Alfred F. Newton

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.

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.

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.

Phylogenetic placement of Arrowinini trib.n. within the subfamily Staphylininae (Coleoptera: Staphylinidae), with revision of the relict South African genus Arrowinus and description of its larva

Abstract.  The rove beetle genus Arrowinus, a flightless endemic of South Africa, is revised. Arrowinus phaenomenalis Bernhauer, the type species of the genus, is redescribed based on an examination of type and additional material. Three new species, A. relictussp.n., A. peckorumsp.n. and A. minutussp.n., are described. For all four species, an identification key for adults and larvae is provided, along with available data on their distribution and bionomics. Phylogenetic relationships among the four species of Arrowinus were sought by means of a cladistic analysis. Based on the presumed larvae of A. peckorum and other species of the genus (identified by association with the respective adults), the first larval description for Arrowinus is provided. By means of a cladistic analysis of morphological characters of both adults and larvae, analysed separately and in combination, the phylogenetic relationships of Arrowinus with other members of the subfamily Staphylininae were revealed and are discussed. To facilitate this discussion, current problems of phylogeny and systematics are summarized for all Staphylininae. In agreement with an earlier hypothesis, the analysis confirmed that Arrowinus does not belong to the subtribe Quediina of the tribe Staphylinini (where it has hitherto resided) but constitutes a separate taxon, sister to the tribe Staphylinini, for which a new tribe Arrowinini is established. The analysis provides support for the monophyly of the tribe Staphylinini, the subfamily Staphylininae, and the clade (Staphylininae + Paederinae). Adult and larval identification keys to all tribes of Staphylininae are provided.

A review of the New Zealand rove beetles (Coleoptera: Staphylinidae)

Abstract A preliminary review of the subfamilies and genera of Staphylinidae known from New Zealand is presented, with data on natural history and selected references. Taxonomic changes involving 15 new combinations and a new synonymy are given in an appendix. The New Zealand staphylinid fauna as reported here includes approximately 936 native and 85 adventive species in approximately 178 genera and 16 subfamilies. The native fauna is characterised by the absence of many major lineages (including half of all staphylinid subfamilies), the high level of endemism of genera (43%) and species (>90%), and the radiation of several groups of genera and species. Staphylinid diversity in New Zealand is compared with that of several other regions. The size of the New Zealand fauna of rove beetles is consistent with species number/land area relationships in a variety of areas around the world.

Larvae of Trichophya and phylogeny of the tachyporine group of subfamilies (Coleoptera: Staphylinidae) with a review, new species and characterization of the Trichophyinae

Abstract. Larvae of the staphylinid subfamily Trichophyinae are described for the first time based on larvae of a new species of Trichophya from the southwestern United States. Adults and larvae of the new species, Trichophya texana Ashe & Newton (type locality Texas, Brewster Co., Big Bend National Park), are described and illustrations of both provided. Also given are a key for separation of the Nearctic species of Trichophya, a checklist of the known World fauna of the Trichophyinae (including first report of the genus from Mexico and Guatemala), and a characterization of the subfamily Trichophyinae based on both larvae and adults. The relationships of major genera and higher taxa in the tachyporine group of staphylinid subfamilies are analysed cladistically using larval characters. No larval characters were found that provide evidence for the monophyly of the tachyporine group; no evidence was found for the monophyly of the Tachyporinae; Charhyphus, Olisthaerus and Phloeocharis (Phloeocharinae + Olisthaerinae) form a monophyletic group; the Trichophyinae and Habrocerinae are sister groups and together probably are the sister group to the Aleocharinae; the Aleocharinae are confirmed to be monophyletic based on larval characters; and Gymnusa + Deinopsis form the sister group to the remainder of the Aleocharinae.

Data Sets, Partitions, and Characters: Philosophies and Procedures for Analyzing Multiple Data Sets

We compared four approaches for analyzing three data sets derived from staphylinoid beetles, a superfamily whose known species diversity is roughly comparable to that of vertebrates. One data set is derived from adult morphology and the two molecular data sets are from 12S ribosomal RNA and cytochrome b mitochondrial DNA. We found that taxonomic congruence following conditional data combination, herein called compatible evidence (CE), resolved more nodes compatible with an initial conservative hypothesis than did total evidence (TE), conditional data combination (CDC), or taxonomic congruence (TC). CE sets a base of nodes obtained by CDC analysis and then investigates what further agreement may arise in a universe where these nodes are accepted as given. We suggest that CE75-75 may be appropriate for future studies that aim to both generate a well-corroborated tree and investigate conflicts between data sets, partitions, and characters. CE75-75 is a 75% bootstrap consensus CDC tree followed by combinable-component consensus of a 75% bootstrap consensus of each homogeneous set of partitions having hierarchical structure.

Remarkable Stasis in a Phloeocharine Rove Beetle from the Late Cretaceous of New Jersey (Coleoptera, Staphylinidae)

The first definitive fossil species of the rove beetle (Staphylinidae) subfamily Phloeocharinae is described and figured from a single individual preserved in Late Cretaceous (Turonian) amber from New Jersey. The species is representative of the extant genus Phloeocharis Mannerheim and is described as Phloeocharis agerata Chatzimanolis, Newton, and Engel, new species. The specimen was imaged using traditional light microscopy as well as synchrotron propagation phase contrast microtomography, permitting a detailed examination of otherwise difficult to observe features. Examination revealed remarkable homogeneity across many characters with those of extant relatives, highlighting considerable morphological stasis in the genus over the last 90 million years.

Larval Description, Adult Feeding Behavior, and Phylogenetic Placement of Megalopinus (Coleoptera: Staphylinidae)

Abstract Larvae of the genus Megalopinus, of the monobasic subfamily Megalopsidiinae, are described for the first time. Larvae can be distinguished from all other known staphylinid larvae by the presence of a mandible with a subapical bladelike process, and are differentiated in detail from other larvae of the staphylinine group of subfamilies in a key. To confirm the identity of Megalopinus larvae, which have not been reared to adults, we compared partitions of 28 adult and 20 larval morphological characters derived from members of the stenine group (Megalopsidiinae, Euaesthetinae, Steninae) and three staphylinid outgroups (Oxyporinae, Pseudopsinae and Piestinae). Partitioned cladograms were similar with the only difference being the placement of Pseudopsinae as sister taxon to the stenine group (adult characters) or included within it (larval characters). Characters were combined and one tree was produced with the following relationships: Megalopsidiinae (Pseudopsinae (Steninae, Euaesthetinae)). Megalopinus is a specialized genus occurring mainly under rotting logs where fungal growth occurs, a habit for which the new term hyphyledic is proposed. Adults digest their food preorally, using a rotary-mill method for extracting liquefied tissues from masticated prey items. A deeply bifurcate labrum bearing modified setae functions as a sieve, while hyaline processes on the labium may be used for tearing captured prey by Megalopinus adults. Implications for the evolution of the adult Stenus “stick-capture” method for prey capture, which involves extruding a rodlike labium that exudes sticky substances at the tip, are discussed based on new phylogenetic information.