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.     

A review of the subgenus <Emphasis Type="Italic">Carpelimus</Emphasis> s. str. (Coleoptera,Staphylinidae) from tropical Africa

A review of the subgenus Carpelimus (s. str.) from tropical Africa is given. The subgenus includes 10 species. A new species C. uhligi sp. n. is described, neotype of Trogophloeus insularis, Kraatz 1858 is designated. T. aequithorax Bernhauer, 1932 is placed in synonymy with C. dieganus (Fauvel, 1904); T. oculatus Wollaston, 1865 and T. meridioafricanus Scheerpeltz, 1974, with C. insularis (Kraatz, 1858), T. rudebecki Scheerpeltz, 1974 with C. memnonius (Erichson, 1840); T. yemenicus Coiffait, 1981 with C. niloticus (Erichson, 1840); T. nigerrimus Coiffait, 1935 and T. mimus Cameron, 1945, with C. rufitarsis (Fauvel, 1907); and T. bredoi Bernhauer, 1943, with C. transmarinus (Fauvel, 1907). Lectotypes of T. aequithorax, T. calidus, T. nigerrimus, and T. bredoi are designated. Twenty species are transferred from the genus Carpelimus, and the following new combinations are formed: Thinodromus brincki Scheerpeltz, 1972; Th. montiumdraconis Scheerpeltz, 1974; Th. rhodesianus Scheerpeltz, 1974; and Th. sudanensis Scheerpeltz, 1974 comb. nn.). The initial generic placement of Carpelimus luzidus Cameron, 1944 is restored.     

Phylogeny of the Homalotina (Coleoptera: Staphylinidae: Aleocharinae) based on morphology

The aleocharine subtribe Homalotina Heer represents one of the most diverse lineages of Staphylinidae. Despite its wide distribution and diversity, the phylogenetic relationships of the subtribe remain poorly understood. Here, we present the first cladistic analysis of the Homalotina based on morphological data. The subtribe is hypothesized to be a monophyletic group consisting of seven genera (Anomognathus Solier, Cephaloxynum Bernhauer, Holisomimus Cameron, Homalota Mannerheim, Neomalota Cameron, Stenomastax Cameron, and Thecturota Casey). The dataset for phylogenetic analysis comprised 83 characters representing 245 character states derived from adult morphology. These data were analysed using equal weighting and implied weighting schemes (k = 1–6) and results support the monophyly of the subtribe based on two synapomorphic characters (complete postoccipital sutures on head, posterolateral margin of metacoxae with macrosetae) and three homoplastic characters (medial setae on prementum not extended to apex of ligula, medial setae on labium contiguous, posterolateral angle of elytron slightly sinuate). Generic relationships differ in each analysis within the Homalotina (EW, IW with k = 1, 2–4, 5–6) although there are some identical topologies among the IW trees. Clades A, B, C, D, H, J and G were resolved as monophyletic in all weighting regimes. The monophyly of the genera is relatively well supported except for the genera Homalota and Stenomastax. Homalota species were recovered in four independent clades (clade C, D, I, K) and the Stenomastax species were recovered in two independent lineages. Candidates for the possible new genera are discussed. We herein transfer Homalota flavomaculata Bernhauer to the genus Stenomastax, resulting in the new combination [Stenomastax flavomaculata (Bernhauer)]. Our preliminary character correlation tests using phylogenetic pairwise comparisons did not support the hypothesis of association between flattened body form, and subcortical habitat and anterior shift of antennal insertion in Homalotini.     

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 .     

Discovery of the Quedius antipodum Sharp larva from New Zealand: phylogenetic test of larval morphology for Staphylinini at the intratribal level (Coleoptera: Staphylinidae)

Quedius antipodum Sharp is an endemic species from New Zealand. Here we describe its larva, the first of the species‐rich group of the south temperate ‘Quedius’ spp. This finding throws light on the controversy between the conventional systematics of Quedius Stephens and newer phylogenetic analyses, both of which are based on non‐larval characters only. We compare the larva of Q. antipodum with those of the north temperate Quedius (Quediina), where it was traditionally placed, and with the known larvae of Amblyopinina, a group where Q. antipodum was placed by recent phylogenetic studies. Sister‐group relationships of Q. antipodum within the tribe Staphylinini are explored based on larvae by means of parsimony analysis: 77 morphological characters scored for 20 species from 17 genera. Consistent with the adult morphology and DNA sequences, larvae‐based cladistic analysis confirms that Q. antipodum should not be placed in the north temperate genus Quedius. However, larval analysis alone remains dubious with respect to finding the exact sister relationships of that species.     

Alesiella gen.n. and a newly discovered relict lineage of Staphylinini (Coleoptera: Staphylinidae)

This study aimed to identify the sister group of the poorly known and morphologically isolated Burmese species Quedius lineipennis within the tribe Staphylinini (Staphylinidae: Staphylininae) using morphological characters. Phylogenetic analysis of a broad taxon sample demonstrated that this Asian species is not a member of the genus Quedius but forms the sister taxon to the Neotropical genus Quediomacrus. Both taxa were shown to be members of a hitherto unrecognized lineage with a highly disjunct distribution. The lineage is hypothesized to be Asian in origin, with dispersals to the Americas during the early Eocene climatic maximum via Beringia and to Australia via land connections in the late Miocene. The current distribution of the lineage is considered to be relictual. New phylogenetic hypotheses within ‘Quediina’ and Staphylinini as a whole are proposed and the general tree topology of Staphylinini recovered by recent morphological studies is refined. Phylogenetic relationships within the Quedius complex remain unclear. Alesiella Brunke and Solodovnikov gen.n. , is erected for Quedius lineipennis and the Quedius subgenus Quedionuchus is reinstated to genus level with new combinations as follows: Quedionuchus atl (Smetana, 1975), comb.n. ; Quedionuchus calli (Smetana, 1976), comb.n. ; Quedionuchus cipactli (Smetana, 1976), comb.n. ; Quedionuchus coatl (Smetana, 1976), comb.n. ; Quedionuchus ehacatl (Smetana, 1976), comb.n. ; Quedionuchus ollin (Smetana, 1976), comb.n. ; Quedionuchus ozomatli (Smetana, 1975), comb.n. ; Quedionuchus reitterianus (Bernhauer, 1944), comb.n. ; Quedionuchus samuraicus (Bernhauer and Schubert, 1916), comb.n. ; Quedionuchus schultzei (Smetana, 1975), comb.n. ; Quedionuchus tecpatl (Smetana, 1976), comb.n. ; Quedionuchus xochitl (Smetana, 1976). Quedius lugubris Lokay, 1913 is transferred from the subgenus Quedionuchus to the subgenus Distichalius and placed in synonymy: Quedius punctatellus (Heer, 1839) = Quedius lugubris Lokay, 1913, syn.n . This published work has been registered in ZooBank, http://zoobank.org/urn:lsid:zoobank.org:pub:9C7D3C90‐FCB9‐414F‐911B‐194A1A6602DE .     

Phylogeny of xanthopygine rove beetles (Coleoptera) based on six molecular loci

The rove beetle subtribe Xanthopygina (Coleoptera: Staphylinidae: Staphylininae: Staphylinini) is a species‐rich group of 27 neotropical genera that contains some of the largest and most brightly coloured of all staphylinid beetles. The monophyly of the subtribe has never been tested before, using a large dataset of taxa and genes. Bayesian and maximum likelihood analyses are used on individual genes (COI, 28S rDNA, wingless, arginine kinase, CAD and topoisomerase I) and the partitioned concatenated dataset to test for monophyly and examine the relationships among Xanthopygina genera. Xanthopygina (excluding Philothalpus) are shown to be a monophyletic group with strong support values. The genus Philothalpus is removed from Xanthopygina and placed in the tribe Staphylinini as incertae sedis. Four distinct clades of Xanthopygina genera are recognized. The origin of Xanthopygina is hypothesized to be in the Late Cretaceous or later and the origin of myrmecophilous adaptations is discussed.     

Towards a natural classification of the subtribe Philonthina (Coleoptera: Staphylinidae: Staphylinini): a phylogenetic analysis of the Neotropical genera

Philonthina, the largest subtribe of the rove beetle tribe Staphylinini, is a hyperdiverse group in the Neotropical Region, accounting for about half of the genera of the subtribe. Despite such diversity, Neotropical Philonthina have never been analysed phylogenetically, deterring formulation of a modern classification of the Staphylinini. A cladistic analysis of Neotropical Philonthina was performed based on 110 morphological characters and 77 terminal taxa. Representatives of Philonthina from other regions and other main lineages of Staphylinini, Arrowinini and Platyprosopini were included to test their relationships with Neotropical Philonthina. The major results are the monophyly of 11 of the 17 endemic Neotropical genera of Philonthina, the placement of Holisus Erichson (Hyptiomina) into this clade showing a sister group relationship to myrmecophile genera, and the position of Erichsonius Fauvel outside of Philonthina within Staphylinini. Six of the current seven species of Endeius Coiffait & Sáiz group with Neotropical species of Philonthus Stephens. The separation of Gondwana about 65 my and major landscape modifications in the vast interior of northern South America during the past 25 my is proposed to explain the evolution of the endemic Neotropical genera of Philonthina. The following taxonomic changes are proposed: Erichsonius Fauvel, 1874 now placed as incertae sedis in Staphylinini; Endeius Coiffait & Sáiz, 1968, n.syn. of Philonthus Stephens, 1929 and Endeius nitidipennis (Solier, 1849) placed as incertae sedis in Philonthina. The following new combinations are proposed: Philonthus franzi (Sáiz, 1971), comb.n. , Philonthus loensis (Coiffait & Sáiz, 1968), comb.n. , Philonthus lugubris (Sáiz, 1971), comb.n. , Philonthus ovaliceps (Coiffait, 1981), comb.n. , Philonthus punctipennis (Solier, 1849), comb.res. and Philonthus subpunctipennis (Coiffait & Sáiz, 1968), comb.n. Philonthus herberti, n.nov., is proposed for Philonthus franzi Schillhammer, 1998 , which is a junior secondary homonym of Philonthus franzi (Sáiz, 1971).     

Molecular phylogeny of the mega‐diverse rove beetle tribe Staphylinini (Insecta,Coleoptera, Staphylinidae)

Chatzimanolis, S., Cohen, I. M., Schomann, A. & Solodovnikov, A. (2010). Molecular phylogeny of the mega‐diverse rove beetle tribe Staphylinini (Insecta, Coleoptera, Staphylinidae). —Zoologica Scripta, 39, 436–449. Phylogeny of the rove beetle tribe Staphylinini is explored by parsimony and Bayesian analyses of sequences of four genes (COI, wingless, Topoisomerase I, and 28S) for 43 ingroup (various genera of Staphylinini) and eight outgroup (two genera of Paederinae, six genera of other tribes of Staphylininae) taxa. Analyses were conducted for each gene independently and for the concatenated data set. Results of the most robust combined analyses were compared with the morphology‐based phylogenies of Staphylinini (‘test phylogeny’), and with the conventional classification of this tribe. Molecular results were congruent with the ‘test phylogeny’ in the following: ancestors of Staphylinini were ‘Quediina‐like’ lineages; formal subtribe Quediina mixes at least two relatively basal groups, ‘Quediina propria’ and ‘southern Quediina’; specialized subtribe Amblyopinina is an internal clade within ‘southern Quediina’; a relatively deeply nested ‘Staphylinini propria’ that unites current subtribes Staphylinina, Eucibdelina, Anisolinina, Xanthopygina and Philonthina is well supported as a monophyletic group. In strong contrast with morphology, molecular data place the tribes Othiini and Xantholinini nested within Staphylinini. Molecular results strongly conflict with morphology by uniting morphologically very different genera Holisus and Atanygnathus in one clade that has uncertain position within Staphylinini. Consistently with the most congruent areas of the morphology‐ and molecular‐based phylogenies, taxonomic changes are implemented for the formal subtribes Quediina and Amblyopinina.     

A revision of the Neotropical species of Bolitogyrus Chevrolat,a geographically disjunct lineage of Staphylinini (Coleoptera,Staphylinidae)

The Neotropical species of the rarely collected genus Bolitogyrus (Coleoptera: Staphylinidae: Staphylininae: Staphylinini) are revised. The genus exhibits an uncommon, disjunct distribution between the Neotropical and Oriental Regions and is of unknown phylogenetic position within Staphylinini. Morphological evolution remarkable for Staphylinini was discovered within Bolitogyrus, including sexually dimorphic modifications of the pronotum that may be involved in male competition for females. rSEM interactive animations were used to establish morphological species boundaries between two highly variable species and are provided to illustrate diagnostic characters of the genitalia in unconventional views. The genus is redescribed based on the world fauna and twenty-eight Neotropical species are considered valid. Of these, nineteen are described as new to science: Bolitogyrus ashei sp. n.; B. apicofasciatus sp. n.; B. brevistellus sp. n.; B. bufo sp. n.; B. cheungi sp. n.; B. cornutus sp. n.; B. divisus sp. n.; B. falini sp. n.; B. gracilis sp. n.; B. inexspectatus sp. n.; B. longistellus sp. n.; B. marquezi sp. n.; B. newtoni sp. n.; B. pseudotortifolius sp. n.; B. pulchrus sp. n.; B. silex sp. n.; B. thomasi sp. n.; B. tortifolius sp. n.; and B. viridescens sp. n. Bolitogyrus sallei (Kraatz), stat. r. is removed from synonymy with B. buphthalmus (Erichson) and the following new synonyms are proposed: Cyrtothorax cyanescens Sharp, 1884, syn. n. = Quedius buphthalmus Erichson, 1840; C. nevermanni Scheerpeltz, 1974, syn. n. = C. costaricensis Wendeler, 1927. A summary of all available bionomic and distributional data, as well as an illustrated identification key to and diagnoses of all Neotropical species are provided.     

Rove beetle subtribes Quediina, Amblyopinina and Tanygnathinina: systematic changes affecting Central European fauna (Coleoptera, Staphylinidae, Staphylinini)

In preparation for the new edition of the identification keys of rove beetles of Central Europe (Volume 4 of the “Die Käfer Mitteleuropas”), the following systematic problems affecting the Central European fauna of the tribe Staphylinini are addressed: phylogeny-based, new concepts for the subtribes Quediina and Amblyopinina; status of the subtribe Tanygnathinina; systematic position of the genus Astrapaeus; status of Quedionuchus, the subgenus of Quedius; identity of some species of Quedius and Heterothops. As a result, new wordwide and Central Europe-based diagnoses are given for the subtribes Quediina and Amblyopinina; earlier recognized but not widely accepted synonymies of the genera Quedius and Velleius, and of the species Heterothops praevius and Heterothops niger, are justified; new synonyms are established for: Quedius pseudonigriceps Reitter, 1909 (= Quedius noricus Bernhauer, 1927, syn. n.); Quedius maurorufus (Gravenhorst, 1806) (= Quedius richteri Korge, 1966, syn. n.); Quedius suturalis Kiesenwetter, 1845 (= Quedius merlini Drugmand & Bruge 1991, syn. n.); lectotypes are designated for Quedius meridiocarpathicus Smetana, 1958, Quedius noricus Bernhauer, 1927, and Quedius pseudonigriceps Reitter, 1909. As a result of synonymy of Quedius and Velleius, the following new combinations are proposed: Quedius amamiensis (Watanabe, 1990), comb. n.; Quedius circumipectus (Cho, 1996), comb. n.; Quedius elongatus (Naomi, 1986), comb. n.; Quedius japonicus (Watanabe, 1990), comb. n.; Quedius pectinatus (Sharp, 1874), comb. n.; Quedius setosus (Sharp, 1889), comb. n.; Quedius simillimus (Fairmaire, 1891), comb. n. As a result of new combinations, Quedius japonicus (Watanabe, 1990) (non Quedius japonicus Sharp, 1874) is replaced with the new name Quedius watanabei Solodovnikov, nom. n., while Quedius pectinatus Lea, 1908 (non Quedius pectinatus (Sharp, 1874)) is replaced with the new name Quedius arthuri Solodovnikov, nom. n.     

Revised concept of the genus Euryporus Erichson (Coleoptera, Staphylinidae, Staphylininae) and phylogenetic significance of Staphylinini from New Guinea

The Staphylinini rove beetle genus Euryporus Erichson from the subtribe Quediina is restricted to include only three species from the Western Palearctic region: Euryporus picipes (Paykull, 1800), Euryporus aeneiventris (Lucas, 1846), and Euryporus princeps Wollaston, 1864. Euryporus argentatus Fauvel, 1881, Euryporus warisensis Last, 1987 and Euryporus multicavus Last, 1980, which do not even belong to the subtribe Quediina, are excluded fromthe genus. Of these, two were transferred to different genera: Tympanophorus argentatus (Fauvel, 1881), comb. nov., from Sumatra;and Hesperus warisensis (Last, 1987), comb. nov.,from New Guinea. "Euryporus" multicavus could not be placed to any of the described genera of Staphylinini and is left as incertae sedis pending a broader study of the relevant fauna of this tribe in New Guinea and adjacent regions. The taxonomic history of Euryporus is reviewed, and an updated diagnosis of this genus is provided.     

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.     

Phylogenetic relationships among diploid species of Symphyotrichum (Asteraceae: Astereae) based on two nuclear markers, ITS and GAPDH

The mostly North American subtribe Symphyotrichinae (Asteraceae: Astereae) comprises Canadanthus, Ampelaster, Psilactis, Almutaster, and Symphyotrichum. Intergeneric and interspecific relationships within the subtribe have been investigated in the past, particularly by Nesom [Nesom, G.L., 1994. Review of the taxonomy of Aster sensu lato (Asteraceae: Astereae), emphasizing the new world species, Phytologia 77, 141–297] and Semple [Semple, J.C., 2005. Classification of Symphyotrichum. Available from: <http://www.jcsemple.uwaterloo.ca/Symphyotrichumclassification.htm/>], using morphological and cytological approaches. Symphyotrichum is the largest and most complex genus within the subtribe and includes four subgenera: Symphyotrichum (x = 7, 8), Virgulus (x = 4, 5), Astropolium (x = 5), and Chapmaniani (x = 7). In this study we used two nuclear markers, the nrDNA internal transcribed spacer (ITS) and the low-copy nuclear gene glyceraldehyde 3-phosphate dehydrogenase (GAPDH), to resolve intergeneric and interspecific relationships within the subtribe at the diploid level, and to determine whether our phylogenies validate the classifications of Nesom or Semple. Our results confirm the distinct generic status of Canadanthus and Ampelaster, whereas Psilactis and Almutaster form a polytomy with Symphyotrichum. Within Symphyotrichum, subg. Virgulus is monophyletic based on ITS and appears polyphyletic based on GAPDH. Neither the ITS nor the GAPDH analyses support a distinct status for subg. Astropolium, which groups within subg. Symphyotrichum. In general, interspecific relationships within Symphyotrichum are unresolved. Lack of resolution may be interpreted as a case of recent and rapid evolutionary radiation.     

Rove beetles of the subtribe Scopaeina Mulsant & Rey (Coleoptera: Staphylinidae) in the West Palaearctic: Phylogeny, biogeography and species catalogue

A cladistic analysis of the West Palaearctic Scopaeina Mulsant & Rey, 1878 (Coleoptera, Staphylinidae: Paederinae) is presented along with bionomic and biogeographic information. A total of 76 morphological characters were coded for the 88 currently known West Palaearctic species, except for S. bifossicapitata (Outerelo & Oromi, 1987). Results show that Scopaeina comprises two well-supported monophyletic groups in the West Palaearctic, Micranops Cameron, 1913 and Scopaeus Erichson, 1840, which are considered to represent distinct genera. Phylogenetic relationships to Orus Casey, 1884, distributed in North and South America, are briefly discussed. Whereas Micranops is only represented by M. pilicornis (Baudi, 1869) in the region under study, 87 species of Scopaeus are currently known from the West Palaearctic. Within Scopaeus, the cladistic analysis yielded many well-supported monophyletic species groups, most of which are restricted to the West Palaearctic. However, except for Hyperscopaeus Coiffait, 1984, they are not in agreement with the widely used subgeneric concept sensu Coiffait (1952–1984). The following polyphyletic subgenera are consequently synonymized: Alloscopaeus Coiffait, 1968, Anomoscopaeus Coiffait, 1968, Geoscopaeus Coiffait, 1960, and Hyposcopaeus Coiffait, 1960 synn. n. = Scopaeus Erichson, 1840. Nivorus Herman, 1965, and Microscopaeus Coiffait, 1981 synn. n. = Micranops Cameron, 1913. The monotypical genus Coecoscopaeus Coiffait, 1984, established for C. coecus (Peyerimhoff, 1906), is excluded from Scopaeina. Scopaeus mitratus perroti Ochs, 1953 is raised to species rank, and S. nigellus Wollaston, 1864, formerly a synonym of S. minimus Erichson, 1939, is revalidated. Finally, we present a catalogue of species and synonyms of West Palaearctic Scopaeina along with distributional data and five new synonymies of species group names: S. bordei Peyerimhoff, 1914 syn. n. = S. portai Luze, 1910; S. tassiliensis Jarrige, 1958, S. mauretanicus Coiffait, 1960 synn. n. = S. crassipes Wollaston, 1867; S. saoudiensis Coiffait, 1981 = S. sinaicus Coiffait, 1970; S. mateui Coiffait, 1953 syn. n. = S. didymus Erichson, 1840. A lectotype is designated for S. didymus Erichson, 1840.See also Electronic Supplement (Parts 13) at http://www.senckenberg.de/odes/02-02.htm     

A review of the <Emphasis Type="Italic">Thinodromus lunatus</Emphasis> species-group (Coleoptera,Staphylinidae)

The Thinodromus lunatus species group is revised. The following new species are described: Thinodromus (s. str.) cattiensis sp. n. from Vietnam, Thinodromus (s. str.) forsteri sp. n. from southern Thailand, Thinodromus (s. str.) himalayensis sp. n. from Nepal and northern India, Thinodromus (s. str.) inconspicuus sp. n. from southern China, Thailand, and Vietnam, and Thinodromus (s. str.) spotus sp. n. from southern China. The following new synonymy is established: Thinodromus (s. str.) deceptor (Sharp, 1889) = Thinodromus (s. str.) gravelyi (Bernhauer, 1926), syn. n.; = Thinodromus (s. str.) reitterianus (Bernhauer, 1938), syn. n. Lectotypes are designated for Trogophloeus lunatus Motschulsky, 1857, Trogophloeus pustulatus Bernhauer, 1904, Trogophloeus socius Bernhauer, 1904, Trogophloeus sumatrensis Bernhauer, 1915, Trogophloeus lewisi Cameron, 1919, Trogophloeus gravelyi Bernhauer, 1926, Trogophloeus reitterianus Bernhauer, 1938, and Trogophloeus unipustulatus Cameron, 1941. A key is presented to all the species of the Thinodromus lunatus group.     

Phylogenetics and biogeography of a spectacular Old World radiation of butterflies: the subtribe Mycalesina (Lepidoptera: Nymphalidae: Satyrini)

Background  

Butterflies of the subtribe Mycalesina (Nymphalidae: Satyrinae) are important model organisms in ecology and evolution. This group has radiated spectacularly in the Old World tropics and presents an exciting opportunity to better understand processes of invertebrate rapid radiations. However, the generic-level taxonomy of the subtribe has been in a constant state of flux, and relationships among genera are unknown. There are six currently recognized genera in the group. Mycalesis, Lohora and Nirvanopsis are found in the Oriental region, the first of which is the most speciose genus among mycalesines, and extends into the Australasian region. Hallelesis and Bicyclus are found in mainland Africa, while Heteropsis is primarily Madagascan, with a few species in Africa. We infer the phylogeny of the group with data from three genes (total of 3139 bp) and use these data to reconstruct events in the biogeographic history of the group.     

Sequence analysis and functional characterization of the promoter of the PiceaglaucaCinnamylAlcoholDehydrogenase gene in transgenic white spruce plants

The enzyme Cinnamyl Alcohol Dehydrogenase (CAD) catalyses the last step of lignin monomer synthesis, and is considered as a molecular marker of cell wall lignification in different plants species. Here, we report the isolation and analysis of 5′ flanking genomic DNA regions upstream to the CAD gene, from two conifers, i.e. white spruce (Picea glauca (Moench) Voss) and loblolly pine (Pinus taeda L.). Sequence comparisons with available CAD gene promoters from angiosperms highlighted the conservation of cis-elements matching MYB, WRKY and bHLH binding sites. Functional characterization of the P. glauca CAD promoter used P. glauca seedlings stably transformed with a DNA fragment of 1,163 base pairs (PgCAD) fused to the β-glucuronidase (GUS) gene. Histochemical observations of different vegetative organs of the transgenic trees showed that this sequence was sufficient to drive GUS expression in lignifying tissues, and more specifically in differentiating xylem cells. Quantitative RT-PCR experiments also indicated that the native CAD gene was preferentially expressed in differentiating xylem both in stems and roots. In addition, GUS expression driven by the PgCAD promoter was wound-inducible which was consistent with the accumulation of CAD mRNA in response to jasmonate application and mechanical wounding. The spruce CAD promoter represents a valuable tool for research and biotechnology applications related to xylem and wood. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.