Hydrophiloidea

Abstract: 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.

Abstract: The phylogeny and evolutionary history of the water scavenger beetles (Coleoptera: Hydrophilidae) are inferred from comprehensive analyses of DNA sequence data from the mitochondrial genes COI, COII and 16S and the nuclear genes 18S, 28S and arginine kinase. Bayesian and maximum parsimony analyses included 151 taxa, representing all subfamilies, tribes and subtribes that have ever been proposed in the family, as well as representatives of the hydrophiloid families Helophoridae, Hydrochidae, Spercheidae, Epimetopidae and Georissidae. The resulting well-supported trees strongly disagree with prior classifications of the Hydrophilidae, suggesting that the smaller subfamilies (Horelophinae, Horelophopsinae and Sphaeridiinae) are derived from within the larger Hydrophilinae. The existing tribal classification is more compatible with our results, but many significant differences are evident. Here, we present a new classification of the Hydrophilidae comprising 6 subfamilies and 12 tribes. Each subfamily and tribe is reviewed in detail with (i) a morphological diagnosis, including known or putative morphological synapomorphies, (ii) its taxonomic circumscription, including genera not included in our analyses, and (iii) a review of its general biology and geographic distribution. A new identification key to subfamily and tribe based on adult morphology is also provided. The newly adopted classification requires the following taxonomic changes: the subfamily Hydrophilinae sensu n. is redefined to include only the tribes Amphiopini stat.n. (removed from the synonymy with the Chaetarthriini), Berosini, Laccobiini, Hydrophilini and Hydrobiusini (= Sperchopsini syn.n.); the subfamily Chaetarthriinae stat.n. is removed from synonymy with the Hydrophilinae and includes the tribes Chaetarthriini and Anacaenini (= Horelophinae syn.n.); the Acidocerinae stat.n. (= Horelophopsinae syn.n.) and Rygmodinae stat.n. (= Andotypini syn.n., Borborophorini syn.n. and Tormissini syn.n.) are elevated to subfamily rank; and the subfamily Enochrinae subfam.n. is established for the genus Enochrus and its relatives. The implications for the morphological evolution, ecological transitions and biogeography of the family are discussed.

Abstract: Additions and corrections to the World Catalogue of Hydrophiloidea (HANSEN 1999) are presented. All new taxa and synonymies as well as all new combinations and taxa with new status are summarized. New distributional data for previously described species are not covered. Aschius MAKHAN, 2004, Rishihydroius MAKHAN, 2001, Satishius MAKHAN, 2004 and Soesilius MAKHAN, 2001 are synonymized with Hydrochus LEACH. There are now 3151 valid species of Hydrophiloidea distributed in 181 genera.

Abstract: . One hundred and twenty-one morphological characters of larvae and adults of the series Staphyliniformia were scored (multistate coding) and analysed to determine the family group relationships of the polyphagan groups Scarabaeoidea, Histeroidea, Hydrophiloidea and Staphylinoidea. Cladograms were rooted with exemplars of Adephaga, Archostemata, Myxophaga and the polyphagan families Dascillidae, Derodontidae, Eucinetidae and Scirtidae. Analyses of the same dataset with multistate characters re-coded as presence/absence (144 characters) produced cladograms that were similar to those produced from analyses of the original characters. Cladograms produced from partitioned larval and adult characters differed strongly, with adult-only trees more similar to those produced by combined data. The results confirm the monophyly of Hydrophiloidea + Histeroidea and of Staphylinoidea (including Hydraenidae). The Epimetopidae + Georissidae are the only strongly supported clade within Hydrophiloidea. A clade comprising Hydrochidae, Spercheidae and Hydrophilidae, and a sister-group relationship between the latter two families were confirmed in analyses of the data with presence/absence coding. Helophoridae, Epimetopidae and Georissidae are probably not a monophyletic unit, and additional evidence is needed for a reliable placement of Helophoridae. Scarabaeoidea are placed as a sister taxon of Hydrophiloidea + Histeroidea, but support for this relationship is weak. The branching pattern ((Hydraenidae + Ptiliidae) + (Leiodidae + Agyrtidae)), and a clade comprising Scydmaenidae, Silphidae and Staphylinidae (= ‘staphylinid group’) are well founded. The branching pattern (Orchymontiinae + (Prosthetopinae + (Ochthebiinae + Hydraeninae))) within Hydraenidae is confirmed. Poor resolution at the base of the trees and the placement of some nonstaphyliniform taxa (Dascillidae, Derodontidae, Scirtidae and Eucinetidae) as a sister group to a clade comprising Scarabaeoidea, Hydrophiloidea and Histeroidea suggests that Staphyliniformia may be paraphyletic. It is recommended that series names are eliminated from the classification of Polyphaga, at least for the more ‘primitive’ groups.