in which 516 adult and larval morphological characters are scored for 359 beetle taxa, representing 314 families or subfamilies plus seven outgroup taxa representing seven holometabolan orders. Many morphological features are discussed at length with accompanying illustrations, and an attempt is made to homologize these and employ a uniform set of terms throughout the order. The resulting data matrix is analyzed using the parsimony ratchet in conjunction with implied weighting. The resulting most parsimonious tree found the order Strepsiptera to be sister to Coleoptera, each of the four coleopteran suborders to be monophyletic and subordinal relationships as follows: (Archostemata + Adephaga) + (Myxophaga + Polyphaga), but without significant support for either clade. The topology of the remainder of the tree is consistent with many prior molecular and morphological analyses, with the monophyly of superfamilies Hydrophiloidea (sensu lato), Scarabaeoidea and Curculionoidea and many currently recognized families and subfamilies are well supported, with weaker support for Elateroidea, Cucujiformia and Phytophaga.

A SYNERGISTIC EFFECT PUTS RARE, SPECIALIZED SPECIES AT GREATER RISK OF EXTINCTION

Theory and empirical evidence have long suggested that some species are extremely vulnerable to extinction because they have combinations of extinction promoting traits. However, ecologists have not considered whether the form of the relationship between traits is additive (not synergistic) or nonadditive (synergistic). We looked at how traits and their interactions were related to the difference in species population growth rates between experimentally fragmented forest and continuous forest. Two traits acted synergistically; natural abundance and degree of specialization interacted so that beetle species that were rare and specialized had a greater reduction in their growth rates in fragments, compared to continuous forest, than the sum of the reductions in growth rates attributable to these traits. In other words, species that were both rare and specialized were especially vulnerable to extinction. From a conservation perspective, an implication of our findings is that making predictions about extinction risk from a single trait, like abundance or population variability, may be risky because traits may act synergistically rendering species more, or less, vulnerable than predicted by that single trait. There is currently a great deal of interest in which traits predict the sensitivity of species to a given threat, but if we are going to look at risks in this way, then we also need to consider how traits interact, because this can alter the vulnerability of species.

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.

Classification of basal Cucujoidea (Coleoptera : Polyphaga): cladistic analysis, keys and review of new families

Phylogenetic relationships among the basal Cucujoidea were reconstructed by a cladistic analysis of a data matrix consisting of 37 exemplar taxa and 99 adult and larval characters. Eight most parsimonious cladograms provided evidence for the polyphyly of Phloeostichidae, the paraphyly of Cucujoidea (with respect to the placement of Trogossitidae), and the monophyly of Protocucujidae + Sphindidae, Biphyllidae + Erotylidae, Cryptophagidae, Cucujidae + Silvanidae, Propalticidae + Laemophloeidae, and the Nitidulidae groups (Nitidulidae, Smicripidae, and Brachypteridae). The following families are elevated from subfamily to family status: Agapythidae (one genus), Phloeostichidae (four genera; the subfamilies Phloeostichinae and Hymaeinae are supressed), Priasilphidae (three genera), Tasmosalpingidae (one genus), and Myraboliidae (one genus). These families are described in detail and adult and larval keys to all families of Cucujoidea are provided. The genus Bunyastichus, gen. nov. (type species: B. monteithi, sp. nov.) is described in the family Phloeostichidae and the family Priasilphidae is revised with the following new taxa: Chileosilpha, gen. nov. (type species: C. elguetai, sp. nov.), Priasilpha (P. angulata, sp. nov., P. aucklandica, sp. nov., P. bufonia, sp. nov., P. carinata, sp. nov., P. earlyi, sp. nov., and P. embersoni, sp. nov.), Priastichus (P. crowsoni, sp. nov. and P. megathorax, sp. nov.).

Relationships and higher classification of some Tenebrionidae and Zopheridae (Coleoptera)

Abstract. Changes in the higher classification of Zopheridae and Tenebrionidae are specified, mostly within the context of Watts (1974b) revised subfamily classification of Tenebrionidae. New or revised diagnoses are provided for adults and/or larvae of several higher taxa, and larval descriptions of several tribes are provided for the first time.

Coleoptera genera of New Zealand

A checklist for the New Zealand genera of Coleoptera is provided and introductory information on the systematics, species number and biology is included. A total of 1091 genera are placed into 82 families and 180 subfamilies.

Three new genera of Indo-Australian Dermestidae (Coleoptera) and their phylogenetic significance

Three new genera are described, providing valuable new information on the phylogeny and biogeography of the family Dermestidae: Derbyana, gen. nov., containing only D. matthewsi, sp. nov. from north-western Australia; Orphilodes, gen. nov., with three Australian species, O. australis, sp. nov. (type of the genus), O. minor, sp. nov. and O. malleecola, sp. nov.; and Trichodryas, gen. nov., with one described species, T. esoterica sp. nov., from the Malay Peninsula and undescribed species from the Sulu Archipelago, Sabah, Kalimantan and Java. A cladistic analysis is included based on a matrix consisting of the three new genera, exemplar genera representing all major groups of Dermestidae and outgroup exemplars from the families Eucinetidae, Derodontidae, Nosodendridae, Endecatomidae and Bostrichidae. Based on the results, a new supergeneric classification is proposed and compared with those in other recent works on Dermestidae and related groups. Recognised subfamilies and tribes include Orphilinae, Attageninnae, Thorictinae, Dermestinae (Marioutini and Dermestini), Megatominae and Trinodinae (Thylodriini and Trinodini). Trichelodini is suppressed and Hexanodes Blair is removed from synonymy with Trichelodes Carter. Keys are given to adults and larvae of the subfamilies of Dermestidae and the genera of Orphilinae, Dermestinae and Trinodinae.

Phylogenetic analysis of the minute brown scavenger beetles (Coleoptera: Latridiidae), and recognition of a new beetle family, Akalyptoischiidae fam.n. (Coleoptera: Cucujoidea)

We infer the first phylogenetic hypothesis for Latridiidae Erichson (Coleoptera: Cucujoidea). Portions of seven genes (18S ribosomal DNA, 28S ribosomal DNA, 12S ribosomal DNA, 16S ribosomal DNA, cytochrome c oxidase I and II and histone III) were analysed. Twenty‐seven latridiid species were included, representing both subfamilies and more than half of the currently recognized genera. Eight outgroup taxa from other families of Cucujoidea were included. Parsimony and partitioned Bayesian analyses were performed on the combined dataset. In both phylogenetic analyses, the enigmatic Akalyptoischion Andrews (Latridiinae) was recovered outside of Latridiidae. The subfamilies Corticariinae and Latridiinae (without Akalyptoischion) were each recovered as monophyletic in both analyses. A new family, Akalyptoischiidae fam.n. is erected based on the results of the phylogenetic study and further support from adult morphology, key features of which are illustrated.

Phylogeny and classification of Corylophidae (Coleoptera: Cucujoidea) with descriptions of new genera and larvae.

Abstract Phylogenetic relationships within the family Corylophidae were investigated. Twenty ingroup taxa and six outgroups were included in a cladistic analysis, based on 48 characters derived from adult and larval morphology. Phylogenetic analysis confirms that Corylophidae are monophyletic within the superfamily Cucujoidea and may be subdivided into two subfamilies: the Australian Periptycinae and the cosmopolitan Corylophinae containing 10 tribes: Foadiini trib.n., Cleidostethini, Aenigmaticini, Parmulini, Sericoderini, Peltinodini, Orthoperini, Corylophini, Teplinini and Rypobiini. All currently recognized family‐group taxa are thoroughly diagnosed, and keys to their identification based on adults and larvae are provided. Two new genera and three species are described: Weirusgen.n., containing only W. tozersp.n. (Australia: Queensland), and Stanusgen.n., with the two species S. bowesteadisp.n. (New Zealand) and S. tasmanicussp.n. (Tasmania). The larvae of Pakalukodes bimaculatusŚlipiński et al. from Queensland and of Stanus bowesteadisp.n. from New Zealand are described and illustrated for the first time.