Phylogeny of the tachyporine group subfamilies and 'basal' lineages of the Aleocharinae (Coleoptera: Staphylinidae) based on larval and adult characteristics

Abstract. This paper reports the conclusions of studies into the phylogeny of tachyporine group subfamilies and the ‘basal’ lineages of the subfamily Aleocharinae (Coleoptera: Staphylinidae) based on both larval and adult morphological data (133 adult characters, twenty-seven larval characters). Representatives of forty species of the tachyporine group were used in the analysis, including representatives of the Aleocharinae, Trichophyinae, Habrocerinae, Phloeocharinae, Olisthaerinae, and Tachyporinae. The Aleocharinae included representatives of the tribes Gymnusini, Deinopsini, Mesoporini, the ‘subfamily’ Trichopseniinae, and representatives of nine major tribes in the ‘higher’ Aleocharinae (Athetini, Hoplandriini, Falagriini, Lomechisini, Oxypodini, Aleocharini, Myllaenini, Homalotini, and Hypocyphtini). Analyses were performed first with adult characters alone and then with both larval and adult characters in a simultaneous analysis. The analysis based on adult characters produced eighty-five equally parsimonious trees (length = 499, consistency index = 42; retention index = 69). In the consensus tree, the Tachyporinae are not monophyletic, and the sister-group relationship between the Trichophyinae + Habrocerinae and the Aleocharinae is not resolved. The Aleocharinae are monophyletic, but, among the ‘basal’ Aleocharinae, the relationships of Gymnusini + Deinopsini, the Mesoporini, and the Trichopseniinae are unresolved. The combined adult and larval data, using Tachinus as the outgroup, produced six equally parsimonious trees (tree length = 588; consistency index = 43; retention index = 69). The strict consensus tree of the combined larval and adult data supports the following conclusions: (1) larval characters substantially stabilize the tree; (2) the subfamily Tachyporinae is not supported to be monophyletic; (3) the subfamilies Trichophyinae and Habrocerinae are sister groups, and together they are sister to the Aleocharinae; (4) the ‘basal’ Aleocharinae are not a monophyletic group, but the ‘higher’ Aleocharinae are monophyletic; (5) the sister group of the remaining Aleocharinae is a lineage made up of genera currently in the tribes Gymnusini and Deinopsini; (6) within the Gymnusini–Deinopsini lineage, the monophyly of the Gymnusini is weakly supported, but the monophyly of the Deinopsini is strongly supported; (7) the subfamily Trichopseniinae is strongly supported to be a member of the ‘basal’ Aleocharinae; (8) the Myllaenini are resolved well within the ‘higher’ Aleocharinae; (9) strong support for the monophyly of some tribes of ‘higher’ Aleocharinae suggests that morphological characters provide substantial phylogenetic signal for analysis of higher-level phylogeny of the Aleocharinae in spite of the preliminary nature of the analysis at this taxonomic level.     

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 .     

Mouthpart structure of Stylogymnusa subantarctica Hammond, 1975 (Coleoptera: Staphylinidae: Aleocharinae) with a reanalysis of the phylogenetic position of the genus

Structure, particularly of the mouthparts, of the unusual aleocharine staphylinid Stylogymnusa subantarctica Hammond was reevaluated. The mouthparts are described in detail, and drawings of their structure are provided. Several features were found to be misinterpreted in previous accounts of the structure of this aleocharine. In particular, the stylate structures found on the prementum are not a highly modified, hollow, stylate glossa, as had been previously suggested. Instead, these structures represent highly modified labial palpi; the segments are completely fused, all signs of typical setae of the labial palpi are missing, and the palpi are very long and slender. In addition, there is no sign that they are hollow. Small, slender, membranous lobes at the base of, and between, the palpi are interpreted as the true glossae (=ligula). Reinterpretation of these characters, and greatly improved phylogenetic techniques in the 20 years since previous phylogenies had been proposed, allowed testing of previous hypotheses of the phylogenetic position of Stylogymnusa and the relationships of the tribes Gymnusini and Deinopsini within the Aleocharinae. Phylogenetic analysis of 7 out‐group and 12 in‐group taxa, based on 84 characters and 231 character states, produced 3 equally most‐parsimonious trees (tree length = 253, C.I. = 0.557, R.I. =0.674, rescaled C.I. = 0.376). These trees provide strong support for a monophyletic lineage consbting of Stylogymnusa, Gymnusa and the Deinopsini but only weak support for a monophyletic Gymnusini (Gymnusa + Stylogymnusa). Bootstrap analysis based on 1000 resampling repetitions showed the following monophyletic clades to be well supported by the dataset: Aleocharinae (79%), Stylogymnusa+Gymnusa+ Deinopsini (99%), Deinopsis+Adinopsis (100%), and representatives of Myllaenini + Athetini + Aleocharini + Homalotini + Oxypodini (the “higher Aleocharinae”) (88%). Weakly supported clades in the bootstrap analysis were: Paraconosoma+ Gymnusini + Deinopsini (56%), Anacyptus+‘higher Aleocharinae’ (54%), and Athetini + Oxypodini + Homalotini + Aleocharini (52%).     

Phylogeny and classification of diving beetles in the tribe Cybistrini (Coleoptera, Dytiscidae, Dytiscinae)

Phylogenetic relationships among members of the diving beetle tribe Cybistrini (Coleoptera: Dytiscidae) were inferred from analysis of 47 adult and larval morphological characters and sequences from portions of the genes cytochrome oxidase I (COI) and II (COII), histone III (H3) and wingless. Thirty‐three species of Cybistrini were included, representing all genus‐groups except Regimbartina Chatanay and Megadytes (Bifurcitus) Brinck, and most historically recognized species groups and subgenera used in the tribe. Outgroups include six species from other tribes within Dytiscinae and Lancetinae. Analyses included parsimony analysis of the combined data, likelihood analysis of combined molecular data and partitioned Bayesian analysis of the combined data. Results indicate that Cybistrini is well supported as a monophyletic group. Within the tribe, all currently recognized genus groups were found to be monophyletic with the exception of Onychohydrus Schaum, which is paraphyletic with respect to Austrodytes Watts in the parsimony analysis, but monophyletic in the likelihood and Bayesian analyses, and Cybister sensu stricto, which is paraphyletic with respect to C. (Melanectes) Brinck and C. (Scaphinectes) Ádám in the parsimony analysis or only the latter in the likelihood and Bayesian analyses. Results also suggest that some, but not all, historically recognized species groups or subgenera in the large genus Cybister Curtis are monophyletic, and this is discussed and compared. To improve the classification, the name Sternhydrus Brinck is elevated from subgenus to genus rank ( new status ). Four subgenera in the genus Cybister are recognized: C. (Melanectes) Brinck, C. (Megadytoides) Brinck ( resurrected ), C. (Neocybister) Miller, Bergsten and Whiting ( new subgenus ) and C. (Cybister) Curtis. The following new synonyms are established: Trochalus Dejean ( new synonym ), and ScaphinectesÁdám = Cybister (Cybister) ( new synonym ). The Neotropical species Cybister parvus Trémouilles (not examined) apparently does not fit any historical or currently recognized genus‐group diagnosis in Cybistrini, so it is retained in Cybister but incertae sedis with respect to subgenus. In addition to classification, the evolution of the unique character combinations present in cybistrines are discussed. A key to the adults of genera and subgenera is presented.     

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.     

COLEOFASCICULUS GEN. NOV. (CYANOBACTERIA): MORPHOLOGICAL AND MOLECULAR CRITERIA FOR REVISION OF THE GENUS MICROCOLEUS GOMONT1

Species currently classified within the cyanobacterial genus Microcoleus were determined to fall into two distinct clades in a 16S rDNA phylogeny, one containing taxa within the Oscillatoriaceae, the other containing taxa within the Phormidiaceae. The two lineages were confirmed in an analysis of the 16S–23S internal transcribed spacer (ITS) region sequences and secondary structures. The type species for Microcoleus is M. vaginatus Gomont, and this taxon belongs in the Oscillatoriaceae. Consequently, Microcoleus taxa in the Phormidiaceae must be placed in separate genera, and we propose the new genus Coleofasciculus to contain marine taxa currently placed in Microcoleus. The type species for Coleofasciculus is the well‐studied and widespread marine mat‐forming species Microcoleus chthonoplastes (Mert.) Zanardini ex Gomont. Other characters separating the two families include type of cell division and thylakoid structure.     

A new sinistral species of the land-snail genus Satsuma (Pulmonata: Camaenidae) from China

ABSTRACT

The species of Satsuma are mostly endemic to East Asia except for one species distributed in Batan Island of the Philippines. More than 99% of the known species of this genus are endemic to the island environment. Only three species are currently known to occur on the mainland. Here we describe a new sinistral Satsuma species from China, Satsuma guandi n. sp. We studied the shell morphology and genital anatomy of the new species and reconstructed the molecular phylogeny of the genus based on partial nucleotide sequences of the mitochondrial gene cytochrome c oxidase subunit I and nuclear markers from the ribosomal RNA cistrons (the internal transcribed spacer and the external transcribed spacer regions). The new species differs from other sinistral Satsuma species by having a strongly angulated shell and an open dark brownish-red umbilicus. The new species also differed from all other sinistral congeners in details of its reproductive anatomy. The molecular analysis supports the validity of the new species within the genus Satsuma.     

TAXONOMY AND MOLECULAR PHYLOGENY OF THE RED ALGAL GENUS LENORMANDIA (RHODOMELACEAE,CERAMIALES)1

The genus Lenormandia Sonder is currently composed of nine species from Australia and New Zealand. Some of these are well known, but others are rare and ill defined. Material of all nine species has been examined and found to fall into three discrete morphological groups forming highly supported clades on analysis of 18S rDNA sequences. The first group contains four Australian‐endemic species and includes the type species L. spectabilis Sonder. Plants have a cleft apex that is not inrolled, a distinctive rhombic surface areolation pattern caused by a one‐ to two‐layered medulla of interlocking cells, lack pseudopericentral cells, and produce their reproductive structures on the blade surfaces. The type species of the genus Lenormandiopsis, L. latifolia (Harvey et Greville) Papenfuss, was found to belong to this group and is thus returned to Lenormandia where it was originally placed. Species falling into the other two groups are removed to new genera that are being described separately. One extremely rare species of Lenormandia from southwestern Australia is transferred to the delesseriacean genus Phitymophora.     

Redefinition of the neotropical genus Anthopterus (Ericaceae: Vaccinieae), including one new species

The taxonomic history of Anthopterus as it relates to Themistoclesia is briefly reviewed. Anthopterus is redefined to include several species formerly placed in Themistoclesia. Two subgenera are established within Anthopterus. Anthopterus costaricensis Luteyn is described as new, and the new combination Anthopterus pterotus (A. C. Sm.) Luteyn is made. A key to the genus, a list of the species currently accepted, and brief notes on distribution and frequency are provided. Most species of Anthopterus are rare and their habitats are endangered.     

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.     

Comparative morphology of the tentorium and hypopharyngeal–premental sclerites in sporophagous and non‐sporophagous adult Aleocharinae (Coleoptera: Staphylinidae)

We elucidate the configuration of the tentorium and the sclerites of the hypopharynx–prementum complex in selected spore‐ (pollen‐) and non‐spore‐feeding Aleocharinae (Staphylinidae) by presenting the first comparative 3D reconstructions of these structures for 19 staphylinoid beetle species (six outgroups, 13 Aleocharinae). General organization of the tentorium follows the groundplan previously proposed for adult Staphylinidae, although some taxa have reduced or lost the dorsal (all Aleocharinae studied, Agathidium mandibulare [Leiodidae]) or anterior (Omalium rivulare [Omaliinae], Anotylus sculpturatus [Oxytelinae]) tentorial arms. All species investigated have premental and hypopharyngeal sclerites that are partly homologizable across taxa. We clarified that Musculus praementopalpalis externus originates from the premental sclerite, resolving its unclear origin reported in our previous publications. Eight of 13 investigated Aleocharinae species are spore/pollen feeders, six obligatorily. Three of these six (Eumicrota, Gyrophaena fasciata, G. gentilis) have grinding pseudomolae and a fully developed hypopharyngeal suspensorium with posterior bridge and anterior elongations; the remaining three (Oxypoda, Pagla, Polylobus) lack pseudomolae and suspensorial bridge, but have the suspensorium elongated anteriorly. The dorsolateral side of the hypopharyngeal sclerite interacts with the pseudomola. Obligate sporophagy/pollinivory apparently arose at least three times in Aleocharinae, not always involving the pseudomola–hypopharynx grinding mechanism.     

Balliella,a new genus of ceramiaceae (Rhodophyta)

Detailed studies of various aspects of the morphology and reproduction ofAntithamnion subcorticatum Itono andAntithamnion crouanioides Itono show that these species are incorrectly referred to as the genusAntithamnion. To include these species, a new genus,Balliella, is proposed. In addition, some other algae possibly referable to this genus are briefly discussed. The genusBalliella is here recognized as a member of the new tribeDelesseriopsieae which is most nearly related to the tribe Antithamnieae in the Ceramiaceae. The tribe Delesseriopsieae is now recognized to include the two genera, i.e.,Delesseriopsis andBalliella.     

A new species of Myotis Kaup, 1829 (Chiroptera, Vespertilionidae) from Ecuador

A new species of bat in the genus Myotis (Vespertilionidae, Myotinae) is described from the Chocó ecoregion on the western slope of the Andes in northwestern Ecuador. The genus Myotis comprises a diverse group of small to large-sized vespertilionid bats distributed worldwide. Twelve South American species are recognized currently, 6 of which occur in Ecuador. Morphological relationships among the new species and the other 12 South American species of Myotis were examined using 15 cranial and 5 external characters. Both qualitative and quantitative analyses found the new species to be distinct. As an aid to future identifications, we provide a key to the Ecuadorian species of Myotis.     

A new species of Mesopropithecus (Primates, Palaeopropithecidae) from Northern Madagascar

We diagnose new subfossil specimens belonging to the extinct palaeopropithecid, Mesopropithecus,from several caves at the Ankarana massif in northern Madagascar. They include three partial crania with mandibles, and the first vertebrae, ribs, clavicle, pelvis, ulna, and hand and foot bones of Mesopropithecusever found. Several other postcranial elements are known for this northern variant of Mesopropithecusand for previously described species from central and southern Madagascar. The new materials cannot be accommodated in either currently recognized species of this genus and are,hereby, given the name Mesopropithecus dolichobrachion. M. dolichobrachionis the second new species of extinct subfossil lemur to be found in the caves of Ankarana. It is unlike the two previously described species of this genus primarily in its larger size, different limb proportions, and aspects of its long bone morphology. Most notable is the fact that this is the only species of Mesopropithecuswith the forelimb longer than the hind limb— hence the name, the “long-armed” Mesopropithecus.Various characteristics place M. dolichobrachionphenetically closest (of species belonging to Mesopropithecus)to Babakotia radofilai, Archaeoindris,and Palaeopropithecus.If these similarities are derived specializations rather than primitive for palaeopropithecids, a new generic name will be required.     

The genus Asiabregma Belokobylskij,Zaldivar & Maeto (Hymenoptera: Braconidae) from China,with description of a new species

The small Asian genus Asiabregma Belokobylskij, Zaldivar & Maeto 2008 is recorded from China for the first time and one new species, A. achterbergi sp. nov., is described and illustrated. A key to species of genus Asiabregma is updated to include A. achterbergi. The relationship between Asiabregma and other members of the subtribe Facitorina and the systematic position of Facitorina are discussed.     

What is Geosaurus? Redescription of Geosaurus giganteus (Thalattosuchia: Metriorhynchidae) from the Upper Jurassic of Bayern,Germany

The holotype and referred specimens of Geosaurus giganteus, a metriorhynchid crocodile from the Tithonian (Upper Jurassic) of Germany, is redescribed, along with a historical overview of the genus and species. This taxon is unique among metriorhynchids as its serrated, strongly lateromedially compressed dentition is arranged as opposing blades, suggesting it was adapted to efficiently slice through fleshy prey. A new phylogenetic analysis of Crocodylomorpha is presented, which finds G. giganteus to be nested within what is currently considered Dakosaurus, whereas the other species currently assigned to Geosaurus form a clade with Enaliosuchus and the holotype of Cricosaurus. The phyletic relationship of G. giganteus with other metriorhynchids indicates that the current definition of the genus Geosaurus is polyphyletic, and that the inclusion of subsequent longirostrine species to this genus is in error. The re‐analysis presented herein demonstrates Geosaurus to be composed of three species sensu stricto. The appropriate taxonomic amendments to the Metriorhynchidae are also provided. © 2009 The Linnean Society of London, Zoological Journal of the Linnean Society, 2009, 157 , 551–585.     

Furcarcturus polarsterni gen. nov., sp. nov., a large deep-sea arcturid isopod from the Drake Passage, with a preliminary molecular characterization

A new arcturid species of unusual size (> 6 cm) is described from a locality in the Drake Passage at 3720 m depth. Due to the unique combination of characters found in these isopods, the new genus Furcarcturus is proposed. Sequences of the 16S rRNA gene obtained from eight species indicate that the new species does not belong to groups of species currently placed in the genera Antarcturus and Chaetarcturus. More sequences are needed to corroborate arcturid phlylogeny. Accepted: 28 May 2000     

The phylogeny and classification of Embioptera (Insecta)

A phylogenetic analysis of the order Embioptera is presented with a revised classification based on results of the analysis. Eighty‐two species of Embioptera are included from all families except Paedembiidae Ross and Embonychidae Navás. Monophyly of each of the eight remaining currently recognized families is tested except Andesembiidae Ross, for which only a single species was included. Nine outgroup taxa are included from Blattaria, Grylloblattaria, Mantodea, Mantophasmatodea, Orthoptera, Phasmida and Plecoptera. Ninety‐six morphological characters were analysed along with DNA sequence data from the five genes 16S rRNA, 18S rRNA, 28S rRNA, cytochrome c oxidase I and histone III. Data were analysed in combined analyses of all data using parsimony and Bayesian optimality criteria, and combined molecular data were analysed using maximum likelihood. Several major conclusions about Embioptera relationships and classification are based on interpretation of these analyses. Of eight families for which monophyly was tested, four were found to be monophyletic under each optimality criterion: Clothodidae Davis, Anisembiidae Davis, Oligotomidae Enderlein and Teratembiidae Krauss. Australembiidae Ross was not recovered as monophyletic in the likelihood analysis in which one Australembia Ross species was recovered in a position distant from other australembiids. This analysis included only molecular data and the topology was not strongly supported. Given this, and because parsimony and the Bayesian analyses recovered a strongly supported clade including all Australembiidae, we regard this family also as monophyletic. Three other families – Notoligotomidae Davis, Archembiidae Ross and Embiidae Burmeister, as historically delimited – were not found to be monophyletic under any optimality criterion. Notoligotomidae is restricted here to include only the genus Notoligotoma Davis with a new family, Ptilocerembiidae Miller and Edgerly, new family, erected to include the genus Ptilocerembia Friederichs. Archembiidae is restricted here to include only the genera Archembia Ross and Calamoclostes Enderlein. The family group name Scelembiidae Ross is resurrected from synonymy with Archembiidae (new status) to include all other genera recently placed in Archembiidae. Embiidae is not demonstrably monophyletic with species currently placed in the family resolved in three separate clades under each optimality criterion. Because taxon sampling is not extensive within this family in this analysis, no changes are made to Embiidae classification. Relationships between families delimited herein are not strongly supported under any optimality criterion with a few exceptions. Either Clothodidae Davis (parsimony) or Australembiidae Ross (Bayesian) is the sister to the remaining Embioptera taxa. The Bayesian analysis includes Australembiidae as the sister to all other Embioptera except Clothididae, suggesting that each of these taxa is a relatively plesiomorphic representatative of the order. Oligotomidae and Teratembiidae are sister groups, and Archembiidae (sensu novum), Ptilocerembiidae, Andesembiidae and Anisembiidae form a monophyletic group under each optimality criterion. Each family is discussed in reference to this analysis, diagnostic combinations and taxon compositions are provided, and a key to families of Embioptera is included.