Phylogeny of the hyper‐diverse rove beetle subtribe Philonthina with implications for classification of the tribe Staphylinini (Coleoptera: Staphylinidae)

With 71 genera and over 2700 described species, Philonthina is the most speciose subtribe of rove beetle tribe Staphylinini and forms a major component of the largest remaining higher systematics challenge in Staphylinini, the ‘Staphylinini propria’ clade. A related systematics issue concerns the position of the genus Holisus (Hyptiomina), which was recovered within the Neotropical philonthine lineage in several recent analyses of morphology. With the aims of resolving the phylogeny of Philonthina and the position and, thus, validity of Hyptiomina, we performed phylogenetic analyses of the tribe Staphylinini based on molecular (six genes, 4471 bp) and morphological (113 characters) data including 138 taxa from all relevant lineages of Staphylinini. We found that ‘Staphylinini propria’ is a monophylum consisting of six lineages: current subtribes Anisolinina, Philonthina, Staphylinina and Xanthopygina; and two new subtribes, Algonina Schillhammer and Brunke and Philothalpina Chatzimanolis and Brunke. While the previously hypothesized Neotropical lineage of Philonthina was corroborated, Holisus was recovered as a separate subtribe, outside of Philonthina, within an informal ‘Southern Hemisphere clade’. Based on our analyses, we propose tentative new concepts of the polyphyletic genera Belonuchus and Philonthus. We propose the following taxonomic changes: synonymy of the subtribes Staphylinina Latreille (valid name) and Eucibdelina Sharp; resurrection of genera Barypalpus Cameron and Trapeziderus Motschulsky from synonymy with Rientis Sharp and Belonuchus Nordmann, respectively; transfer of 38 Belonuchus species, 16 Hesperus Fauvel species and one Philonthus Stephens species to Trapeziderus as new combinations; transfer of two Hesperus species to Eccoptolonthus Bernhauer as new combinations; transfer of one Belonuchus species to Paederomimus Sharp as a new combination; and transfer of Pridonius Blackwelder new status from its position as a subgenus of Quedius (subtribe Quediina) to Philonthina as a genus, and new combinations for its two described species.     

Molecular phylogeny of the beetle tribe Oxypodini (Coleoptera: Staphylinidae: Aleocharinae)

This is the first study to comprehensively address the phylogeny of the tribe Oxypodini Thomson and its phylogenetic relationships to other tribes within the staphylinid subfamily Aleocharinae. Using the hitherto largest molecular dataset of Aleocharinae comprising of 4599 bp for representatives of 22 tribes, the Oxypodini are recovered as non‐monophyletic. Members of the tribe belong to three distantly related lineages within the Aleocharinae: (i) the Amarochara group as sister clade to the tribe Aleocharini, (ii) the subtribe Tachyusina within a clade that also includes the tribes Athetini and Hygronomini, (iii) all other Oxypodini in a clade that also includes the tribes Placusini, Hoplandriini and Liparocephalini. Based on the inferred phylogeny, five subtribes of the Oxypodini are recognized: Dinardina Mulsant & Rey, Meoticina Seevers, Microglottina Fenyes, Oxypodina Thomson and Phloeoporina Thomson. The following changes in the classification of the Aleocharinae are proposed: (i) Amarochara Thomson is removed from the Oxypodini and placed in the tribe Aleocharini; (ii) the subtribe Taxicerina Lohse of the Athetini is reinstated as tribe Taxicerini to include Discerota Mulsant & Rey, Halobrecta Thomson (both removed from the Oxypodini) and Taxicera Mulsant & Rey; (iii) the subtribe Tachyusina Thomson is excluded from the Oxypodini and provisionally treated as tribe Tachyusini; (iv) the oxypodine subtribe name Blepharhymenina Klimaszewski & Peck is placed in synonymy with the subtribe name Dinardina Mulsant & Rey.     

Revision of the rove beetle genus Antimerus (Coleoptera, Staphylinidae, Staphylininae), a puzzling endemic Australian lineage of the tribe Staphylinini

The genus Antimerus Fauvel, 1878, endemic to eastern Australia and Tasmania and a phylogenetically enigmatic member of the large rove beetle tribe Staphylinini, is revised. The genus and each of its four previously known species are redescribed, and a lectotype is designated for Antimerus punctipennis Lea, 1906. Five species are described as new: Antimerus metallicussp. n., Antimerus jamesrodmanisp. n., Antimerus gracilissp. n., Antimerus bellussp. n. and Antimerus monteithisp. n., so that the number of known species in this genus now totals nine. For the first time Antimerus larvae are described, tentatively identified as Antimerus smaragdinus Fauvel, 1878, Antimerus punctipennis and Antimerus metallicus. Available distributional and bionomic data are provided for each species and summarized in the discussion. Adult and larval morphology of Antimerus and its distribution patterns are discussed in the broader context of new data on the evolution of the entire tribe Staphylinini, and with respect to the formation of the Australian fauna of this tribe. The phylogenetic position of Antimerus within Staphylinini remains unresolved pending a targeted formal study. However, a majority of currently available data suggests that it could be a basal member of the recently recovered monophyletic clade of Staphylinini tentatively called "Staphylinini propria".     

Early evolution of the hyperdiverse rove beetle tribe Staphylinini (Coleoptera: Staphylinidae: Staphylininae) and a revision of its higher classification

The rove beetle tribe Staphylinini (Staphylinidae: Staphylininae) is a monophyletic lineage of over 5500 relatively large and charismatic species, yet its higher classification remains deeply rooted in historical concepts. Despite recent progress toward inferring phylogenetic relationships within this group using morphological and molecular datasets, relationships among taxa that were united under a polyphyletic “Quediina” remain largely unknown. To infer these relationships, we analysed a six‐gene dataset (4370 bp) using parsimony and model‐based analyses and the results were placed in the context of morphology. While all genes contributed synapomorphies for major lineages or relationships between them, carbamoyl synthetase (CAD), topoisomerase I (TP) and wingless (Wg) were the most informative. TP was generally most informative at the level of subtribe, Wg above this level and CAD throughout the tree. The monophyly of Staphylinini was strongly supported and analyses support seven clades that correspond to higher taxonomic levels, four of which are formally described as subtribes here: Acylophorina stat. rev., Cyrtoquediina new subtribe, Erichsoniina new subtribe and Indoquediina new subtribe. The majority of Staphylinini taxa were recovered within a well‐supported “northern hemisphere clade” that is weakly represented in the southern hemisphere. The composition and morphological diagnosis of the “Staphylinini propria” clade are revised, and the pronotum shape historically associated with this group is shown to have evolved multiple times elsewhere in Staphylinini. The genus Stevensia is moved from Staphylinina to Acylophorina based on morphological evidence. Cyrtoquedius stat. nov., previously a subgenus of Quedius, is raised to the genus level. The following 32 new combinations (from Quedius) are proposed: Cyrtoquedius anthracinus (Solsky); C. arrogans (Sharp); C. basiventris (Sharp); C. bolivianus (Sharp); C. bruchi (Bernhauer); C. clypealis (Sharp); C. concolor (Sharp); C. flavicaudus (Sharp); C. flavinasis (Bernhauer); C. frenatus (Erichson); C. graciliventris (Sharp); C. jacobi (Scheerpeltz); C. jocosus (Sharp); C. labiatus (Erichson); C. laeviventris (Bernhauer); C. mexicanus (Sharp); C. ochropygus (Bernhauer); C. ogloblini (Bernhauer); C. ornatocollis (Bierig); C. protensus (Sharp); C. rufinasus (Sharp); C. verecundus (Sharp); C. verres (Smetana); Indoquedius borneensis (Cameron); I. dispersepunctatus (Scheerpeltz); I. javanus (Cameron); I. malaisei (Scheerpeltz); I. micantiventris (Scheerpeltz); I. parallelicollis (Scheerpeltz); I. philippinus (Cameron); I. recticollis (Scheerpeltz); and I. sanguinipennis (Scheerpeltz). Cyrtoquedius verres is recorded from the state of Georgia (USA) for the first time, which, together with its transfer from Quedius, extends the distribution of the Cyrtoquediina significantly northward into the Nearctic.     

Revised systematics and biogeography of ‘Quediina’ of sub‐Saharan Africa: new phylogenetic insights into the rove beetle tribe Staphylinini (Coleoptera: Staphylinidae)

Abstract Quediina, a mega‐diverse conventional subtribe of the rove beetle tribe Staphylinini, is remarkably species rich in the north and south temperate regions of the world. Tropical faunas of this group, and the fauna of the entire Afrotropical biogeographical region (= Ethiopian region, = sub‐Saharan Africa), in contrast, are remarkably poor. The taxonomic study of the quediine genera of Staphylinini from the Afrotropical region reveals misidentifications for many of them. Their phylogenetic study demonstrates polyphyly of Quediina and reveals a new evolutionary pattern for the entire tribe Staphylinini. In particular, the formerly quediine genera Euristus Fauvel, 1899 , Ioma Blackwelder, 1952, Natalignathus Solodovnikov, 2005 , all endemic in the Afrotropical region, belong to the non‐related ‘Staphylinina’, ‘Philonthina propria’ and ‘Tanygnathinina sensu novo’ lineages of Staphylinini, respectively. Contrary to earlier records, the genus Quedius Stephens, 1929 does not occur in Africa south of Sahara: Quedius angularis Cameron, 1948 and Quedius cinctipennis Cameron, 1951 are moved to the genus Philonthus Stephens, 1829. The same is established for the Asian genus Algon Sharp, 1874, formerly for a long time associated with Quediina: African species Algon robustus Wendeler, 1928 is moved to the genus Moeocerus Fauvel, 1899 (here in the ‘Philonthina propria’ lineage); and the misidentification of Algon africanus Bernhauer, 1915, a species that probably belongs to a new genus, is discussed. The phylogenetic affiliation of Afroquedius Solodovnikov, 2006 , a South African endemic, is still ambiguous. Overall, the formerly seen bipolar distribution pattern for the ‘Quediina’ is demonstrated to be an artefact, not a reality to explain. Historical biogeographical explanations are proposed for some of the Afrotropical endemics, partly as an attempt to apply biogeography as an external criterion for the evaluation of the new phylogenetic pattern revealed for Staphylinini. The monotypic genera Euristus and Ioma, as well as Heterothops megalops Cameron, 1959 , the only representative of this widespread genus in the Afrotropical region, are redescribed. Limits and synapomorphies of the genus Heterothops are discussed. The following new combinations and new names are proposed: Philonthus cinctipennis ( Cameron, 1951 ) comb.n. (preoccupied by Philonthus cinctipennis Fauvel, 1875), here replaced by Philonthus pseudoquedius Solodovnikov nom.n. ; Philonthus angularis ( Cameron, 1948 ) comb.n. ; Moeocerus robustus ( Wendeler, 1928 ) comb.n. [preoccupied by Moeocerus robustus (Gestro, 1881)], here replaced by Moeocerus wendeleri Solodovnikov nom.n. A lectotype is designated for Heterothops megalops Cameron, 1959 .     

A new genus and species of the rove beetle tribe Mesoporini from Baltic amber (Coleoptera: Staphylinidae: Aleocharinae)

A new extinct rove beetle, Palaeomesoporus electiricus gen. et sp.n., is described from a single specimen of Eocene Baltic amber. This fossil beetle is placed in the tribe Mesoporini, a ‘basal’ group of the mega-diverse subfamily Aleocharinae. Palaeomesoporus is easily discriminated from other mesoporine genera due to its less-developed antennal club, longer and slender elytron lacking a sinuate posterior margin, and prominently shorter mesotarsus. This finding sheds light on the paleodiversity and evolutionary history of the tribe and ‘basal’ Aleocharinae.     

Revision and phylogenetic assessment of the rove beetle genus Pseudohesperus Hayashi,with broad reference to the subtribe Philonthina (Coleoptera: Staphylinidae: Staphylinini)

This paper studies the phylogeny of the rove beetle subtribe Philonthina, to test its hypothetical monophyly and to unravel the evolutionary relationships of the subtribe and its included genus‐level taxa, with emphasis on the genus Pseudohesperus and its close‐allied relatives. The phylogenetic analyses are based on 105 adult morphological characters and 66 terminal taxa, i.e., all six members of Pseudohesperus, 51 species to represent 29 other genera of the subtribe Philonthina, seven species to represent the other six subtribes of Staphylinini, one species of the tribes Arrowinini, and one of the Platyprosopini. According to the phylogenetic results obtained, the genus Erichsonius should move out from the hitherto‐defined subtribe Philonthina and thus the monophyly of this taxon is challenged. The phylogenetic tree suggests that the genera Hesperus and Belonuchus might not be monophyletic, but the monophyly of Pseudohesperus and the sister relationship between it and Bisnius are well supported. The species‐level phylogenetic relationships of the genus Pseudohesperus reveal a clear pattern of species diversification that can be correlated well with the species' zoogeographical patterns. The paper also revises the taxonomy of Pseudohesperus and describes five new species from China: Pseudohesperus luteus Li & Zhou sp. nov. , Pseudohesperus pedatiformis Li & Zhou sp. nov. , Pseudohesperus tripartitus Li & Zhou sp. nov. , Pseudohesperus sparsipunctatus Li & Zhou sp. nov. , and Bisnius lubricus Li & Zhou sp. nov. An identification key to the species of Pseudohesperus is provided and their geographical distributions are mapped. © 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 163 , 679–722.     

Structure and energy pathways of spermatozoa of the rove beetle Aleochara bilineata (Coleoptera, Staphylinidae)

The morphology of mature spermatozoa of the rove beetle Aleochara bilineata was examined by using scanning and transmission electron microscopy. They are about 1000 mum long and filiform. The acrosome and the nucleus are elongate and each about 20 mum long. A well-developed centriole adjunct region connects the nucleus with the sperm tail. The axoneme reveals the 9 + 9 + 2 pattern of the pterygote sperm flagellum. Two accessory bodies and two mitochondrial derivatives with paracrystalline inclusions are present. Cristae are reduced to the cortical zone of the derivatives. Cytochrome-c oxidase activity was detected within the cristae by DAB-reaction. The energy metabolism of the spermatozoa was investigated by using different inhibitors affecting the mitochondrial and cytoplasmic metabolic pathways. Sperm movement was used as an indicator for the utilization of ATP by the axoneme. In control experiments, the duration of motility was longer than 45 min. In the presence of atractyloside or potassium cyanide the motility duration was not affected. On the other hand, iodoacetic acid in the medium stopped sperm motility within 15 min. This indicates that sperm energy metabolism mainly depends on the glycolytic pathway.     

A basal oxyteline rove beetle (Coleoptera: Staphylinidae) from the Early Cretaceous of China: Oldest record for the tribe Euphaniini

A remarkable new rove beetle, Protodeleaster glaber gen. et sp. nov, is described and illustrated based on two well-preserved specimens from the Lower Cretaceous Yixian Formation of Liaoning Province, China. The new genus is placed in the extant staphylinid subfamily Oxytelinae, and recent tribe Euphaniini, based on several characteristic features (e.g. a single pair of wide paratergites on abdominal segments; open procoxal fissures; contiguous mesocoxae; abdominal sternite II short and poorly sclerotized). This find from the Early Cretaceous documents the oldest fossil representative of the tribe Euphaniini. Morphologically, it resembles most closely the recent genus Platydeleaster Schülke, 2003, an unusual member of the extant Oxytelinae. According to the currently accepted hypothesis of the phylogenetic position of Euphaniini and the prior discovery of other taxa from the Late Jurassic, we suggest the tribe might have first appeared at least as early as the Late Jurassic.     

Food preference of the rove beetle, Atheta coriaria Kraatz （Coleoptera： Staphylinidae） under laboratory conditions

A study, involving laboratory choice tests, was conducted to determine the feeding behavior, based on food preference, of the adult and larval stage of the rove beetle, Atheta coriaria Kraatz when presented with both fresh moistened oatmeal and second instar fungus gnat, Bradysia sp. nr. coprophila （Lintner） larvae in Petri dishes. Rove beetles used in this study came from a laboratory-reared colony. A rating scale from 1 to 5, based on percent missing （1 = 0 to 10%, 2 = 11 to 30%, 3 = 31 to 50%, 4 = 51 to 75%, and 5 = 76 to 100%）, was used to objectively assess the amount of oatmeal and number of fungus gnat larvae consumed by each rove beetle adult and larva. In all the choice tests, A. coriaria adults and larvae preferred to feed on fungus gnat larvae （78% and 69%, respectively） significantly more so than oatmeal （9% and 5%, respectively） based on the amount of oatmeal and number of fungus gnat larvae consumed after 4 and 6 hours. There were relatively minimal differences in the amount of food consumed for both adults and larvae after 4 and 6 hours. The results of this study indicate that oatmeal may be an inexpensive supplemental food source, during the rearing process, which will not inhibit the effectiveness of rove beetles to control fungus gnat larvae when released into greenhouses.     

Multilocus phylogeny defines a new classification of Staphylininae (Coleoptera,Staphylinidae), a rove beetle group with high lineage diversity

We provide the first multilocus molecular phylogeny of a group corresponding to the former subfamily Staphylininae. Results are corroborated by the morphological, biogeographical and palaeobiological evidence to serve as a baseline for an updated suprageneric classification. The former subfamily Staphylininae is proven to be a lineage sister to the monophyletic Paederinae and reclassified according to a robust phylogeny resolving a number of long-standing controversies. The subfamily Xantholininae (revised status) is reinstated to contain the tribes Xantholinini, Othiini, Maorothiini and Diochini. Subfamily Platyprosopinae (revised status) is reinstated for the tribes Platyprosopinini, Arrowinini and †Thayeralinini. For a highly peculiar genus Coomania Cameron, formerly in Diochini, a new subfamily Coomaniinae subfam.n. is established and the composition of Diochini (revised status) is changed accordingly. The subfamily Staphylininae (revised status) is reduced to contain the former tribe Staphylinini only. Elevating this mega-diverse tribe to the subfamily rank opened up an opportunity for its more fractional classification by raising several subtribes to the tribal level as follows: Acylophorini, Afroquediini, Amblyopinini, Antimerini, †Baltognathini, Cyrtoquediini, Erichsoniini, Hyptiomini, Indoquediini, Quediini and Tanygnathinini (revised status for all). As a result, the most species-rich tribe Staphylinini (revised status) is reduced to the more homogeneous lineage containing the subtribes Algonina, Anisolinina, Philonthina, Philothalpina, Staphylinina and Xanthopygina. Morphological synapomorphies and diagnostic characters supporting all newly defined higher taxa are provided. This published work has been registered on ZooBank, http://zoobank.org/urn:lsid:zoobank.org:pub:DED8B042-83C9-4D10-B0CB-B50372B067A9 .     

Molecular phylogeny illuminates Amblyopinini (Coleoptera: Staphylinidae) rove beetles as a target for systematic and evolutionary research

The rove beetle tribe Amblyopinini (Coleoptera: Staphylinidae: Staphylininae) is a recently discovered monophyletic lineage comprising an estimated 1000 or more species of mainly leaf- and log litter-dwelling predatory insects found throughout the southern hemisphere. Of these, a single genus Heterothops Stephens somehow conquered all continents in the northern hemisphere as well. A few lineages of amblyopinines independently evolved into highly derived predators of mammal ectoparasites from free-living ancestors. In return, they are tolerated in the mammal fur and nests, which is a unique example of cleaning symbiosis between insects and vertebrates. For over a century the great majority of free-living southern amblyopinines were incorrectly placed in the northern hemisphere-restricted, and superficially similar, rove beetle genera from the subtribe Quediina. Only their mammal-associated derived forms were understood as amblyopinines, a nonmonophyletic taxon of volatile status and enigmatic sister-group relationships of its various members. Here we present the first comprehensive phylogeny of Amblyopinini inferred with Bayesian analysis of a six-gene molecular dataset (4672 bp) across a broad sample of taxa (90 species). This phylogeny provides a framework for the badly needed taxonomic inventory of this group and, in particular, reveals at least two independent origins of mammal association within the tribe. It frames the upcoming in-depth interdisciplinary exploration of a variety of phenomena such as evolution of the austral biota in response to continental drift and climate change, biotic exchange between southern and northern continents, origin and evolution of beetle–mammal symbiosis, and pathways and constraints of the evolutionary parallelisms.     

Morphology and ultrastructure of paired prototergal glands in the adult rove beetle Philonthus varians (Coleoptera, Staphylinidae)

Philonthus and other genera of Philonthina possess a pair of prototergal glands located in the first abdominal tergum and hidden at rest by hind wings and elytra. In Philonthus varians they occupy the whole length of the tergum and form a pouch-like invaginated reservoir with a scaly glandular zone and a smooth outlet. A grille of long setae covers the opening of each gland. The fine structure of these glands is given for the first time. Three types of cells are found in the glandular epithelium. Epidermal cells underlie the cuticular scales, numerous class 1 secretory cells open in the centre of calyces made of finger-like processes of the cuticle, and class 3 cells are connected to pored tubercles. A cytological comparison is made with the diverse class 1 cells described to date in Coleoptera. In these cells different evolutionary trends are shown in the structure of the cuticular apparatus, particularly in the number, size and position of the cuticular apertures as well as in the length and abundance of epicuticular filaments. A possible defensive function of the prototergal glands against pathogens and their interest for the phylogenetic study of Staphylininae are discussed.     

Termitariophily: expanding the concept of termitophily in a physogastric rove beetle (Coleoptera: Staphylinidae)

1. Termitophily in some rove beetles is commonly attributed to the striking termite worker resemblance that is provided by the beetles' hypertrophic (‘physogastric’) abdomen. However, a termite nest may offer to a termitophile additional benefits, such as a continuously repaired shelter. 2. This could apply to Corotoca melantho (Aleocharinae: Corotocini), a viviparous obligatory termitophile staphylinid beetle species. While conferring morphological congruence to its host worker termites, its physogastry may impair mobility, leading to vulnerability and the need for a secure environment. It seems plausible to hypothesise that physogastry in C. melantho would imply in interactions between this termitophile and its host termites as well as its host termitarium. 3. This study provides evidence to build such a hypothesis by inspecting the morpho‐anatomical reproductive traits of this termitophile. It was found that a gradient of growth stages of embryos and larvae in the oviduct explains physogastry in females while pointing to iteroparity. The asynchronous development of oocytes in females, combined with a full developing sequence of sperm cells indicative of continuous spermatogenesis in males, suggests frequent matings. 4. While improving guest–host similarity, physogastry and flightlessness should confer vulnerability to pregnant females, forcing C. melantho to seek close and sheltered environments. These could facilitate the frequent male–female contacts demanded by iteroparity. It is possible, therefore, that physogastry in C. melantho is not only associated with the termites themselves but also with the physical structure of the termitarium. Thus, C. melantho can be hypothesised to be a termitariophile in addition to being a termitophile.     

Biology and distribution of the littoral rove beetle Psamathobledius punctatissimus (Le Conte) (Coleoptera: Staphylinidae)

The intertidal macrofauna on a small sheltered marine beach at Pawley's Island, South Carolina, was dominated by insects, primarily Psamathobledius punctatissimus, a small beetle which attained densities of up to 2 260 adults m⁻². Both adult and larval beetles remained buried in the sand when covered by the tide, then emerged to form mole-like surface trails and feed upon diatoms when exposed. Eggs and young larvae were maintained within special maternal burrows, while larger larvae and non-breeding adults occupied their own individual burrows. Both adults and larvae became comatose when in direct contact with water, but recovered after as long as six hours submergence. In the field, burrows retained air during tidal coverage, allowing the beetles to avoid direct contact with sea water. Seaward extension of the population is probably limited by time available for feeding, while longshore distribution is restricted by sand texture and diatom concentration, or by sediment mobility in more exposed locations. Although very locally distributed, P. punctatissimus has achieved considerable success in a habitat poorly exploited by competitors of either marine or terrestrial origin. Contribution No. 434 of the Belle W. Baruch Institute for Marine Biology and Coastal Research Contribution No. 434 of the Belle W. Baruch Institute for Marine Biology and Coastal Research     

The ecology of the marine rove beetle,Bryothinusa spp. (Coleoptera: Staphylinidae) in Hong Kong

Bryothinusa spp. are common marine insect in Hong Kong. They occur in the intertidal zone of sandy shores between 0.6 and 1.2 m tide level. They emerge when the tide recedes, possibly to mate and feed, then burrow again at the advance of the incoming tide.Bryothinusa has a special respiratory apparatus for long submersions and therefore is able to inhabit the changeable intertidal zone where the competition with other living organisms is comparatively low.     

Molecular phylogeny of Omaliinae (Coleoptera: Staphylinidae) and its implications for evolution of atypically long elytra in rove beetles

Staphylinidae, or rove beetles, are a megadiverse family known for their typically very short elytra exposing most of the abdomen, but the putatively early-derived subfamily Omaliinae and its relatives have been known to include multiple taxa with unusually long elytra. The ancestral elytral length of the family and of this subfamily have long been debated. We present a phylogenetic analysis of Omaliinae based on partial mitochondrial COI (1488 bp), COII (366 bp), 12S rDNA (353 bp), nuclear 18S rDNA (1814 bp), 28S rDNA (876 bp) and CAD (869 bp) data. In all, 51 species in 31 genera and four outgroup species were included. The concatenated sequences were analysed by both parsimony- and model-based (Bayesian and maximum likelihood) methods. The subfamily Omaliinae was not supported as a monophyletic group. The model-based analyses (Bayesian and maximum likelihood trees) showed Empelinae nested within Omaliinae (excluding Corneolabiini), whereas parsimony analysis found all three putative ingroup subfamilies, Empelinae, Glypholomatinae and Microsilphinae, grouped within Omaliinae. Within the Omaliinae, the tribes Coryphiini and Eusphalerini were each supported as monophyletic, whereas Anthophagini and Omaliini were each nonmonophyletic. We hypothesize that there have been at least four independent origins of long elytra from short elytra in the omaliine lineage.     

Phylogeny of the rove beetle tribe Gymnusini sensu n. (Coleoptera: Staphylinidae: Aleocharinae): implications for the early branching events of the subfamily

The rove beetle subfamily Aleocharinae is the largest subfamily of animals known in terms of species richness. Two small aleocharine tribes, Gymnusini and Deinopsini, are believed to be a monophyletic clade, sister to the rest of the Aleocharinae. Although the phylogenetic relationships of the extant lineages have been well investigated, the monophyly of Gymnusini has been questioned due to a series of previous studies and the recent discovery of the aleocharine †Cretodeinopsis Cai & Huang (Deinopsini) from mid‐Cretaceous Burmese amber. Using an additional specimen of †Cretodeinopsis and well‐preserved specimens of †Electrogymnusa Wolf‐Schwenninger from Eocene Baltic amber, we present here two types of morphology‐based phylogenetic analyses, employing all extant/extinct genera of Gymnusini and Deinopsini for the first time. The maximum parsimony and Bayesian analyses recovered a monophyletic clade of the two tribes combined, but each analysis suggested nonmonophyly of Gymnusini. In agreement with the results of the present study, we synonymize Deinopsini syn.n. under Gymnusini sensu n. , by priority. This published work has been registered in ZooBank, http://zoobank.org/urn:lsid:zoobank.org:pub:F09EB444‐C6CA‐4525‐A986‐3CFC826F5877 .