Phylogeny and historical biogeography of leafhopper subfamily Iassinae (Hemiptera: Cicadellidae) with a revised tribal classification based on morphological and molecular data

Phylogenetic relationships among major lineages of the leafhopper subfamily Iassinae were explored by analysing a dataset of 91 discrete morphological characters and DNA sequence data from nuclear 28S rDNA and histone H3 genes and mitochondrial 12S rDNA. Bayesian, maximum‐likelihood and maximum parsimony analyses yielded similar tree topologies that were well resolved with strong branch support except at the base of the tree, resulting in equivocal support for inclusion of Bythoniini as a tribe of Iassinae but strong support for the monophyly of Iassinae (excluding Bythoniini) and most previously recognized iassine tribes. Divergence times for recovered nodes were estimated using a Bayesian relaxed clock method with two fossil calibration points. The results suggest that the deepest divergences coincided with Gondwanan vicariant events but that more recent divergences resulted from long‐range dispersal and colonization. Biogeographical analyses suggest that the group most likely has a Neotropical origin. The following changes to the taxonomic classification are proposed: establishment of three new tribes, Batracomorphini trib.n. (based on type genus Batracomorphus Lewis), Hoplojassini trib.n. (based on type genus Hoplojassus Dietrich and including one other South American genus), Lipokrisnini trib.n. (based on type genus Lipokrisna Freytag and including two other endemic Caribbean genera); Krisnini is redefined to include only the Old World genera Krisna and Gessius; Iassini is redefined to include only the type genus and four endemic Afrotropical genera; Bascarrhinus Fowler and Platyhynna Berg, recently treated as genera incertae sedis, are placed in Hyalojassini; Thalattoscopus Kirkaldy is added to the previously monobasic tribe Trocnadini. Iassinae now includes 12 tribes, all of which appear to be monophyletic. Revised morphological diagnoses of the subfamily and each of the included tribes are provided and a key to tribes is also given. This published work has been registered in ZooBank, http://zoobank.org/urn:lsid:zoobank.org:pub:41295B68‐2DAB‐4C4F‐B260‐F7C054922173 .     

A molecular phylogeny of the tribe Aphidini (Insecta: Hemiptera: Aphididae) based on the mitochondrial tRNA/COII, 12S/16S and the nuclear EF1α genes

Abstract A phylogeny of the tribe Aphidini (Hemiptera: Aphididae) was reconstructed from three gene fragments: two mitochondrial regions, partial tRNA‐leucine + cytochrome oxidase II (tRNA/COII), partial 12S rRNA + tRNA‐valine + 16S rRNA (12S/16S) and one nuclear gene, the elongation factor‐1 alpha (EF1α). Bayesian phylogenetic (BP) analyses were performed on each individual dataset of tRNA/COII, 12S/16S and EF1α, and maximum parsimony (MP), Bremer support test, maximum likelihood (ML) and BP analysis were performed on the combined dataset. After comparing our molecular phylogenetic results with the classic classification based on morphological and ecological data, we analysed three main issues: the monophyletic relationships among tribes and subtribes, the validities of the latest taxonomic positions of genera and species and the status of certain Aphis species groups. Our results indicate that 36 of the species analysed, with the exception of Cryptosiphum artemisiae, are clustered within the clade of Aphidini. Also, the 28 species representative of the subtribe Aphidina were separated from the eight species representative of Rhopalosiphina; each monophyletic subtribe was supported by significant P‐values in the combined analysis. According to our results, Cryptosiphum should be moved to Macrosiphini because it is more closely related to the genera Lipaphis and Brevicoryne. The genus Toxoptera was recovered as non‐monophyletic. In Rhopalosiphina, three genera, Hyalopterus, Rhopalosiphum and Schizaphis, were relatively closer to each other than to the genus Melanaphis. In the relationships between species‐groups among Aphis, most species were separated into two main lineages; the fabae group seemed to be more closely related to the spiraecola and craccivora group rather than to the gossypii group.     

Molecular phylogenetics and generic assessment in the tribe Morindeae (Rubiaceae–Rubioideae): How to circumscribe Morinda L. to be monophyletic?

Most of the species of the family Rubiaceae with flowers arranged in head inflorescences are currently classified in three distantly related tribes, Naucleeae (subfamily Cinchonoideae) and Morindeae and Schradereae (subfamily Rubioideae). Within Morindeae the type genus Morinda is traditionally and currently circumscribed based on its head inflorescences and syncarpous fruits (syncarps). These characters are also present in some members of its allied genera, raising doubts about the monophyly of Morinda. We perform Bayesian phylogenetic analyses using combined nrETS/nrITS/trnT-F data for 67 Morindeae taxa and five outgroups from the closely related tribes Mitchelleae and Gaertnereae to rigorously test the monophyly of Morinda as currently delimited and assess the phylogenetic value of head inflorescences and syncarps in Morinda and Morindeae and to evaluate generic relationships and limits in Morindeae. Our analyses demonstrate that head inflorescences and syncarps in Morinda and Morindeae are evolutionarily labile. Morinda is highly paraphyletic, unless the genera Coelospermum, Gynochthodes, Pogonolobus, and Sarcopygme are also included. Morindeae comprises four well-supported and morphologically distinct major lineages: Appunia clade, Morinda clade (including Sarcopygme and the lectotype M. royoc), Coelospermum clade (containing Pogonolobus and Morinda reticulata), and Gynochthodes–Morinda clade. Four possible alternatives for revising generic boundaries are presented to establish monophyletic units. We favor the recognition of the four major lineages of Morindeae as separate genera, because this classification reflects the occurrence of a considerable morphological diversity in the tribe and the phylogenetic and taxonomic distinctness of its newly delimited genera.     

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 .     

A comprehensive molecular phylogeny of tiger beetles (Coleoptera,Carabidae, Cicindelinae)

Tiger beetles are a remarkable group that captivates amateur entomologists, taxonomists and evolutionary biologists alike. This diverse clade of beetles comprises about 2300 currently described species found across the globe. Despite the charisma and scientific interest of this lineage, remarkably few studies have examined its phylogenetic relationships with large taxon sampling. Prior phylogenetic studies have focused on relationships within cicindeline tribes or genera, and none of the studies have included sufficient taxon sampling to conclusively examine broad species patterns across the entire subfamily. Studies that have attempted to reconstruct higher‐level relationships of Cicindelinae have yielded conflicting results. Here, we present the first taxonomically comprehensive molecular phylogeny of Cicindelinae to date, with the goal of creating a framework for future studies focusing on this important insect lineage. We utilized all available published molecular data, generating a final concatenated dataset including 328 cicindeline species, with molecular data sampled from six protein‐coding gene fragments and three ribosomal gene fragments. Our maximum‐likelihood phylogenetic inferences recover Cicindelinae as sister to the wrinkled bark beetles of the subfamily Rhysodinae. This new phylogenetic hypothesis for Cicindelinae contradicts our current understanding of tiger beetle phylogenetic relationships, with several tribes, subtribes and genera being inferred as paraphyletic. Most notably, the tribe Manticorini is recovered nested within Platychilini including the genera Amblycheila Say, Omus Eschscholtz, Picnochile Motschulsky and Platychile Macleay. The tribe Megacephalini is recovered as paraphyletic due to the placement of the monophyletic subtribe Oxycheilina as sister to Cicindelini, whereas the monophyletic Megacephalina is inferred as sister to Oxycheilina, Cicindelini and Collyridini. The tribe Collyridini is paraphyletic with the subtribes Collyridina and Tricondylina in one clade, and Ctenostomina in a second one. The tribe Cicindelini is recovered as monophyletic although several genera are inferred as para‐ or polyphyletic. Our results provide a novel phylogenetic framework to revise the classification of tiger beetles and to encourage the generation of focused molecular datasets that will permit investigation of the evolutionary history of this lineage through space and time.     

Phylogeny of a new supertribe Cephenniitae with generic review of Eutheiini and description of a new tribe Marcepaniini (Coleoptera: Staphylinidae: Scydmaeninae)

Genera of Eutheiini are reviewed and Eutheimorphus is removed from this tribe of ant‐like stone beetles (Scydmaeninae) and transferred to Cephenniini. A monogeneric Marcepaniini trib.n. is described to accommodate Marcepania gen.n. from Malaysia, with five species: M. semengohensis sp.n. (the type species of Marcepania), M. tuberculata sp.n. , M. seramaensis sp.n. , M. minutissima sp.n. and M. elongata sp.n. A phylogenetic analysis of all genera of Cephenniini, Eutheiini and Marcepaniini based on adult morphological characters resulted in recovering a well‐supported monophyletic clade Eutheiini + (Marcepaniini + Cephenniini) and these tribes are included in Cephenniitae stat.n. (Eutheiini and Cephenniini are therefore removed from Scydmaenitae). Only a weak support for monophyly of Eutheiini was found, but morphological characters allow for maintaining this presumably relic group as a separate tribe. Previously proposed monophyletic groups within Cephenniini were recovered as such, but after inclusion of Eutheimorphus, a sister taxon to the ‘Cephennomicrus group’, the latter lineage gained weak statistical support. The evolutionary history of Cephenniitae is discussed, with focus on known northern hemisphere fossils classified in Scydmaenitae and Hapsomelitae, but possibly closely allied to Cephenniitae. Establishing the supertribe Cephenniitae is the first step toward a profound reclassification of Scydmaeninae on a robust phylogenetic basis. This published work has been registered in ZooBank, http://zoobank.org/urn:lsid:zoobank.org:pub:B0E1B12D-9587-4C4F-A908-A12A0C424A8C .     

Phylogeny of Mesoamerican freshwater mussels and a revised tribe‐level classification of the Ambleminae

Evolutionary and ecological hypotheses of the freshwater mussel subfamily Ambleminae are intensely geographically biased—a consequence of the complete exclusion of Mesoamerican taxa in phylogenetic reconstructions of the clade. We set out to integrate a portion of the Mesoamerican freshwater mussel assemblage into existing hypotheses of amblemine classification and evolution by generating a molecular phylogeny that includes four previously unsampled Mesoamerican genera and nine species endemic to that region. Given the traditionally hypothesized affinity to Nearctic mussels and the understanding that classification should reflect common ancestry, we predicted that (a) Mesoamerican genera would be recovered as members of the recognized tribes of the Ambleminae, and (b) genera would be supported as monophyletic. The mutilocus phylogeny (COI + 28S + 16S) reported herein does not fully support either of those hypotheses. Neither Cyrtonaias nor Psorula were supported as monophyletic and we predict several other Mesoamerica genera are also non‐monophyletic. The reconstructed phylogeny recovered four independent lineages of Mesoamerican freshwater mussels and these clades are distributed across the phylogeny of the Ambleminae, including the tribe Quadrulini (Megalonaias), Lampsilini (two lineages: Cyrtonaias explicata/Sphenonaias microdon, and Pachynaias), and a previously unrecognized, exclusively Mesoamerican and Rio Grande clade consisting of the genera Psoronaias, Psorula and Popenaias. The latter clade possesses several morphological characteristics that distinguish it from its sister taxon, tribe Lampsilini, and we recognize this newly identified Mesoamerican clade as a fifth tribe of the Ambleminae attributable to the Popenaiadini Heard & Guckert, 1970. This revised classification more completely recognizes the suprageneric diversity of the Ambleminae.     

Mitochondrial phylogeny of Arvicolinae using comprehensive taxonomic sampling yields new insights

Comprehensive taxonomic sampling can vastly improve the accuracy of phylogenetic reconstruction. Here, we present the most inclusive phylogenetic analysis of Arvicolinae (Mammalia, Rodentia) to date, combining all published cytochrome  b gene sequences of greater than 1097 bp and new sequences from two monotypic genera. Overall, the phylogenetic relationships between 69 species of voles and lemmings, representing 18 genera and 10 tribes, were studied. By applying powerful modern approaches to phylogenetic reconstruction, such as maximum likelihood and Bayesian analysis, we provide new information on the early pulse of evolution within the Arvicolinae. While the position of two highly divergent lineages, Phenacomys and Ondatra , could not be resolved, the tribe Lemmini, appeared as the most basal group of voles. The collared lemmings (Dicrostonychini) grouped together with all of the remaining tribes. The two previously unstudied monotypic genera Dinaromys and Prometheomys form a moderately well-supported monophyletic clade, possibly a sister group to Ellobius (Ellobiusini). Furthermore, with one exception, all tribes ( sensu Musser & Carleton, 2005) proved to be monophyletic and can thus be regarded as meaningful evolutionary entities. Only the tribe Arvicolini emerged as paraphyletic in both analyses because of the unresolved phylogenetic position of Arvicola terrestris . Steppe voles of the genus Lagurus were solidly supported as a sister group to the Microtus and allies clade.  © 2008 The Linnean Society of London, Biological Journal of the Linnean Society , 2008, 94 , 825–835.     

Phylogeny and classification of whirligig beetles (Coleoptera: Gyrinidae): relaxed‐clock model outperforms parsimony and time‐free Bayesian analyses

Phylogenetic relationships among members of the family Gyrinidae (Coleoptera: Adephaga) were inferred from analysis of 42 morphological characters and DNA sequence data from the genes 12S rRNA, cytochrome c oxidase I and II, elongation factor 1 alpha (2 different copies) and histone III. Eighty‐nine species of Gyrinidae were included representing all known subfamilies, tribes and genera. Outgroups include species from Noteridae, Paelobiidae and Dytiscidae. Analyses include parsimony analysis, and partitioned time‐free and relaxed‐clock Bayesian analyses of the combined data using reversible‐jump MCMC to simultaneously integrate over all possible 4 × 4 nucleotide substitution models. Analyses resulted in conflicting topologies between the combined parsimony and Bayesian analyses on the one hand, and the relaxed‐clock analysis on the other. The marginal likelihoods of competing models were calculated with stepping‐stone sampling and used in a Bayes factor test, which, along with arguments from morphology, supported the topology generated by the relaxed‐clock analysis. This phylogenetic hypothesis is adopted to revise the higher classification of Gyrinidae. Major taxonomic conclusions include: (i) monophyletic Gyrinidae, (ii) the Nearctic Spanglerogyrinae Folkerts (with one species, Spanglerogyrus albiventris Folkerts) sister to all other Gyrinidae, (iii) the Madagascar endemic Heterogyrinae Brinck stat. n. (with one species, Heterogyrus milloti Legros) sister to all Gyrinidae except Spanglerogyrinae, (iv) monophyletic Gyrininae Latreille including three monophyletic tribes with the following relationship: Orectochilini Régimbart + (Gyrinini Latreille + Enhydrini Régimbart), (v) monophyletic Orectochilini comprising four monophyletic genera with the following relationships: (Gyretes Brullé + Patrus Aubé stat. n. ) + (Orectogyrus Régimbart + Orectochilus Dejean), (vi) monophyletic Gyrinini comprising three genera with the following relationships: Gyrinus Geoffroy + (Metagyrinus Brinck + Aulonogyrus Motschulsky), each monophyletic except Metagyrinus with only one included species and not tested for monophyly, and (vii) monophyletic Enhydrini comprising five genera with the following relationships: (Porrorhynchus Laporte + Dineutus MacLeay) + (Enhydrus Laporte + (Andogyrus Ochs + Macrogyrus Régimbart)), each monophyletic except Porrorhynchus, Enhydrus and Andogyrus each with one included species and untested for monophyly. Each subfamily, tribe and genus is diagnosed and discussed. The female reproductive tract of each group is presented, illustrated and discussed with respect to the phylogenetic conclusions.     

A new genus of mantidflies discovered in the Oriental region,with a higher‐level phylogeny of Mantispidae (Neuroptera) using DNA sequences and morphology

A remarkable new genus and two new species of Mantispidae (Neuroptera) are described from the Oriental region. Allomantispa Liu, Wu, Winterton & Ohl gen.n. , currently including A. tibetana Liu, Wu & Winterton sp.n. and A. mirimaculata Liu & Ohl sp.n. The new genus is placed in the subfamily Drepanicinae based on a series of morphological characteristics and on the results of total evidence phylogenetic analyses. Bayesian and Parsimony analyses were undertaken using three gene loci (CAD, 16S rDNA and COI) combined with 74 morphological characters from living and fossil exemplars of Mantispidae (17 genera), Rhachiberothidae (two genera) and Berothidae (five genera), with outgroup taxa from Dilaridae and Osmylidae. The resultant phylogeny presented here recovered a monophyletic Mantispidae with ?Mesomantispinae sister to the rest of the family. Relationships among Mantispidae, Rhachiberothidae and Berothidae support Rhachiberothidae as a separate family sister to Mantispidae. Within Mantispidae, Drepanicinae are a monophyletic clade sister to Calomantispinae and Mantispinae. In a combined analysis, Allomantispa gen.n. was recovered in a clade comprising Ditaxis McLachlan from Australia, and two fossil genera from the Palaearctic, ?Promantispa Panfilov (Kazakhstan; late Jurassic) and ?Liassochrysa Ansorge & Schlüter (Germany; Jurassic), suggesting a highly disjunct and relictual distribution for the family. This published work has been registered in ZooBank, http://zoobank.org/urn:lsid:zoobank.org:pub:464B06E8‐47E6‐482E‐8136‐83FE3B2E9D6B .     

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). This published work has been registered in ZooBank, http://zoobank.org/urn:lsid:zoobank.org:pub:440E7FA4‐C859‐47E0‐8335‐30D478CBA8FA .     

Morphology-based phylogeny of the treehopper family Membracidae (Hemiptera: Cicadomorpha: Membracoidea)

A parsimony‐based phylogenetic analysis of eighty‐three morphological characters of adults and immatures of seventy representatives of the tribes and subfamilies of Membracidae and two outgroup taxa was conducted to evaluate the status and relationships of these taxa. Centrotinae apparently gave rise to Nessorhinini and Oxyrhachini (both formerly treated as subfamilies, now syn.n. and syn.reinst., respectively, of Centrotinae). In contrast to previous analyses, a clade comprising Nicomiinae, Centronodinae, Centrodontinae, and the unplaced genera Holdgatiella Evans, Euwalkeria Goding and Antillotolania Ramos was recovered, but relationships within this clade were not well resolved. Nodonica bispinigera, gen.n. and sp.n., is described and placed in Centrodontini based on its sister‐group relationship to a clade comprising previously described genera of this tribe. Membracinae and Heteronotinae were consistently monophyletic. Neither Darninae nor Smiliinae, as previously defined, was monophyletic on the maximally parsimonious cladograms, but constraining both as monophyletic groups required only one additional step. The monophyly of Stegaspidinae, including Deiroderes Ramos (unplaced in Membracidae), was supported on some but not all equally parsimonious cladograms. More detailed analyses of individual subfamilies, as well as morphological data on the undescribed immatures of several membracid tribes and genera, will be needed to elucidate relationships among tribes and genera. A key to the subfamilies and tribes is provided.     

Phylogenetic analysis of the Mexican freshwater crabs of the tribe Pseudothelphusini (Decapoda: Brachyura: Pseudothelphusidae)

The Mexican pseudothelphusid crabs are classified in one subfamily, three tribes, and 13 genera. Up to now, 56 species have been recognized, distributed in a strictly Neotropical pattern, with some of them reaching the state of Sonora on the western slope of Mexico. The tribe Pseudothelphusini is the most diverse, with five genera and 35 species, all of them endemic to Mexico: the two most species‐rich genera are Pseudothelphusa, with 23 species, and Tehuana, with eight species; Epithelphusa includes two species, whereas Disparithelphusa and Smalleyus are monotypic. The Pseudothelphusini lack an updated systematic revision, which could serve as a framework to analyse the monophyletic origin of the group, to clarify the relationships among genera and species, as well as to resolve the taxonomic status of various species complexes. In the present study, an exhaustive morphological revision was conducted using somatic and sexual characters. A phylogenetic analysis was performed using 77 characters and 183 character states, taken from 41 species. Ten trees of the same length were obtained using PAUP 4.0 through a heuristic search. The results show that the tribe as it is actually known constitutes a paraphyletic group, in which the species of Epithelphusa and Pseudothelphusa puntarenas are excluded from the internal group. According to the obtained results, the tribe Pseudothelphusini s.s. includes five genera: Smalleyus, Pseudothelphusa, Tehuana, and two new ones to accommodate Pseudothelphusa galloi and Pseudothelphusa sulcifrons, respectively. This new arrangement considers the provisional suppression of the genus Disparithelphusa, which remained as another species of Pseudothelphusa throughout the cladistic analysis. The phylogenetic results show a strong congruence with the distribution of the species, in several cases grouping species that form morphological clines along a geographical gradient. The previously proposed southern origin of the tribe Pseudothelphusini gains support with the results obtained. © 2010 The Linnean Society of London, Zoological Journal of the Linnean Society, 2010, 160 , 457–481     

Independent origins of coastal colonization in the tribe Athetini (Coleoptera,Staphylinidae)

Less than 1% of Staphylinidae are known to be confined to coastal habitats. To explore the origins of coastal colonization within the tribe Athetini Casey, we present a revised molecular phylogeny. The dataset comprised partial mitochondrial COI, COII, 16S rDNA, NADH1, partial nuclear 18S rDNA and 28S rDNA. We chose a total of 95 species in 51 genera, including 14 coastal species in eight genera and 21 outgroup species from other aleocharine tribes. The concatenated dataset was analysed simultaneously by both parsimony‐ and model‐based (Bayesian and maximum likelihood) methods. The tribe Athetini was not supported as a monophyletic group, but together with the tribes Tachyusini, Ecitocharini and Hygronomini did form a monophylum. The ecological association of species with a coastal habitat was mapped onto a phylogeny to assess the evolution of habitat specialization in the Athetini lineage. The results reveal that five independent origins of coastal colonization have occurred throughout the tribe Athetini: (a) Osakatheta + Adota minuta + coastal Atheta (Badura) (clade A); (b) Adota (clade B); (c) Pontomalota + Tarphiota + Thinusa (clade C); (d) Iotarphia (clade D); and (e) Psammostiba (clade E). The low species number of the coastal Athetini compared with the entire Athetini lineage indicates that coastal habitats are harsh environments and so only a few species were able to colonize this habitat. The following changes in classification are proposed: (a) Ad. minuta Lee and Ahn is removed from the genus Adota and tentatively included in Atheta (Badura); (b) The genus Saphocallus Sharp is transferred from Athetini to Geostibini.     

Reevaluation of Blapimorpha and Opatrinae: addressing a major phylogeny‐classification gap in darkling beetles (Coleoptera: Tenebrionidae: Blaptinae)

The taxonomic concepts of Blapimorpha and Opatrinae (informal and traditional, morphology‐based groupings among darkling beetles) are tested using molecular phylogenetics and a reassessment of larval and adult morphology to address a major phylogeny‐classification gap in Tenebrionidae. Instead of a holistic approach (family‐level phylogeny), this study uses a bottom‐up strategy (tribal grouping) in order to define larger, monophyletic lineages within Tenebrioninae. Sampling included representatives of 27 tenebrionid tribes: Alleculini, Amarygmini, Amphidorini, Blaptini, Bolitophagini, Branchini, Cerenopini, Coniontini, Caenocrypticini, Dendarini, Eulabini, Helopini, Lagriini, Melanimini, Opatrini, Pedinini, Phaleriini, Physogasterini, Platynotini, Platyscelidini, Praociini, Scaurini, Scotobiini, Tenebrionini, Trachyscelini, Triboliini and Ulomini. Molecular analyses were based on DNA sequence data from four non‐overlapping gene regions: carbamoyl‐phosphate synthetase domain of rudimentary (CAD) (723 bp), wingless (wg) (438 bp) and nuclear ribosomal 28S (1101 bp) and mitochondrial ribosomal 12S (363 bp). Additionally, 15 larval and imaginal characters were scored and subjected to an ancestral state reconstruction analysis. Results revealed that Amphidorini, Blaptini, Dendarini, Pedinini, Platynotini, Platyscelidini and Opatrini form a clade which can be defined by the following morphological features: adults—antennae lacking compound/stellate sensoria; procoxal cavities externally and internally closed, intersternal membrane of abdominal ventrites 3–5 visible; paired abdominal defensive glands present, elongate, not annulated; larvae—prolegs enlarged (adapted for digging); ninth tergite lacking urogomphi. To accommodate this monophyletic grouping (281 genera and ～4000 species), the subfamily Blaptinae sens. nov. is resurrected. Prior to these results, all of the tribes within Blaptinae were classified within the polyphyletic subfamily Tenebrioninae. The non‐monophyletic nature of Terebrioninae has already been postulated by previous authors, yet no taxonomic decisions were made to fix its status. The reinstatement of Blaptinae, which groups ～50% of the former Tenebrioninae, helps to clarify phylogenetic relations among the whole family and is the first step towards a complete higher‐level revision of Tenebrionidae. The Central Asian tribe Dissonomini (two genera, ～30 species) was not included in Blaptinae due to a lack of representatives in the performed phylogenetic analyses; however, based on morphological features, the tribe is listed as a potential addition to the subfamily.     

Evolution of blindness in scolopendromorph centipedes (Chilopoda: Scolopendromorpha): insight from an expanded sampling of molecular data

Relative to its diversity (34 genera, 700 species), Scolopendromorpha has been undersampled in molecular phylogenetic analyses compared with the other chilopod orders. Previous analyses based on morphology have not resolved several key controversies in systematics and evolutionary morphology unambiguously. Here we apply new molecular and morphological data to scolopendromorph phylogenetics, with a focus on the evolution of blindness. The taxonomic sample includes 19 genera, many lacking previous molecular data, and diverse, cosmopolitan genera of Scolopendridae are sampled by multiple species. Phylogenetic analysis with Direct Optimization used 94 morphological characters and ca. 4.5 kb of sequence data from two nuclear (18S and 28S rRNA) and two mitochondrial (16S rRNA and COI) loci. A single most‐parsimonious cladogram selected after sensitivity analyses resolves Scolopendromorpha as monophyletic, and divides it into a blind clade of three families (Plutoniumidae, Cryptopidae, Scolopocryptopidae) and its ocellate sister group, Scolopendridae. Some species‐rich, cosmopolitan genera (Cormocephalus, Otostigmus, Scolopendra) in Scolopendridae are non‐monophyletic, and in several instances (e.g. New and Old World Scolopendra) relationships are more congruent with geographical distributions than with traditional classifications. The tribe Asanadini is particularly subject to parameter‐sensitivity, nesting in the combined analysis within Scolopendrini but as sister to all other Scolopendrinae for molecular data alone. The total‐evidence tree unambiguously optimizes trunk segmentation: a 23‐segmented trunk has a single origin in the blind clade. © The Willi Hennig Society 2011.     

A remarkable new opsiine leafhopper genus (Hemiptera: Cicadellidae: Deltocephalinae) from Southern China,with notes on its phylogenetic position

A remarkable new genus of tribe Opsiini (Deltocephalinae), Yinformibus gen. nov. including Yinformibus menglaensis sp. nov. is described from Yunnan of China. The new genus is placed in the tribe Opsiini based on male adults having a pair of aedeagal shafts, each with its own gonopore. Partial 28S rDNA and Histone H3 sequences are provided for the new species, and a phylogenetic analysis based on these markers for Deltocephalinae retrieved from GenBank suggests Opsiini is monophyletic and supports the placement of Yinformibus gen. nov. in Opsiini. The molecular results placed the new genus in a clade comprising Hishimonus Ishihara, Opsius Fieber, and an undescribed genus from Zambia. Additionally, we also analysed the relationships of Yinformibus gen. nov. with other genera based on morphology.

http://www.zoobank.org/urn:lsid:zoobank.org:pub:85C23AC3-8B19-4F29-9F55-BFC392242B51     

Evolution of the Canthonini Longitarsi (Scarabaeidae) in Madagascar

Dung beetle species belonging to the worldwide tribe Canthonini (Scarabaeidae) and occurring in Madagascar are all endemic to that island. The Malagasy Canthonini form three lineages, one of which is the group Longitarsi that includes five genera. The phylogenetic relationships of Malagasy Canthonini are not fully resolved and only few species of Longitarsi have been included in previous studies. Here we infer the phylogenetic relationships within the Longitarsi group using molecular data and together with morphological examination revise the systematics of the group. The five genera of the Longitarsi group form one monophyletic clade and thus we suggest the synonymization of the younger genera Sikorantus, Phacosomoides, Madaphacosoma and Aleiantus; with the oldest genus belonging to this clade Epactoides. We describe two new species: Epactoides jounii sp. n and Epactoides mangabeensis sp. n. Most of the species of Longitarsi inhabit the eastern rainforests, with very low local species diversity and highly restricted geographical ranges. In the group Longitarsi four species are wingless. The loss of wings has evolved at least twice, at high altitude along the mountain range.     

Molecular phylogeny of Palaearctic genera of Gomphocerinae grasshoppers (Orthoptera, Acrididae)

Abstract.  Molecular phylogenetic methods were used to examine morphologically based hypotheses concerning the taxonomic structure and relationships of the grasshopper subfamily Gomphocerinae. Two mitochondrial gene (cytochrome b and cytochrome oxidase subunit I) sequences were determined for twenty-five species representing eleven Palaearctic genera. The studied Gomphocerinae species constituted a monophyletic group; furthermore, the earlier division of Gomphocerinae into tribes was supported, with each tribe monophyletic. There was no support for various systems uniting Stenobothrini and Gomphocerini into one tribe. Two separate clusters were discerned in Gomphocerini and two tribes were distinguished – Gomphocerini (genera Aeropus , Stauroderus , Chorthippus ) and Stenobothrini (genera Omocestus , Stenobothrus ).     

Phylogenetic reassessment of the Exophthalmus genus complex (Curculionidae: Entiminae: Eustylini,Geonemini)

The monophyly of the Neotropical entimine weevil genus Exophthalmus Schoenherr, 1823 (Curculionidae: Entiminae: Eustylini Lacordaire) is reassessed. Exophthalmus presently includes more than 80 species, approximately half of which are restricted to either the Caribbean archipelago or the continental Neotropics. The taxonomic composition and position of Exophthalmus have been subject to longstanding disagreements; in particular, authors have questioned the relationship of Exophthalmus to other Caribbean genera such as Diaprepes Schoenherr, 1834 (Eustylini) and Lachnopus Schoenherr, 1840 (Geonemini Gistel), as well as to the speciose Central and South American genera Compsus Schoenherr, 1823, Eustylus Schoenherr, 1842, and Exorides Pascoe, 1881 (all Eustylini), among others. The present study scrutinizes these traditional perspectives, based on a cladistic analysis of 143 adult morphological characters and 90 species, representing 30 genera and seven tribes of Neotropical entimine weevils. The character matrix yielded eight most‐parsimonious cladograms (length = 239 steps; consistency index = 66; retention index = 91), with mixed clade support that remains particularly wanting for some of the deeper in‐group divergences. The strict consensus supports the existence of a paraphyletic Geonemini ‘grade’ that includes Lachnopus and related Caribbean genera such as Apotomoderes Dejean, 1834, followed by a monophyletic Eustylini in‐group clade. Within the latter, a monophyletic South American Eustylini clade – including Compsus, Eustylus, Exorides, and related genera – is sister to a major clade that contains a ‘grade’ of heterogeneous and often misclassified Caribbean members of the Eustylini, Geonemini (Tetrabothynus Labram & Imhoff, 1852 and Tropirhinus Schoenherr, 1823), and Tanymecini Lacordaire (Pachnaeus Schoenherr, 1826), as well as two major clades: one with the majority of Central American Exophthalmus species, and the other with most Caribbean members of Exophthalmus. The Central American Exophthalmus clade is paraphyletic with respect to Chauliopleurus Champion, 1911 (Geonemini) and Rhinospathe Chevrolat, 1878 (Phyllobiini Schoenherr). The Caribbean clade, in turn, contains two subclades: i.e. (1) the Greater Antillean Exophthalmus s.s. clade, including the type species Exophthalmus quadrivittatus (Olivier, 1807); and (2) the primarily Lesser Antillean Diaprepes. The latter genus is therefore nested within Central American and Caribbean species of a highly paraphyletic Exophthalmus, yet may be rendered monophyletic if several Lesser Antillean Exophthalmus species are (re‐)assigned to Diaprepes. The results thus provide a suitable basis for a revision of all Exophthalmus species, and furthermore suggest that historical biographic factors, including colonization via temporary continental Neotropics‐to‐Caribbean land connections, were important in the evolution of major eustyline lineages. Based on these preliminary insights, the following taxonomic and nomenclatural adjustments are made. Compsoricus gen. nov. is erected to accommodate two Puerto Rican species erroneously assigned to Compsus: i.e. the herein designated type species Compsoricus maricao comb. nov. and Compsoricus luquillo comb. nov. Eustylus dentipes comb. nov. is transferred from Compsus. Diaprepes marginicollis Chevrolat, 1880 is reinstated from synonymy under Exophthalmus. Lastly, the following five transfers are proposed: (1) Chauliopleurus Champion, 1911, from Geonemini to Eustylini; (2) Tetrabothynus Labram & Imhoff, 1852, from Geonemini to Eustylini; (3) Tropirhinus Schoenherr, 1823, from Geonemini to Eustylini; (4) Rhinospathe Chevrolat, 1878, from Phyllobiini to Eustylini; and (5) Pachnaeus Schoenherr, 1826, from Tanymecini to Eustylini. © 2012 The Linnean Society of London, Zoological Journal of the Linnean Society, 2012, 164 , 510–557.