The systematics of right whales (Mysticeti: Balaenidae)

Balaenidae (right whales) are large, critically endangered baleen whales represented by four living species. The evolutionary relationships of balaenids are poorly known, with the number of genera, relationships to fossil taxa, and position within Mysticeti in contention. This study employs a comprehensive set of morphological characters to address aspects of balaenid phylogeny. A sister‐group relationship between neobalaenids and balaenids is strongly supported, although this conflicts with molecular evidence, which may be an artifact of long‐branch attraction (LBA). Monophyly of Balaenidae is supported, and three major clades are recognized: (1) extinct genus Balaenula, (2) extant and extinct species of the genus Eubalaena, and (3) extant and extinct species of the genus Balaena plus the extinct taxon, Balaenella. The relationships of these clades to one another, as well as to the early Miocene stem balaenid, Morenocetus parvus, remain unresolved. Pliocene taxa, Balaenula astensis and Balaenula balaenopsis, form a clade that is the sister group to the Japanese Pliocene Balaenula sp. Eubalaena glacialis and Pliocene Eubalaena belgica, are in an unresolved polytomy with a clade including E. japonica and E. australis. Extant and fossil species of Balaena form a monophyletic group that is sister group to the Dutch Pliocene Balaenella, although phylogenetic relationships within Balaena remain unresolved.     

Phylogeny of Neotropical oryzomyine rodents (Muridae: Sigmodontinae) based on the nuclear IRBP exon

Sigmodontine rodents are the most diverse family-level mammalian clade in the Neotropical region, with about 70 genera and 320 recognized species. Partial sequences (1266 bp) from the first exon of the nuclear gene encoding the Interphotoreceptor Retinoid Binding Protein (IRBP) were used to infer the phylogenetic relationships among 44 species representing all 16 currently recognized genera of the largest sigmodontine tribe, the Oryzomyini. Monophyly of the tribe was assessed relative to 15 non-oryzomyine sigmodontine taxa representing all major sigmodontine lineages. Twelve taxa from seven muroid subfamilies were used as outgroups. The resulting matrix included 71 taxa and 386 parsimony-informative characters. Phylogenetic analysis of this matrix resulted in 16 equally parsimonious cladograms, which contained the following well-supported groups: (i). a monophyletic Oryzomyini, (ii). a clade containing all oryzomyines except Scolomys and Zygodontomys, (iii). a clade containing Oecomys, Handleyomys, and several species of forest-dwelling Oryzomys, and (iv). a clade containing the remaining oryzomyine taxa. The last clade is composed of two large subclades, each with lower nodal support, containing the following taxa: (i). Microryzomys, Oligoryzomys, Neacomys, and Oryzomys balneator; (ii). Holochilus, Lundomys, Pseudoryzomys, Nectomys, Amphinectomys, Sigmodontomys, and several species of open-vegetation or semiaquatic Oryzomys. Regarding relationships among non-oryzomyine taxa, sigmodontines, neotomines, and tylomyines do not form a monophyletic group; a clade containing Rheomys and Sigmodon is basal relative to all other sigmodontines; and the remaining sigmodontines are grouped in three clades: the first containing Thomasomyini, Akodontini, and Reithrodon; the second containing Abrothrichini, and Phyllotini, plus Wiedomys, Juliomys, Irenomys, and Delomys; and the third containing the oryzomyines. No conflict is observed between IRBP results and previous robust hypotheses from mitochondrial data, while a single case of incongruence is present between the IRBP topology and robust hypothesis from morphological studies.     

Phylogenetic relationships within the South American fish family Anostomidae (Teleostei,Ostariophysi, Characiformes)

Analysis of a morphological dataset containing 152 parsimony‐informative characters yielded the first phylogenetic reconstruction spanning the South American characiform family Anostomidae. The reconstruction included 46 ingroup species representing all anostomid genera and subgenera. Outgroup comparisons included members of the sister group to the Anostomidae (the Chilodontidae) as well as members of the families Curimatidae, Characidae, Citharinidae, Distichodontidae, Hemiodontidae, Parodontidae and Prochilodontidae. The results supported a clade containing Anostomus, Gnathodolus, Pseudanos, Sartor and Synaptolaemus (the subfamily Anostominae sensu Winterbottom) albeit with a somewhat different set of relationships among the species within these genera. Anostomus as previously recognized was found to be paraphyletic and is split herein into two monophyletic components, a restricted Anostomus and the new genus Petulanos gen. nov. , described herein. Laemolyta appeared as sister to the clade containing Anostomus, Gnathodolus, Petulanos, Pseudanos, Sartor and Synaptolaemus. Rhytiodus and Schizodon together formed a well‐supported clade that was, in turn, sister to the clade containing Anostomus, Gnathodolus, Laemolyta, Petulanos, Pseudanos, Sartor and Synaptolaemus. Anostomoides was sister to the clade formed by these nine genera. Leporinus as currently defined was not found to be monophyletic, although certain clades within that genus were supported, including the species with subterminal mouths in the former subgenus Hypomasticus which we recognize herein as a genus. Abramites nested in Leporinus, and Leporellus was found to be the most basal anostomid genus. The presence of cis‐ and trans‐Andean species in Abramites, Leporellus, Leporinus and Schizodon, all relatively basal genera, suggests that much of the diversification of anostomid species pre‐dates the uplift of the Andean Cordilleras circa 11.8 million years ago. Several important morphological shifts in anostomid evolution are illustrated and discussed, including instances of convergence and reversal. © 2008 The Linnean Society of London, Zoological Journal of the Linnean Society, 2008, 154 , 70–210.     

The Taxonomic and Evolutionary History of Fossil and Modern Balaenopteroid Mysticetes

Balaenopteroids (Balaenopteridae + Eschrichtiidae) are a diverse lineage of living mysticetes, with seven to ten species divided between three genera (Megaptera, Balaenoptera and Eschrichtius). Extant members of the Balaenopteridae (Balaenoptera and Megaptera) are characterized by their engulfment feeding behavior, which is associated with a number of unique cranial, mandibular, and soft anatomical characters. The Eschrichtiidae employ suction feeding, which is associated with arched rostra and short, coarse baleen. The recognition of these and other characters in fossil balaenopteroids, when viewed in a phylogenetic framework, provides a means for assessing the evolutionary history of this clade, including its origin and diversification. The earliest fossil balaenopterids include incomplete crania from the early late Miocene (7–10 Ma) of the North Pacific Ocean Basin. Our preliminary phylogenetic results indicate that the basal taxon, “Megaptera” miocaena should be reassigned to a new genus based on its possession of primitive and derived characters. The late late Miocene (5–7 Ma) balaenopterid record, except for Parabalaenoptera baulinensis and Balaenoptera siberi, is largely undescribed and consists of fossil specimens from the North and South Pacific and North Atlantic Ocean basins. The Pliocene record (2–5 Ma) is very diverse and consists of numerous named, but problematic, taxa from Italy and Belgium, as well as unnamed taxa from the North and South Pacific and eastern North Atlantic Ocean basins. For the most part Pliocene balaenopteroids represent extinct species and genera and reveal a greater degree of morphological diversity than at present. The Pleistocene record is very limited and, unfortunately, fails to document the evolutionary details leading to modern balaenopteroid species diversity. It is evident, however, that most extant species evolved during the Pleistocene. Morphological and molecular based phylogenies support two competing hypotheses concerning relationships within the Balaenopteroidea: (1) balaenopterids and eschrichtiids as sister taxa, and (2) eschrichtiids nested within a paraphyletic Balaenopteridae. The addition of fossil taxa (including a new Pliocene species preserving a mosaic of balaenopterid and eschrichtiid characters) in morphological and “total evidence” analyses, offers the potential to resolve the current controversy concerning the possible paraphyly of Balaenopteridae.     

Phylogeny of the bee family Megachilidae (Hymenoptera: Apoidea) based on adult morphology

Phylogenetic relationships within the bee family Megachilidae are poorly understood. The monophyly of the subfamily Fideliinae is questionable, the relationships among the tribes and subtribes in the subfamily Megachilinae are unknown, and some extant genera cannot be placed with certainty at the tribal level. Using a cladistic analysis of adult external morphological characters, we explore the relationships of the eight tribes and two subtribes currently recognised in Megachilidae. Our dataset included 80% of the extant generic‐level diversity, representatives of all fossil taxa, and was analysed using parsimony. We employed 200 characters and selected 7 outgroups and 72 ingroup species of 60 genera, plus 7 species of 4 extinct genera from Baltic amber. Our analysis shows that Fideliinae and the tribes Anthidiini and Osmiini of Megachilinae are paraphyletic; it supports the monophyly of Megachilinae, including the extinct taxa, and the sister group relationship of Lithurgini to the remaining megachilines. The Sub‐Saharan genus Aspidosmia, a rare group with a mixture of osmiine and anthidiine features, is herein removed from Anthidiini and placed in its own tribe, Aspidosmiini, new tribe . Protolithurgini is the sister of Lithurgini, both placed herein in the subfamily Lithurginae; the other extinct taxa, Glyptapina and Ctenoplectrellina, are more basally related among Megachilinae than Osmiini, near Aspidosmia, and are herein treated at the tribal level. Noteriades, a genus presently in the Osmiini, is herein transferred to the Megachilini. Thus, we recognise four subfamilies (Fideliinae, Pararhophitinae, Lithurginae and Megachilinae) and nine tribes in Megachilidae. We briefly discuss the evolutionary history and biogeography of the family, present alternative classifications, and provide a revised key to the extant tribes of Megachilinae.     

Phylogeny and diversification of the cloud forest Morpho sulkowskyi group (Lepidoptera,Nymphalidae) in the evolving Andes

The monophyletic Morpho sulkowskyi butterfly group, endemic of Andean cloud forests, was studied to test the respective contributions of Mio‐Pliocene intense uplift period and Pleistocene glacial cycles on Andean biodiversity. We sampled nine taxa covering the whole geographical range of the group. Two mitochondrial and two nuclear genes were analysed using a Bayesian method. We established a dated phylogeny of the group using a relaxed clock method and a wide‐outgroup approach. To discriminate between two hypotheses, we used a biogeographical probabilistic method. Results suggest that the ancestor of the M. sulkowskyi group originated during the Middle–Late Miocene uplift of the Eastern Cordillera in northern Peru. Biogeographical inference suggests that the M. sulkowskyi and Morpho lympharis clades diverged in the northern Peruvian Andes. The subsequent divergences, from the Late Miocene to the Late Pliocene, should have resulted from a dispersal towards the Northern Andes (M. sulkowskyi clade), after the closure of the West Andean Portal separating the Central and Northern Andes, and a southwards dispersal along the Peruvian and Bolivian Eastern Cordilleras (M. lympharis clade). Only a few divergences occurred at the very end of the Pliocene or during the Pleistocene, a period when the more recent uplifts interfered with Pleistocene glacial cycles.     

Evolution of wing polymorphism and genital asymmetry in the thread‐legged bugs of the tribe Metapterini Stål (Hemiptera,Reduviidae, Emesinae) based on morphological characters

Wing polymorphism and asymmetric male genitalia are intriguing morphological phenomena occurring in insects. Among Emesinae, or thread‐legged bugs, the tribe Metapterini Stål exhibits these two interesting morphological attributes. Nonetheless, evolutionary interpretations of these phenomena cannot be put forward because phylogenetic hypotheses for Emesinae are lacking. Thread‐legged bugs are easily recognized among assassin bugs due to their elongated and seemingly delicate body. The tribe Metapterini has 28 genera and approximately 280 described species. The only available phylogenetic hypothesis among Emesinae tribes was proposed by Wygodzinsky (1966), and it hypothesized Deliastini Villiers as the sister group of Metapterini, although this hypothesis has never been tested with cladistic approaches. Recent analyses using character sets of genitalia and prolegs suggest that Metapterini might not be monophyletic. In order to test these ideas, we compiled a morphological dataset of 138 characters that includes external morphological characters, detailed features of prolegs and genitalia of both sexes for Metapterini, which were analysed cladistically including 55 terminals, comprising 24 genera (85.7% of the generic diversity), 43 species of Metapterini and 12 outgroups. Metapterini was recovered as paraphyletic by the inclusion of Bergemesa Wygodzinsky, Palacus Dohrn and Stalemesa Wygodzinsky, all currently assigned to Deliastini. Gardena Dohrn (Emesini) was recovered as the sister group of Metapterini + Deliastini as suggested by Wygodzinsky (1966). Based on these results, we synonymize Deliastini syn. n. with Metapterini sensu n. and propose two new genera: Bacata Castro‐Huertas & Forero gen. n. , for three Andean species previously placed in Liaghinella Wygodzinsky, and Valkyriella Castro‐Huertas & Forero gen. n. for Ghilianella borgmeieri Wygodzinsky. Ancestral state reconstruction of wing polymorphism indicates that males and females were fully winged in the ancestor of Metapterini sensu n. with two independent evolutionary transitions to the apterous and brachypterous conditions. The analysis of the symmetry of the male genitalia shows an ancestor with symmetric male genitalia and two independent emergences of asymmetrical male genitalia in Metapterini.     

Relationships within Siphini (Hemiptera,Aphidoidea: Chaitophorinae) in light of molecular and morphological research

The aim of this paper was to further explore the phylogeny of Siphini by analysing molecular data (two mitochondrial genes and two nuclear markers), together with morphological (29) and ecological (two) characters, for comprehensive analyses concerning the evolution of Siphini, relationships within the tribe, and between Siphini and other Chaitophorinae. Nine Siphini species, which represent all the genera of this tribe, as well as 12 out‐group species (mainly Chaitophorini representatives of the genera Chaitophorus and Periphyllus), were used in the analyses. Molecular phylogenetic trees were reconstructed by the Bayesian inference (BI) phylogenetic analysis and maximum parsimony (MP) criterion. The cladistic analysis was performed using nona . The monophyly of Siphini was confirmed. Species belonging to subgenera Sipha and Rungsia were clustered together, and this clade was a sister with reference to a clade including the genera Atheroides and Chaetosiphella. Monophyly of Atheroides was confirmed by the molecular data; however, in cladistic analysis Atheroides seemed to be paraphyletic because Atheroides hirtellus was placed as sister to Atheroides serrulatus and Chaetosiphella. The monotypic genera Caricosipha and Laingia formed separate lineages, and Laingia was sister to all other Siphini. Chaitophorini was not retrieved by the molecular and combined data: Periphyllus was sister to a clade containing Chaitophorus and Siphini.     

A remarkable autosomal heteromorphism in Pseudoryzomys simplex 2n = 56; FN = 54–55 (Rodentia,Sigmodontinae)

Pseudoryzomys simplex, the false rice rat, is a monotypic genus of the Oryzomyini tribe (Sigmodontinae) distributed in part of Bolivia, Paraguay, Argentina and Brazil. Its diploid number has been described as 56 acrocentric chromosomes decreasing in size and no karyotype figure has been depicted. Herein, we present karyotypic data on P. simplex, including chromosome banding and molecular fluorescent in situ hybridization using telomeric sequences and the whole X-chromosome of its sister clade Holochilus brasiliensis (HBR) as probes. A case of remarkable autosomal heteromorphism due to the presence of a whole heterochromatic arm leading to the variability of FN is reported, as well as the occurrence of regions of homology between the X and Y chromosomes (pseudoautosomal regions) after chromosome painting with the HBR X probe on P. simplex metaphases.     

Phylogeny of the North American catfish family Ictaluridae (Teleostei: Siluriformes) combining morphology,genes and fossils

We performed the first combined‐data phylogenetic analysis of ictalurids including most living and fossil species. We sampled 56 extant species and 16 fossil species representing outgroups, the seven living genera, and the extinct genus ?Astephus long thought to be an ictalurid. In total, 209 morphological characters were curated and illustrated in MorphoBank from published and original work, and standardized using reductive coding. Molecular sequences harvested from GenBank for one nuclear and four mitochondrial genes were combined with the morphological data for total evidence analysis. Parsimony analysis recovers a crown clade Ictaluridae composed of seven living genera and numerous extinct species. The oldest ictalurid fossils are the Late Eocene members of Ameiurus and Ictalurus. The fossil clade ?Astephus placed outside of Ictaluridae and not as its sister taxon. Previous morphological phylogenetic studies of Ictaluridae hypothesized convergent evolution of troglobitic features among the subterranean species. In contrast, we found morphological evidence to support a single clade of the four troglobitic species, the sister taxon of all ictalurids. This result holds whether fossils are included or not. Some previously published clock‐based age estimates closely approximate our minimum ages of clades.     

PHYLOGENY OF THE RUBIACEAE AND THE LOGANIACEAE: CONGRUENCE OR CONFLICT BETWEEN MORPHOLOGICAL AND MOLECULAR DATA?

Phylogenetic analyses of 33 genera of Rubiaceae were performed using morphological and a few chemical characters. Parsimony analysis based on 29 characters resulted in eight equally parsimonious trees, with a consistency index of 0.40 and a retention index of 0.69. These results were compared to a phylogenetic analysis of the same genera based on chloroplast DNA restriction site data. There are discrepancies between the two analyses, but if we consider groupings reflected in the present classification there is much congruency. With the exception of four genera, all the genera are positioned in the same group of taxa in the two analyses. Clades of taxa representing three of the four subfamilies (~the Antirheoideae, ~the Rubioideae, and the ~Ixoroideae) are monophyletic, while the fourth subfamily Cinchonoideae is shown to be paraphyletic. Both analyses support a widened tribe Chiococceae, including the former subtribe Portlandiinae (Condamineeae). Furthermore, in both analyses the tribe Hamelieae is placed outside the subfamily Rubioideae where it is now housed. In search for the most plausible sister group to the Rubiaceae, the genus Cinchona (Rubiaceae) was analyzed together with 13 genera of the Loganiaceae, Nerium (Apocynaceae), and Exacum (Gentianaceae). Cornus (Comaceae), Olea (Oleaceae), and these two genera together were used as outgroups. The analysis, including 25 characters, 16 taxa, and with Cornus and Olea together as an outgroup, resulted in four equally parsimonious trees, with a consistency index of 0.53 and a retention index of 0.62. The non-Loganiaceae taxa Cinchona (Rubiaceae), Nerium (Apocynaceae), and Exacum (Gentianaceae) were all found to have their closest relatives within the Loganiaceae indicating that the Loganiaceae are paraphyletic and ought to be reclassified. As a result of the morphological data the most plausible sister group to the Rubiaceae is the tribe Gelsemieae of the Loganiaceae.     

Phylogeny of the tribe Abrotrichini (Cricetidae,Sigmodontinae): integrating morphological and molecular evidence into a new classification

The tribe Abrotrichini (five genera and 14 living species) is a small clade within the speciose subfamily Sigmodontinae (Rodentia, Cricetidae), representing one of the extant successful radiations of mammals at southern high latitudes of the Neotropics. Its distribution is mostly Andean, reaching its greatest diversity in southern Argentina and Chile. We evaluate the phylogenetic relationships within this tribe through parsimony and Bayesian approaches based on 99 morphological characters (including 19 integumental characters, 38 skull characters, 31 dental characters, three postcranial skeletal characters, seven from the male accessory glands and phallus and one from the digestive system) and six molecular markers (one mitochondrial and five nuclear). We include representatives of all, except one, of the currently recognized species of living Abrotrichini plus one fossil form. Based on total evidence, we recovered a primary division between the genus Abrothrix and a group including the long‐clawed Abrotrichini, Chelemys, Geoxus, Notiomys and Pearsonomys. Both clades are recognized and named here as subtribes. The large degree of morphological variation observed within Abrothrix suggests that species in the genus fall into four groups, which we recognize as subgenera. In addition, the two known species of Chelemys do not form a monophyletic group, and Geoxus was recovered as paraphyletic with respect to Pearsonomys. To reconcile classification and phylogenetics, we describe a new genus for Chelemys macronyx and include Pearsonomys as a junior synonym of Geoxus. Our results highlight the importance of both morphology and molecules in resolving the phylogenetic relationships within this tribe. Based on biogeographical analyses, we hypothesize that Abrotrichini originated in south‐western South America by vicariance and then diversified mostly by successive dispersal events.     

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 .     

Radiation of the Australian Salicornioideae (Chenopodiaceae)--based on evidence from nuclear and chloroplast DNA sequences

In phylogenetic analyses of nuclear ITS and chloroplast trnL DNA sequences, the mostly endemic Australian genera; Halosarcia, Pachycornia, Sclerostegia, Tecticornia, and Tegicornia of the subfamily Salicornioideae (Chenopodiaceae) together form a monophyletic group, congruent with the hypothesis that they evolved from a common ancestor. However, limited genetic differentiation evident in both nrDNA and cpDNA sequences among these taxa suggests a possible rapid radiation. Based on fossil pollen records and climatic models of other authors, it is hypothesized that the expansion of the Australian endemic Salicornioideae most likely occurred during the Late Miocene to Pliocene, when increasing aridity caused the formation of extensive salt lakes along endorheic paleodrainage channels. Moreover, Australian Sarcocornia representatives were supported as monophyletic, nested within a paraphyletic Sarcocornia clade that also comprised European Salicornia in the ITS analysis. This suggests that Sarcocornia arrived in Australia subsequent to the ancestor of the Australian endemic genera most likely via long-distance dispersal.     

Phylogeny ofRubiaceae-Rubieae inferred from the sequence of a cpDNA intergene region

A phylogenetic analysis of 25 species, representing eight genera of theRubieae tribe (Rubiaceae), has been made using the DNA sequence of the chloroplastatp B-rbc L intergene region. Six tropical genera from other tribes ofRubiaceae have been used as outgroups. Whatever the method of analysis (distance, parsimony or maximum likelihood), five groups are clearly separated and described as informal clades. Their relative relationships are not clearly resolved by the parsimony analysis, resulting in eight equally parsimonious trees, 327 steps long, with a consistency index (CI) of 0.749 (excluding uninformative sites). TheRubieae tribe appears monophyletic from the data available. Some new and partly unexpected phylogenetic relationships are suggested. The genusRubia forms a separate clade and appears to be the relatively advanced sister group of the remaining taxa. TheSherardia clade also includes the generaCrucianella andPhuopsis. Galium sect.Aparinoides appears closely attached to theAsperula sect.Glabella clade. The remaining taxa ofGalium are paraphyletic:Galium sect.Platygalium (in theCruciata clade) is linked to the advanced generaCruciata andValantia; the more apomorphic groups ofGalium form theGalium sect.Galium clade, including the perennial sectionsGalium, Leiogalium, andLeptogalium as well as the annual (and possibly polyphyletic) sect.Kolgyda.     

Higher‐level phylogeny of diving beetles (Coleoptera: Dytiscidae) based on larval characters

A comprehensive higher‐level phylogeny of diving beetles (Dytiscidae) based on larval characters is presented. Larval morphology and chaetotaxy of a broad range of genera and species was studied, covering all currently recognized subfamilies and tribes except for the small and geographically restricted Hydrodytinae, where the larva is unknown. The results suggest several significant conclusions with respect to the systematics of Dytiscidae including the following: monophyly of all currently recognized subfamilies, although Dytiscinae when considered in a broad context is rendered paraphyletic by Cybistrinae; currently recognized tribes are monophyletic except for Agabini, Hydroporini and Laccornellini; inter‐subfamily and inter‐tribe relationships generally show weak support, except for a few well supported clades; three distinct clades are recognized within Dytiscinae [Dytiscini sensu lato (i.e. including the genera Dytiscus Linnaeus and Hyderodes Hope), Hydaticini sensu lato, and Cybistrini]; and recognition of Pachydrini as a distinct tribe. Other less robust results include: Methlini sister to the rest of Hydroporinae; relative basal position of Laccornini, Hydrovatini and Laccornellini within Hydroporinae; close relationship of Agabinae and Copelatinae; Matinae nested deep within Dytiscidae, as sister to a large clade including Colymbetinae, Coptotominae, Lancetinae and Dytiscinae sensu lato; the sister‐group relationship of Agabetini and Laccophilini is confirmed. The results presented here are discussed and compared with previous phylogenetic hypotheses based on different datasets, and the evolution of some significant morphological features is discussed in light of the proposed phylogeny. All suprageneric taxa are diagnosed, including illustrations of all relevant synapomorphies, and a key to separate subfamilies and tribes is presented, both in traditional (paper) format and as an online Lucid interactive identification key.     

Phylogenetic analysis of higher-level relationships of Odonata

Abstract. This is the most comprehensive analysis of higher‐level relationships in Odonata conducted thus far. The analysis was based on a detailed study of the skeletal morphology and wing venation of adults, complemented with a few larval characters, resulting in 122 phylogenetically informative characters. Eighty‐five genera from forty‐five currently recognized families and subfamilies were examined. In most cases, several species were chosen to serve as exemplars for a given genus. The seven fossil outgroup taxa included were exemplar genera from five successively more distant odonatoid orders and suborders: Tarsophlebiidae (the closest sister group of Odonata, previously placed as a family within ‘Anisozygoptera’), Archizygoptera, Protanisoptera, Protodonata and Geroptera. Parsimony analysis of the data, in which characters were treated both under equal weights and implied weighting, produced cladograms that were highly congruent, and in spite of considerable homoplasy in the odonate data, many groupings in the most parsimonious cladograms were well supported in all analyses, as indicated by Bremer support. The analyses supported the monophyly of both Anisoptera and Zygoptera, contrary to the well known hypothesis of zygopteran paraphyly. Within Zygoptera, two large sister clades were indicated, one comprised of the classical (Selysian) Calopterygoidea, except that Amphipterygidae, which have traditionally been placed as a calopterygoid family, nested within the other large zygopteran clade comprised of Fraser's ‘Lestinoidea’ plus ‘Coenagrionoidea’ (both of which were shown to be paraphyletic as currently defined). Philoganga alone appeared as the sister group to the rest of the Zygoptera in unweighted cladograms, whereas Philoganga + Diphlebia comprised the sister group to the remaining Zygoptera in all weighted cladograms. ‘Anisozygoptera’ was confirmed as a paraphyletic assemblage that forms a ‘grade’ towards the true Anisoptera, with Epiophlebia as the most basal taxon. Within Anisoptera, Petaluridae appeared as the sister group to other dragonflies.     

Phylogenetic relationships among the phyllotini (rodentia: Sigmodontinae) using morphological characters

Thirty-three species representing all 14 genera of the South American rodent tribe Phyllotini and 5 problematic genera are surveyed for 96 multistate and binary dental, cranial, skeletal, external, and male reproductive tract characters. Wagner parsimony analysis confirmsCalomys as the most basal phyllotine genus, and as currently constituted it is likely paraphyletic. The results are consistent with the exclusion ofPseudoryzomys from the phyllotines and the separation ofReithrodon andNeotomys fromHolochilus at the tribal level. Several highly differentiated generic groups that include a radiation of altiplano endemics centered onAuliscomys and the largely southern Andean/PatagonianReithrodon group appear to form a clade. AGraomys generic group that includesAndalgalomys andEligmodontia is also apparent, but its relationships to other phyllotines are obscured by poorly resolved internal nodes in the more species-rich and probably paraphyletic genusPhyllotis. The significance and consequences of more intensive taxonomic sampling are discussed. The taxonomic consequences of the phylogeny are presented.     

The gastropod–crustacean connection: towards the phylogeny and evolution of the parasitic copepod family Splanchnotrophidae

Amongst the most significant metazoan taxa associated with gastropod molluscs is the endoparasitic copepod family Splanchnotrophidae. Currently it contains five genera with highly modified morphology and exclusively infesting nudibranch and sacoglossan sea slug hosts. The present study is a first approach towards reconstructing their phylogeny and evolution. Cladistic analysis of 109 morphological characters including 24 known splanchnotrophid species resulted in a fully resolved strict consensus tree that is discussed in morphological, functional, and geographical frameworks. Alternative topologies are also explored. Originating from paraphyletic Philoblennidae, the Splanchnotrophidae emerge as sister group to the genus Briarella. Unique synapomorphies, such as the bizarre body shapes and successive reduction of mouthparts, are discussed as adaptive traits to endoparasitism that evolved only once within copepods infesting shell‐less heterobranch gastropods. The ancestrally Indo‐Pacific Splanchnotrophidae split up into a clade of the still Indo‐Pacific genera Ceratosomicola and Arthurius, sister to a clade composed of the monophyletic amphi‐American genus Ismaila and European Splanchnotrophus emerging from paraphyletic Lomanoticola. Although initial radiation of Briarella and Splanchnotrophidae is likely to have involved chromodoridid nudibranch hosts, later phylogenies of parasites and their hosts are incongruent; intriguingly, host shifts from nudibranch to only distantly related sacoglossan species occurred at least two times independently. Such remarkable ecological plasticity is assumed to have driven splanchnotrophid diversification. Topological hypotheses and historical biogeographical and evolutionary scenarios inferred herein can be tested by future molecular research. © 2013 The Linnean Society of London     

New fossil species of ommatids (Coleoptera: Archostemata) from the Middle Mesozoic of China illuminating the phylogeny of Ommatidae

ABSTRACT: BACKGROUND: Ommatidae is arguably the "most ancestral" extant beetle family. Recent species of this group are only found in South America and Australia, but the fossil record reveals a much broader geographical distribution in the Mesozoic. Up to now, thirteen fossil genera with more than 100 species of ommatids have been described. However, the systematic relationships of the extant and extinct Ommatidae have remained obscure. RESULTS: In this study, four new species, Pareuryomma ancistrodonta sp. nov., Pareuryomma cardiobasis sp. nov., Omma delicata sp. nov., and Tetraphalerus decorosus sp. nov., are described. Based on well-preserved fossil specimens and previously published data the phylogenetic relationships of extant and extinct lineages of Ommatidae were analyzed for the first time cladistically. Based on the results we propose a new classification with six tribes of Ommatidae: Pronotocupedini, Notocupedini, Lithocupedini, Brochocoleini, Ommatini and Tetraphalerini. These taxa replace the traditional four subfamilies. CONCLUSION: There is good support for the monophyly of the ingroup. Notocupedini, as defined by Ponomarenko, are paraphyletic. Notocupoides + Eurydictyon are the sister group of the remaining fossil and extant ommatids. Together they form the clade Pronotocupedini. Notocupedini and Lithocupedini are the next two branches. The tribe Brochocoleini is the sister group of a clade comprising Tetraphalerini and Ommatini.