Wioletta Tomaszewska

Phylogeny and classification of Cucujoidea and the recognition of a new superfamily Coccinelloidea (Coleoptera: Cucujiformia)

A large‐scale phylogenetic study is presented for Cucujoidea (Coleoptera), a diverse superfamily of beetles that historically has been taxonomically difficult. This study is the most comprehensive analysis of cucujoid taxa to date, with DNA sequence data sampled from eight genes (four nuclear, four mitochondrial) for 384 coleopteran taxa, including exemplars of 35 (of 37) families and 289 genera of Cucujoidea. Maximum‐likelihood analyses of these data present many significant relationships, some proposed previously and some novel. Tenebrionoidea and Lymexyloidea are recovered together and Cleroidea forms the sister group to this clade. Chrysomeloidea and Curculionoidea are recovered as sister taxa and this clade (Phytophaga) forms the sister group to the core Cucujoidea (Cucujoidea s.n.). The nitidulid series is recovered as the earliest‐diverging core cucujoid lineage, although the earliest divergences among core Cucujoidea are only weakly supported. The cerylonid series (CS) is recovered as monophyletic and is supported as a major Cucujiform clade, sister group to the remaining superfamilies of Cucujiformia. Currently recognized taxa that were not recovered as monophyletic include Cucujoidea, Endomychidae, Cerylonidae and Bothrideridae. Biphyllidae and Byturidae were recovered in Cleroidea. The remaining Cucujoidea were recovered in two disparate major clades: one comprising the nitidulid series + erotylid series + Boganiidae and Hobartiidae + cucujid series, and the other comprising the cerylonid series. Propalticidae are recovered within Laemophloeidae. The cerylonid series includes two major clades, the bothriderid group and the coccinellid group. Akalyptoischiidae are recovered as a separate clade from Latridiidae. Eupsilobiinae are recovered as the sister taxon to Coccinellidae. In light of these findings, many formal changes to cucujiform beetle classification are proposed. Biphyllidae and Byturidae are transferred to Cleroidea. The cerylonid series is formally recognized as a new superfamily, Coccinelloidea stat.n. Current subfamilies elevated (or re‐elevated) to family status include: Murmidiidae stat.n., Teredidae stat.n., Euxestidae stat.n., Anamorphidae stat.rev., Eupsilobiidae stat.n., and Mycetaeidae stat.n. The following taxa are redefined and characterized: Cleroidea s.n., Cucujoidea s.n., Cerylonidae s.n., Bothrideridae s.n., Endomychidae s.n. A new subfamily, Cyclotominae stat.n., is described. Stenotarsinae syn.n. is formally subsumed within a new concept of Endomychinae s.n.

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.

Two new genera of Epilachnini Mulsant from New Guinea and Aru Islands (Coleoptera: Coccinellidae)

The species formerly classified in the genus Afidentula Kapur from New Guinea and Aru Islands are revised. As a result of detailed analysis of mouth parts and genital structures of the species, two new genera are proposed: monotypic Lalokia gen. nov. from Aru Islands and New Guinea, with Lalokia aruensis (Crotch) (comb. nov.), and Papuaepilachna gen. nov. from New Guinea including Papuaepilachna bivacana (Bielawski) (comb. nov.), P. kapuri (Bielawski) (comb. nov.), P. nasti (Bielawski) (comb. nov.), P. tenmana (Bielawski) (comb. nov.), P. watalai (Jadwiszczak) (comb. nov.) and P. wiebesi (Bielawski) (comb. nov.). All species are illustrated and an identification key to the species of Papuaepilachna is provided. A key to the genera of Epilachnini from New Guinea and Aru Islands is also provided. http://www.zoobank.org/urn:lsid:zoobank.org:pub:F721B07F-CBF0-4516-853D-1FD434FE06F1

Molecular phylogeny reveals food plasticity in the evolution of true ladybird beetles (Coleoptera: Coccinellidae: Coccinellini)

BackgroundThe tribe Coccinellini is a group of relatively large ladybird beetles that exhibits remarkable morphological and biological diversity. Many species are aphidophagous, feeding as larvae and adults on aphids, but some species also feed on other hemipterous insects (i.e., heteropterans, psyllids, whiteflies), beetle and moth larvae, pollen, fungal spores, and even plant tissue. Several species are biological control agents or widespread invasive species (e.g., Harmonia axyridis (Pallas)). Despite the ecological importance of this tribe, relatively little is known about the phylogenetic relationships within it. The generic concepts within the tribe Coccinellini are unstable and do not reflect a natural classification, being largely based on regional revisions. This impedes the phylogenetic study of important traits of Coccinellidae at a global scale (e.g. the evolution of food preferences and biogeography).ResultsWe present the most comprehensive phylogenetic analysis of Coccinellini to date, based on three nuclear and one mitochondrial gene sequences of 38 taxa, which represent all major Coccinellini lineages. The phylogenetic reconstruction supports the monophyly of Coccinellini and its sister group relationship to Chilocorini. Within Coccinellini, three major clades were recovered that do not correspond to any previously recognised divisions, questioning the traditional differentiation between Halyziini, Discotomini, Tytthaspidini, and Singhikaliini. Ancestral state reconstructions of food preferences and morphological characters support the idea of aphidophagy being the ancestral state in Coccinellini. This indicates a transition from putative obligate scale feeders, as seen in the closely related Chilocorini, to more agile general predators.ConclusionsOur results suggest that the classification of Coccinellini has been misled by convergence in morphological traits. The evolutionary history of Coccinellini has been very dynamic in respect to changes in host preferences, involving multiple independent host switches from different insect orders to fungal spores and plants tissues. General predation on ephemeral aphids might have created an opportunity to easily adapt to mixed or specialised diets (e.g. obligate mycophagy, herbivory, predation on various hemipteroids or larvae of leaf beetles (Chrysomelidae)). The generally long-lived adults of Coccinellini can consume pollen and floral nectars, thereby surviving periods of low prey frequency. This capacity might have played a central role in the diversification history of Coccinellini.

Phylogeny and evolution of phytophagous ladybird beetles (Coleoptera: Coccinellidae: Epilachnini), with recognition of new genera

Phytophagous ladybird beetles of the tribe Epilachnini are a cosmopolitan, species‐rich group of significant economic importance as pests of agricultural crops. The tribe is well characterized morphologically and clearly monophyletic, but very little is known about its internal phylogenetic relationships and their genus‐level taxonomy. In order to infer the evolutionary history of Epilachnini, test its monophyly and provide a phylogeny‐based classification, we assembled a comprehensive dataset, consisting of four DNA markers (18S and 28S rRNA and 16S, COI mtDNA) and a matrix of 104 morphological characters for 153 species of Epilachnini representing all previously recognised genera, ∼11% of the known species, and 14 outgroup taxa. Molecular, morphological and combined datasets were analysed using maximum likelihood, parsimony and Bayesian inference. Bayes factors and Approximately Unbiased tests (AU) were used to compare alternative phylogenetic hypotheses of unconstrained and backbone‐constrained analysis. Only 14 of the 25 included genera were recovered monophyletic, as originally defined. Afidentula Kapur, Afidenta Dieke, Afissula Kapur, Epilachna Chevrolat, Henosepilachna Li Toxotoma Weise and Mada Mulsant are shown to be poly‐ or paraphyletic; Chnootriba Chevrolat, Subafissa Bielawski, Lalokia Szawaryn & Tomaszewska and Papuaepilachna Szawaryn & Tomaszewska form monophyletic groups within larger clades of genus level. All of these genera are redefined here. The two largest genera of Epilachnini, Epilachna Chevrolat and Henosepilachna Li were represented by multiple monophyletic clades, which we described as new genera: Chazeauiana Tomaszewska & Szawaryn gen.n.; Diekeana Tomaszewska & Szawaryn gen.n.; Fuerschia Tomaszewska & Szawaryn gen.n. and Ryszardia Tomaszewska & Szawaryn gen.n. The following new synonyms are proposed: Afissa Dieke (=Afissula Kapur); Henosepilachna Li in Li & Cook (=Subafissa Bielawski); Papuaepilachna Szawaryn & Tomaszewska (=Lalokia Szawaryn & Tomaszewska).

Chapinaria, New Genus of Chilocorini for Endochilus meridionalis Sicard from Africa (Coleoptera: Coccinellidae)

Abstract. Chapinaria, a new genus of Chilocorini (Coleoptera: Coccinellidae) from Tanzania and Zambia is described and illustrated. It is established for Endochilus meridionalis Sicard, 1929 (comb. nov.). Adult characters concerning similarities to other Chilocorini genera are discussed. Lecototype for Endochilus meridionalis Sicard is designated. Key to Afrotropical genera of Chilocorini is presented. A checklist to the known species of the tribe from Afrotropic Ecozone is also provided.

Revision of the Genus Orcus Mulsant (Coleoptera: Coccinellidae: Chilocorini)

Abstract. Species of the genus Orcus Mulsant, 1850 are revised, keyed and illustrated. Orcus biroi var. ruficollis Weise, 1902 is regarded as a synonym of Orcus biroi Weise, 1902 new synonym. Lectotypes are designated for Orcus biroi Weise, Orcus biroi var. ruficollis Weise, Orcus cinctus Weise and Orcus nigricollis Weise. Three new species, all from New Guinea, are described: Orcus cordiformis, O. tetrafasciatus, O. viridulus. Orcus carinicus Gorham, O. bipunctatus Gorham and O. quadriguttatus Gorham are removed from Chilocorini and transferred to the genus Sticholotis Crotch (Sticholotidini) (new combinations). Distribution, nomenclatural history and diagnoses are provided for each species.

Descriptions of new Neotropical Lycoperdininae (Coleoptera: Endomychidae) and their phylogenetic placement

Abstract Two new genera and three new species of Lycoperdininae (Coleoptera: Endomychidae) from South America, are described and illustrated based on adults: Hylaperdina n. g., along with H. brevicornis n. sp. from Ecuador and H. costariciensis n. sp. from Costa Rica, and Chileanus n. g., along with C. talca n. sp. from Chile. Their most likely placement within the Lycoperdininae is discussed. A key to genera of the neotropical Lycoperdininae is provided.

Fauna Europaea: Coleoptera 2 (excl. series Elateriformia, Scarabaeiformia, Staphyliniformia and superfamily Curculionoidea)

Abstract Fauna Europaea provides a public web-service with an index of scientific names (including synonyms) of all living European land and freshwater animals, their geographical distribution at country level (up to the Urals, excluding the Caucasus region), and some additional information. The Fauna Europaea project covers about 230,000 taxonomic names, including 130,000 accepted species and 14,000 accepted subspecies, which is much more than the originally projected number of 100,000 species. This represents a huge effort by more than 400 contributing specialists throughout Europe and is a unique (standard) reference suitable for many users in science, government, industry, nature conservation and education. Coleoptera represent a huge assemblage of holometabolous insects, including as a whole more than 200 recognized families and some 400,000 described species worldwide. Basic information is summarized on their biology, ecology, economic relevance, and estimated number of undescribed species worldwide. Little less than 30,000 species are listed from Europe. The Coleoptera 2 section of the Fauna Europaea database (Archostemata, Myxophaga, Adephaga and Polyphaga excl. the series Elateriformia, Scarabaeiformia, Staphyliniformia and the superfamily Curculionoidea) encompasses 80 families (according to the previously accepted family-level systematic framework) and approximately 13,000 species. Tabulations included a complete list of the families dealt with, the number of species in each, the names of all involved specialists, and, when possible, an estimate of the gaps in terms of total number of species at an European level. A list of some recent useful references is appended. Most families included in the Coleoptera 2 Section have been updated in the most recent release of the Fauna Europaea index, or are ready to be updated as soon as the FaEu data management environment completes its migration from Zoological Museum Amsterdam to Berlin Museum für Naturkunde.

Epilachnini (Coleoptera: Coccinellidae)—A Revision of the World Genera

Based on the recent revised generic classification of the tribe Epilachnini (Szawaryn et al. 2015), all 27 genera are re-described, diagnosed, illustrated, and included in an identification key. The following nomenclatural changes are made: Epilachna (Hypsa) Mulsant 1850, Epilachna (Cleta) Mulsant 1850, Solanophila Weise 1898, Epilachna (Aparodentata) Wang and Cao 1993, and Epilachna (Uniparodentata) Wang and Cao 1993 are removed from synonymy of Epilachna Chervolat 1837. The subgenus Cleta of Epilachna is raised to the genus level, as Cleta Mulsant 1850 stat. nov.; the subgenus Uniparodentata of Epilachna is raised to the genus level, as Uniparodentata Wang and Cao 1993 stat. nov. Chazeauiana Tomaszewska and Szawaryn 2015 (type species, Epilachna sahlbergi Mulsant 1850), and Epilachna (Hypsa) Mulsant 1850 (type species, Epilachna nigrolimbata Thomson 1875) are synonymized here under the name Cleta Mulsant 1850 (type species, Epilachna eckloni Mulsant 1850)—new synonyms; Fuerschia Tomaszewska and Szawaryn 2015 (type species, Coccinella canina Fabricius 1781) is synonymized with Solanophila Weise 1898 (type species, Epilachna gibbosa Crotch 1874)—new synonym; Ryszardia Tomaszewska and Szawaryn 2015 (type species, Epilachna decipiens Crotch 1874) and Epilachna (Aparodentata) Wang and Cao, 1993 (type species, Epilachna yongshanensis Cao and Xiao 1984) are synonymized under the name Uniparodentata Wang and Cao 1993 (type species, Epilachna paramagna Pang and Mao 1979)—new synonyms. Henosepilachna (Elateria) Fürsch 1964 (type species: Coccinella elaterii Rossi 1794) is removed from synomyms of Henosepilachna Li 1961 [type species, Coccinella sparsa Herbst 1786 (=Coccinella vigintioctopunctata Fabricius 1775)] and is synonymized here with Chnootriba Chevrolat 1837 (type species: Coccinella similis Thunberg 1781)—new synonym. Coccinella flavofasciata Laporte 1840, Epilachna aequatorialis Gordon 1975, E. bizonata Crotch 1874, E. convergens Crotch 1874, E. cruciata Mulsant 1850, E. dubia Crotch 1874, E. monovittata Gordon 1975, E. orthostriata Gordon 1975, E. paracuta Gordon 1975, E. patricia Mulsant 1850, E. satipensis Gordon 1975, and E. univittata Crotch 1874 are transferred to Toxotoma Weise 1900 (comb. nov.); Afissa chapini Dieke 1947, A. complicata Dieke 1947, A. convexa Dieke 1947, A. magna Dieke 1947, A. militaris Dieke 1947, A. quadricollis Dieke 1947, A. subacuta Dieke 1947, A. szechuana Dieke 1947, Epilachna boymi Jadwiszczak and Węgrzynowicz 2003, E. crepida Pang and Ślipiński 2012, E. decipiens Crotch 1874, E. dorotae Bielawski 1979, E. hamulifera Pang and Ślipiński 2012, E. malleforma Peng, Pang and Ren 2002, E. siphodenticulata Hoàng 1983, E. angusta Li 1961, E. bifibra Li 1961, E. chingjing Yu and Wang 1999, E. circumdata Hoàng 1978, E. circummaculata Pang and Mao 1977, E. clematicola Cao and Xiao 1984, E. exornata Bielawski 1965, E. folifera Pang and Mao 1979, E. fugongensis Cao and Xiao 1984, E. glochisifoliata Pang and Mao 1979, E. gressiti Li 1961, E. lata Li 1961, E. madanensis Zeng 2002, E. media Li 1961, E. mobliteratiae Li 1961, E. yongshanensis Cao and Xiao 1984, Solanophila acuta Weise 1900, and S. saginata Weise 1902 are transferred to Uniparodentata Wang and Cao 1993 (comb. nov.); Coccinella canina Fabricius 1781, Epilachna dregei Mulsant 1850, E. infirma Mulsant 1850, E. murrayi Crotch 1874 and E. paykullii Mulsant 1850 are transferred to Solanophila Weise 1898 (comb. nov.); Afissula antennata Bielawski 1967, A. rana Kapur 1958, A. uniformis Pang and Mao 1979, Epilachna ampliata Pang and Mao 1979, E. flavimarginalis Hoàng 1978, E. max Pang and Ślipiński 2012, E. parvula Crotch, E. plicata Weise 1889, and E. sanscrita Crotch 1874 are transferred to Afissa Dieke 1947 (comb. nov.); Epilachna papuensis Crotch 1874 and Subafissa brittoni Bielawski 1963 are transferred to Henosepilachna Li 1961 (comb. nov.); Epilachna admirabilis Crotch 1874, E. alternans Mulsant 1850, E. glochinosa Pang and Mao 1979, E. hopeiana Miyatake 1985, E. insignis Gorham 1892, E. macularis Mulsant 1850, E. parainsignis Pang and Mao 1979, and Solanophila maxima Weise 1898 are transferred to Diekeana Tomaszewska and Szawaryn 2015 (comb. nov.); Epilachna fulvohirta Weise 1895, E. nigrolimbata Thomson 1875, Henosepilachna griveaudi Chazeau 1975, H. vadoni Chazeau 1976, Solanophila consignata Weise 1909, S. coquereli Sicard 1907, and S. gyldenstolpei Weise 1924 are transferred to Cleta Mulsant 1850 (comb. nov.); Afidenta janczyki Fürsch 1986, Epilachna capicola Mulsant 1850, E. godarti Mulsant 1850, E. scitula Weise 1898, Henospeilachna acervata Chazeau 1975, and Solanophila blaesa Weise 1905 are transferred to Afidentula Kapur 1958 (com. nov.); Coccinella elaterii Rossi 1794, C. hirta Thunberg 1781, C. pavonia Olivier 1808, Epilachna annulata Kolbe 1897, E. biplagiata Kolbe 1897, E. cinerascens Weise 1907, E. connectens Weise 1912, E. erichi Weise, 1897, E. occellata Bertoloni, 1849, E. pauli Weise, 1897, E. tetracycla Gerstaecker, 1871, E. umbratilis Weise 1909, E. vulgaris Weise 1901, Henosepilachna bigemmata Fürsch 1991, Solanophila guttifera Weise 1899, S. hova Weise 1905, and S. kaffaeensis Weise 1906 are transferred to Chnootriba Chevrolat 1837 (com. nov.); Coccinella guttatopustulata Fabricius 1775, Epilachna aruensis Crotch 1874, E. biroi Weise 1902, E. buqueti Montrouzier 1861, E. fulvimana Weise 1903, E. immaculata Bielawski 1963, E. karapensis Bielawski 1963, E. orrori Bielawski 1963, E. samuelsoni Jadwiszczak 1991, and E. slipinskii Jadwiszczak 1987 are transferred to Papuaepilachna Tomaszewska and Szawaryn, 2013 (comb. nov.). The history of classification, the known aspects of the biology and distributional data of the tribe are summarized.