Ingroup Species Research Articles

A family-level Tree of Life for bivalves based on a Sanger-sequencing approach

The systematics of the molluscan class Bivalvia are explored using a 5-gene Sanger-based approach including the largest taxon sampling to date, encompassing 219 ingroup species spanning 93 (or 82%) of the 113 currently accepted bivalve families. This study was designed to populate the bivalve Tree of Life at the family level and to place many genera into a clear phylogenetic context, but also pointing to several major clades where taxonomic work is sorely needed. Despite not recovering monophyly of Bivalvia or Protobranchia—as in most previous Sanger-based approaches to bivalve phylogeny—our study provides increased resolution in many higher-level clades, and supports the monophyly of Autobranchia, Pteriomorphia, Heteroconchia, Palaeoheterodonta, Heterodonta, Archiheterodonta, Euheterodonta, Anomalodesmata, Imparidentia, and Neoheterodontei, in addition to many other lower clades. However, deep nodes within some of these clades, especially Pteriomorphia and Imparidentia, could not be resolved with confidence. In addition, many families are not supported, and several are supported as non-monophyletic, including Malletiidae, Nuculanidae, Yoldiidae, Malleidae, Pteriidae, Arcidae, Propeamussiidae, Iridinidae, Carditidae, Myochamidae, Lyonsiidae, Pandoridae, Montacutidae, Galeommatidae, Tellinidae, Semelidae, Psammobiidae, Donacidae, Mactridae, and Cyrenidae; Veneridae is paraphyletic with respect to Chamidae, although this result appears to be an artifact. The denser sampling however allowed testing specific placement of species, showing, for example, that the unusual Australian Plebidonax deltoides is not a member of Donacidae and instead nests within Psammobiidae, suggesting that major revision of Tellinoidea may be required. We also showed that Cleidothaerus is sister group to the cementing member of Myochamidae, suggesting that Cleidothaeridae may not be a valid family and that cementation in Cleidothaerus and Myochama may have had a single origin. These results highlight the need for an integrative approach including as many genera as possible, and that the monophyly and relationships of many families require detailed reassessment. NGS approaches may be able to resolve the most recalcitrant nodes in the near future.

Application of RNA-seq for mitogenome reconstruction, and reconsideration of long-branch artifacts in Hemiptera phylogeny.

Hemiptera make up the largest nonholometabolan insect assemblage. Despite previous efforts to elucidate phylogeny within this group, relationships among the major sub-lineages remain uncertain. In particular, mitochondrial genome (mitogenome) data are still sparse for many important hemipteran insect groups. Recent mitogenomic analyses of Hemiptera have usually included no more than 50 species, with conflicting hypotheses presented. Here, we determined the nearly complete nucleotide sequence of the mitogenome for the aphid species of Rhopalosiphum padi using RNA-seq plus gap filling. The 15,205 bp mitogenome included all mitochondrial genes except for trnF. The mitogenome organization and size for R. padi are similar to previously reported aphid species. In addition, the phylogenetic relationships for Hemiptera were examined using a mitogenomic dataset which included sequences from 103 ingroup species and 19 outgroup species. Our results showed that the seven species representing the Aleyrodidae exhibit extremely long branches, and always cluster with long-branched outgroups. This lead to the failure of recovering a monophyletic Hemiptera in most analyses. The data treatment of Degen-coding for protein-coding genes and the site-heterogeneous CAT model show improved suppression of the long-branch effect. Under these conditions, the Sternorrhyncha was often recovered as the most basal clade in Hemiptera.

Planthopper family Issidae (Insecta: Hemiptera: Fulgoromorpha): linking molecular phylogeny with classification

A molecular phylogeny of the planthopper family Issidae (Hemiptera, Fulgoroidea) is provided using both Maximum likelihood and Bayesian inference analyses. The phylogeny is based on 18S, two parts of 28S, COXI and Cytb genes from 50 genera and 79 ingroup species (including 8 species recently excluded from Issidae). As with the only few previous studies, an important taxonomic impediment is observed with the sampling; however for the first time, all analyses depict several fully supported lineages, which challenge the recent proposed “modern classification” of the family. It also highlights a strong coherence between these lineages and their respective geographical distribution. All previously excluded taxa are confirmed as not being part of the Issidae as recently defined which monophyly is confirmed. Accordingly, a new classification of the family is proposed with 3 subfamilies and 7 tribes as follows. Neotropical issid Thioniini in Thioniinae stat. rev. is re-established as an independent lineage sister to all other Issidae. Palaearctic Issidae are weakly supported as a monophyletic lineage, Issinae stat. nov., including 2 tribes: Issini stat. nov. (genera Issus and Latissus) and Hysteropterini stat. rev. (all other Palaearctic genera). Oriental Issidae form a strongly supported monophyletic subfamily group Hemisphaeriinae stat. rev. including 4 tribes: Kodaianellini trib. nov., Sarimini trib. nov., Parahiraciini Cheng & Yang, 1991, and Hemisphaeriini Melichar, 1906, the latter including 2 subtribes: Mongolianina s.trib. nov., and Hemisphaeriina Melichar, 1906. A Neotropical lineage including the genus Picumna is provisionally placed in incertae sedis within the Hemisphaeriinae stat. nov.

Maximum Likelihood Implementation of an Isolation-with-Migration Model for Three Species.

We develop a maximum likelihood (ML) method for estimating migration rates between species using genomic sequence data. A species tree is used to accommodate the phylogenetic relationships among three species, allowing for migration between the two sister species, while the third species is used as an out-group. A Markov chain characterization of the genealogical process of coalescence and migration is used to integrate out the migration histories at each locus analytically, whereas Gaussian quadrature is used to integrate over the coalescent times on each genealogical tree numerically. This is an extension of our early implementation of the symmetrical isolation-with-migration model for three species to accommodate arbitrary loci with two or three sequences per locus and to allow asymmetrical migration rates. Our implementation can accommodate tens of thousands of loci, making it feasible to analyze genome-scale data sets to test for gene flow. We calculate the posterior probabilities of gene trees at individual loci to identify genomic regions that are likely to have been transferred between species due to gene flow. We conduct a simulation study to examine the statistical properties of the likelihood ratio test for gene flow between the two in-group species and of the ML estimates of model parameters such as the migration rate. Inclusion of data from a third out-group species is found to increase dramatically the power of the test and the precision of parameter estimation. We compiled and analyzed several genomic data sets from the Drosophila fruit flies. Our analyses suggest no migration from D. melanogaster to D. simulans, and a significant amount of gene flow from D. simulans to D. melanogaster, at the rate of ~0.02 migrant individuals per generation. We discuss the utility of the multispecies coalescent model for species tree estimation, accounting for incomplete lineage sorting and migration.

Phylogeography of the Subgenus Drosophila (Diptera: Drosophilidae): Evolutionary History of Faunal Divergence between the Old and the New Worlds

The current subgenus Drosophila (the traditional immigrans-tripunctata radiation) includes major elements of temperate drosophilid faunas in the northern hemisphere. Despite previous molecular phylogenetic analyses, the phylogeny of the subgenus Drosophila has not fully been resolved: the resulting trees have more or less varied in topology. One possible factor for such ambiguous results is taxon-sampling that has been biased towards New World species in previous studies. In this study, taxon sampling was balanced between Old and New World species, and phylogenetic relationships among 45 ingroup species selected from ten core species groups of the subgenus Drosophila were analyzed using nucleotide sequences of three nuclear and two mitochondrial genes. Based on the resulting phylogenetic tree, ancestral distributions and divergence times were estimated for each clade to test Throckmorton’s hypothesis that there was a primary, early-Oligocene disjunction of tropical faunas and a subsequent mid-Miocene disjunction of temperate faunas between the Old and the New Worlds that occurred in parallel in separate lineages of the Drosophilidae. Our results substantially support Throckmorton’s hypothesis of ancestral migrations via the Bering Land Bridge mainly from the Old to the New World, and subsequent vicariant divergence of descendants between the two Worlds occurred in parallel among different lineages of the subgenus Drosophila. However, our results also indicate that these events took place multiple times over a wider time range than Throckmorton proposed, from the late Oligocene to the Pliocene.

Phylogeny and revision of a colorful Neotropical genus of rove beetles: Xenopygus Bernhauer (Coleoptera: Staphylinidae).

Xenopygus Bernhauer is one of the most colorful staphylinid beetles and widespread in the Neotropical region. The aim of this study was to test the monophyly of Xenopygus based on adult morphology and to review the current species. Cladistic analysis was performed with six ingroup species, including two new species. Xenopygus is a monophyletic group supported by: antennomere V as wide as long to slightly wider, antennomere VII two times wider than long and superior line of pronotal hypomere developed, continuous on the anterior angle of pronotum. Xenopygus is composed of six species, four previously described and two new species, with the following topology: ((X. analis+(X. bicolor+X. confusus))+(X. cordovensis+(X. sancticamillus, sp. nov.+ X. petilicolis, sp. nov.))). Xenopygus is the sister group to a clade formed by species of Dysanellus and Xanthopygus, corroborating previous phylogenetic studies. The genus, and all its species were redescribed, an identification key was produced and illustration diagnostic plates and distribution maps were also provided.

An evaluation of transcriptome-based exon capture for frog phylogenomics across multiple scales of divergence (Class: Amphibia, Order: Anura).

Custom sequence capture experiments are becoming an efficient approach for gathering large sets of orthologous markers in nonmodel organisms. Transcriptome-based exon capture utilizes transcript sequences to design capture probes, typically using a reference genome to identify intron-exon boundaries to exclude shorter exons (<200bp). Here, we test directly using transcript sequences for probe design, which are often composed of multiple exons of varying lengths. Using 1260 orthologous transcripts, we conducted sequence captures across multiple phylogenetic scales for frogs, including outgroups ~100Myr divergent from the ingroup. We recovered a large phylogenomic data set consisting of sequence alignments for 1047 of the 1260 transcriptome-based loci (~561000bp) and a large quantity of highly variable regions flanking the exons in transcripts (~70000bp), the latter improving substantially by only including ingroup species (~797000bp). We recovered both shorter (<100bp) and longer exons (>200bp), with no major reduction in coverage towards the ends of exons. We observed significant differences in the performance of blocking oligos for target enrichment and nontarget depletion during captures, and differences in PCR duplication rates resulting from the number of individuals pooled for capture reactions. We explicitly tested the effects of phylogenetic distance on capture sensitivity, specificity, and missing data, and provide a baseline estimate of expectations for these metrics based on a priori knowledge of nuclear pairwise differences among samples. We provide recommendations for transcriptome-based exon capture design based on our results, cost estimates and offer multiple pipelines for data assembly and analysis.

The borage family (Boraginaceae s.str.): A revised infrafamilial classification based on new phylogenetic evidence, with emphasis on the placement of some enigmatic genera

AbstractBoraginaceae s.str. is a subcosmopolitan family of 1600 to 1700 species in around 90 genera, and recent phylogenetic studies indicate that the infrafamilial classification as currently used is highly obsolete. The present study addresses the relationships of the major clades in Boraginaceae s.str. with an emphasis on monophyly of, and relationships between previously recognized clades and the position of various unplaced genera such as Afrotysonia, Anoplocaryum, Brachybotrys, Chionocharis, Craniospermum, Thyrocarpus, and Trigonocaryum using three plastid markers and a taxon sampling with four outgroup and 170 ingroup species from 73 genera. The phylogeny shows high statistical support for most nodes on the backbone and within individual clades. Echiochileae are confirmed as sister to the remainder of Boraginaceae s.str., which, in turn, fall into two well‐supported clades, the Boragineae + Lithospermeae and the Cynoglosseae s.l. The latter is highly resolved and includes the Lasiocaryum‐clade (Chionocharis, Lasiocaryum, Microcaryum) and the Trichodesmeae (Caccinia, Trichodesma) as sister to the remainder of the group. Rochelieae (formerly the Eritrichieae s.str., also including Eritrichium, Hackelia, and Lappula) form a poorly supported polytomy together with the Mertensia‐clade (also including Anoplocaryum, Asperugo, and Memoremea) and the Omphalodes‐clade. The enigmatic genus Craniospermum (Craniospermeae) is sister to an expanded Myosotideae (also including Brachybotrys, Decalepidanthus, Trigonocaryum, and Trigonotis) and these two clades are in turn sister to the Core‐Cynoglosseae, in which Afrotysonia glochidiata and Thyrocarpus sampsonii are included. Core‐Cynoglosseae again fall into two pairs of well‐supported subclades. The majority of generic placements are now resolved satisfactorily and the remaining phylogenetic questions can be clearly delimited. Based on the extensive phylogenetic data now available we propose a new infrafamilial classification into three subfamilies and 11 tribes, representing a consensus among the participating authors, according to which major clades are renamed.

Untangling the phylogeny of Leandra s.str. (Melastomataceae, Miconieae)

Phylogenetic studies in Melastomataceae have demonstrated the need for taxonomic rearrangements in the current classification. Nonetheless, melastomes are among the most diverse groups of plants and several cases of known artificial taxa have been observed and awaiting further resolution. The Leandra s.str. clade, with ca. 200 species, includes the majority of the taxa traditionally treated in the genus Leandra and is almost restricted to eastern Brazil. In earlier studies, some attempts have been made to infer the relationships within Leandra s.str., but the sampling was sparse and the resolution low inside the clade. Here, we attempt to provide an improved phylogenetic hypothesis for this group on which to base further studies. Specifically, we provide a comprehensive taxon sampling and attempt to infer a species tree for this group, dissecting potential noise in the phylogenetic reconstruction, such as paralogy, rogue taxa, hybridization and incomplete lineage sorting. Our data set includes 126 ingroup species (192 terminals) and four partitions (six markers). We implement the ∗BEAST model for species tree inference and perform several simulation methods to assess model fit and to discuss potential causes for the observed patterns. Major lineages of Leandra s.str. were delineated, a strictly bifurcating species tree model seems to not account for the observed data, and hybridization is very likely an important evolutionary force in this group.

Ingroup Bias and Ingroup Projection in the Furry Fandom

&lt;p&gt;In two studies, we examine ingroup favoritism within the furry community. Furry fans construct personas (“fursonas”) that are often related to a species of animal (e.g., fox, wolf). In Study 1, furries were found to rate their fursona species more favorably than other species. In Study 2, we examined whether the ingroup projection model may aid in understanding this ingroup species bias. Participants’ ratings of prototypicality of their species within the fandom was associated with greater favoritism of one’s species. A serial mediation model showed that identification with the fursona and perceived benefits of one’s fursona mediated the association between ingroup projection and ingroup species bias.&lt;/p&gt;

Recircumscription ofOxera(Lamiaceae: Ajugoideae) to includeFaradayabased on molecular and anatomical data

Oxera is a genus of c. 20 species from New Caledonia and Vanuatu. Its phylogenetic placement in Lamiaceae (Ajugoideae) remained uncertain until studies showed close affinities with the large pantropical genus Clerodendrum, the Indo-Pacific Faradaya (three species) and the monotypic Hosea from Borneo. We reassessed the relationships of Oxera with these genera by building a phylogenetic framework based on six plastid and six nuclear loci and on a sampling including 30 ingroup species. The results show that Clerodendrum and its closely related genera are sister to the alliance encompassing Faradaya, Hosea and Oxera. Hosea is sister to all species of Oxera and Faradaya. Faradaya is recovered as polyphyletic as it is partly nested within Oxera. Extensive morphological variability is observed in Oxera. However, Oxera also shares strong similarities with the three Faradaya species. Polyphyletic Faradaya is therefore placed in synonymy with Oxera and we propose three new combinations: O. amicorum, O. lehuntei and O. splendida. A unique synapomorphy supports this new circumscription of Oxera: the petiole vascularization consisting of several small vascular bundles arranged in an interrupted full circle (vs. a large bundle in a half circle in other ajugoids). Oxera thus comprises c. 37 species, with a diversity centre in New Caledonia. Biogeographical processes, plastid paralogy and vascularization shift are discussed. © 2015 The Linnean Society of London, Botanical Journal of the Linnean Society, 2015, 179, 693–711.

A morphological supermatrix-based phylogeny for the Neotropical fish superfamily Anostomoidea (Ostariophysi: Characiformes): phylogeny, missing data and homoplasy.

Although 11 studies have addressed the systematics of the four families and 281 fish species of the ecomorphologically diverse Anostomoidea, none has proposed a global hypothesis of relationships. We synthesized these studies to yield a supermatrix with 463 morphological characters among 174 ingroup species, and inferred phylogeny with parsimony and Bayesian optimization. We evaluated the applicability of the supermatrix approach to morphological datasets, tested its sensitivity to missing data, determined the impact of homoplastic characters on phylogenetic resolution, and determined the distribution of homologies and homoplasies on the topology. Despite more than 60% missing data, analyses supported the monophyly of all families, and phylogenetic structure degraded only with inclusion of species with high percentages of missing data and in analyses limited to homoplasies. The latter differs modestly from the full matrix indicating phylogenetic signal in homoplastic characters. Character distributions differ across the phylogeny, with a greater prevalence of homologies at deeper nodes and homoplasies nearer the tips than expected by chance. This may suggest early diversification into distinct bauplans with subsequent diversification of faster evolving character systems. The morphological supermatrix approach is powerful and allows integration of classical data with modern methods to examine the evolution of multiple character systems.

Resolving the phylogeny of a speciose spider group, the family Linyphiidae (Araneae)

For high-level molecular phylogenies, a comprehensive sampling design is a key factor for not only improving inferential accuracy, but also for maximizing the explanatory power of the resulting phylogeny. Two standing problems in molecular phylogenies are the unstable placements of some deep and long branches, and the phylogenetic relationships shown by robust supported clades conflict with recognized knowledge. Empirical and theoretical studies suggest that increasing taxon sampling is expected to ameliorate, if not resolve, both problems; however, sometimes neither the current taxonomic system nor the established phylogeny can provide sufficient information to guide additional sampling design. We examined the phylogeny of the spider family Linyphiidae, and selected ingroup species based on epigynal morphology, which can be reconstructed in a phylogenetic context. Our analyses resulted in seven robustly supported clades within linyphiids. The placements of four deep and long branches are sensitive to variations in both outgroup and ingroup sampling, suggesting the possibility of long branch attraction artifacts. Results of ancestral state reconstruction indicate that successive state transformations of the epigynal plate are associated with early cladogenetic events in linyphiid diversification. Representatives of different subfamilies were mixed together within well supported clades and examination revealed that their defining characters, as per traditional taxonomy, are homoplastic. Furthermore, our results demonstrated that increasing taxon sampling produced a more informative framework, which in turn helps to study character evolution and interpret the relationships among linyphiid lineages. Additional defining characters are needed to revise the linyphiid taxonomic system based on our phylogenetic hypothesis.

DNA barcoding supports identification of Malacobdella species (Nemertea: Hoplonemertea).

Nemerteans of the genus Malacobdella live inside of the mantle cavity of marine bivalves. The genus currently contains only six species, five of which are host-specific and usually found in a single host species, while the sixth species, M. grossa, has a wide host range and has been found in 27 different bivalve species to date. The main challenge of Malacobdella species identification resides in the similarity of the external morphology between species (terminal sucker, gut undulations number, anus position and gonad colouration), and thus, the illustrations provided in the original descriptions do not allow reliable identification. In this article, we analyse the relationships amongthree species of Malacobdella:M.arrokeana,M.japonica andM.grossa,adding new data for the M.grossa and reporting the first for M. japonica, analysing 658 base pairs of the mitochondrial cytochrome c oxidase subunit I gene(COI).Based on these analyses, we present and discuss the potential of DNA barcoding for Malacobdellaspecies identification. Sixty-four DNA barcoding fragments of the mitochondrial COI gene from three different Malacobdella species (M. arrokeana, M. japonica and M. grossa) are analysed (24 of them newly sequenced for this study, along with four outgroup specimens) and used to delineate species. Divergences, measured as uncorrected differences, between the three species were M.arrokeana-M. grossa11.73%,M.arrokeana-M.japonica 10.62%and M.grossa-M. japonica 10.97%. The mean intraspecific divergence within the ingroup species showed a patent gap with respect to the interspecific ones: 0.18% for M.arrokeana,0.13% for M.grossa and0.02% for M.japonica (rangesfrom 0 to 0.91%). We conclude that there is a clear correspondence between the molecular data and distinguishing morphological characters. Our results thus indicate that some morphological characters are useful for species identification and support the potential of DNA barcoding for species identification in a taxonomic group with subtle morphological external differences.

Systematics of some Late Ordovician encrinurine trilobites from Laurentian North America

Classification and relationships of the Ordovician encrinurines Frencrinoides Lespérance and Desbiens and Walencrinoides Lespérance and Desbiens are poorly understood, with little evidence for monophyly of either genus. We revise the type species of both genera, F. capitonis (Frederickson) and W. rarus (Walcott), using new and archival material. We explore their species composition and phylogenetic relationships with a parsimony analysis that includes 17 well-documented ingroup species that can be coded readily, and which is rooted with Encrinuroides regularis Parnaste, the oldest known encrinurine. The results support monophyly of Frencrinuroides and Walencrinuroides, albeit with more limited species membership than proposed by Lespérance and Desbiens. Previous suggestions that both E. uncatus Evitt and Tripp and E. neuter Evitt and Tripp should be assigned to Erratencrinurus Kruger are also supported by our analysis, as is monophyly of Physemataspis Evitt and Tripp. New species are W. rolfi and W. tremblayi.

Large‐scale molecular phylogeny of metallic wood‐boring beetles (Coleoptera: Buprestoidea) provides new insights into relationships and reveals multiple evolutionary origins of the larval leaf‐mining habit

AbstractThe family Buprestidae (jewel beetles or metallic wood‐boring beetles), contains nearly 15 000 species in 522 genera. Together with the small family Schizopodidae (seven species, three genera), they form the superfamily Buprestoidea. Adult Buprestoidea feed on flowers or foliage, whereas larvae are mostly internal feeders, boring in roots or stems, or mining the leaves of woody or herbaceous plants. The subfamilial and tribal classification of Buprestoidea remains unsettled, with substantially different schemes proposed by different workers based on morphology. Here we report the first large‐scale molecular phylogenetic study of the superfamily Buprestoidea based on data from four genes for 141 ingroup species. We used these data to reconstruct higher‐level relationships and to assess the current classification and the origins of the larval leaf‐mining habit within Buprestoidea. In our analyses, the monophyly of Buprestoidea was strongly supported, as was the monophyly of Schizopodidae and its placement sister to Buprestidae. Our results are largely consistent with the generally accepted major lineages of buprestoids, including clearly‐defined agrilines, buprestines–chrysochroines and early‐branching julodines–polycestines. In addition to Schizopodidae, three of the six subfamilies were monophyletic in our study: Agrilinae, Julodinae and the monogeneric Galbellinae (Galbella). Polycestinae was monophyletic with the exception of the enigmatic Haplostethini. Chrysochroinae and Buprestinae were not monophyletic, but were recovered together in a large mixed clade along with Galbella. The interrelationships of Chrysochroinae and Buprestinae were not well resolved; however they were clearly polyphyletic, with chrysochroine genera falling into several different well‐supported clades otherwise comprising buprestine genera. All Agrilinae were contained in a single strongly supported clade. Coraebini were dispersed throughout Agrilinae, with strong nodal support for several clades representing subtribes. Neither Agrilini nor Tracheini were monophyletic. The leaf‐mining genus Paratrachys (Paratracheini) was recovered within the Acmaeoderioid clade, consistent with the current classification, and confirming the independent origins of leaf‐mining within Polycestinae and Agrilinae. Additionally, our results strongly suggest that the leaf‐mining agriline tribe Tracheini is polyphyletic, as are several of its constituent subtribes. External root feeding was likely the ancestral larval feeding habit in Buprestoidea. The apparent evolutionary transitions to internal feeding allowed access to a variety of additional plant tissues, including leaves. Interestingly, the several genera of leaf‐mining agrilines do not form a monophyletic group. Many of these genera are diverse and highly specialized, possibly indicating adaptive radiations.

Morphometric study of phylogenetic and ecologic signals in procyonid (mammalia: carnivora) endocasts.

Endocasts provide a proxy for brain morphology but are rarely incorporated in phylogenetic analyses despite the potential for new suites of characters. The phylogeny of Procyonidae, a carnivoran family with relatively limited taxonomic diversity, is not well resolved because morphological and molecular data yield conflicting topologies. The presence of phylogenetic and ecologic signals in the endocasts of procyonids will be determined using three-dimensional geometric morphometrics. Endocasts of seven ingroup species and four outgroup species were digitally rendered and 21 landmarks were collected from the endocast surface. Two phylogenetic hypotheses of Procyonidae will be examined using methods testing for phylogenetic signal in morphometric data. In analyses of all taxa, there is significant phylogenetic signal in brain shape for both the morphological and molecular topologies. However, the analyses of ingroup taxa recover a significant phylogenetic signal for the morphological topology only. These results indicate support for the molecular outgroup topology, but not the ingroup topology given the brain shape data. Further examination of brain shape using principal components analysis and wireframe comparisons suggests procyonids possess more developed areas of the brain associated with motor control, spatial perception, and balance relative to the basal musteloid condition. Within Procyonidae, similar patterns of variation are present, and may be associated with increased arboreality in certain taxa. Thus, brain shape derived from endocasts may be used to test for phylogenetic signal and preliminary analyses suggest an association with behavior and ecology.

Phylogeny and evolution of pharmacophagy in tiger moths (Lepidoptera: Erebidae: Arctiinae).

The focus of this study was to reconstruct a phylogenetic hypothesis for the moth subfamily Arctiinae (tiger moths, woolly bears) to investigate the evolution of larval and adult pharmacophagy of pyrrolizidine alkaloids (PAs) and the pathway to PA chemical specialization in Arctiinae. Pharmacophagy, collection of chemicals for non-nutritive purposes, is well documented in many species, including the model species Utetheisa ornatrix L. A total of 86 exemplar ingroup species representing tiger moth tribes and subtribes (68 genera) and nine outgroup species were selected. Ingroup species included the most species-rich generic groups to represent the diversity of host-plant associations and pharmacophagous behaviors found throughout Arctiinae. Up to nine genetic markers were sequenced: one mitochondrial (COI barcode region), one nuclear rRNA (D2 region, 28S rRNA), and seven nuclear protein-coding gene fragments: elongation factor 1-α protein, wingless, ribosomal protein subunit S5, carbamoylphosphate synthase domain regions, glyceraldehyde-3-phosphate dehydrogenase, isocitrate dehydrogenase and cytosolic malate dehydrogenase. A total of 6984 bp was obtained for most species. These data were analyzed using model-based phylogenetic methods: maximum likelihood (ML) and Bayesian inference (BI). Ancestral pharmacophagous behaviors and obligate PA associations were reconstructed using the resulting Bayes topology and Reconstructing Ancestral States in Phylogenies (RASP) software. Our results corroborate earlier studies on the evolution of adult pharmacophagous behaviors, suggesting that this behavior arose multiple times and is concentrated in the phaegopterine-euchromiine-ctenuchine clade (PEC). Our results suggest that PA specialization may have arisen early in the phylogeny of the subfamily and that facultative larval pharmacophagous behaviors are the derived condition.

A preliminary phylogenetic analysis of the New World Helopini (Coleoptera, Tenebrionidae, Tenebrioninae) indicates the need for profound rearrangements of the classification.

Helopini is a diverse tribe in the subfamily Tenebrioninae with a worldwide distribution. The New World helopine species have not been reviewed recently and several doubts emerge regarding their generic assignment as well as the naturalness of the tribe and subordinate taxa. To assess these questions, a preliminary cladistic analysis was conducted with emphasis on sampling the genera distributed in the New World, but including representatives from other regions. The parsimony analysis includes 30 ingroup species from America, Europe and Asia of the subtribes Helopina and Cylindrinotina, plus three outgroups, and 67 morphological characters. Construction of the matrix resulted in the discovery of morphological character states not previously reported for the tribe, particularly from the genitalia of New World species. A consensus of the 12 most parsimonious trees supports the monophyly of the tribe based on a unique combination of characters, including one synapomorphy. None of the subtribes or the genera of the New World represented by more than one species (Helops Fabricius, Nautes Pascoe and Tarpela Bates) were recovered as monophyletic. Helopina was recovered as paraphyletic in relation to Cylindrinotina. One Nearctic species of Helops and one Palearctic species of Tarpela (subtribe Helopina) were more closely related to species of Cylindrinotina. A relatively derived clade, mainly composed by Neotropical species, was found; it includes seven species of Tarpela, seven species of Nautes, and three species of Helops, two Nearctic and one Neotropical. Our results reveal the need to deeply re-evaluate the current classification of the tribe and subordinated taxa, but a broader taxon sampling and further character exploration is needed in order to fully recognize monophyletic groups at different taxonomic levels (from subtribes to genera).

Morphology-Based Phylogeny and Biogeography of Wockia (Lepidoptera: Urodidae) with Description of a New Species from Japan and South Korea

Wockia, one of the six genera within the lepidopteran family Urodidae, currently includes nine species distributed in Holarctic, Oriental, and Neotropical regions. A new species of the genus, W. magna sp. nov., is described from Japan and South Korea. This is the first record of Urodidae from Japan. A cladistic analysis was constructed based on 27 morphological characters from five ingroup species and two outgroup taxa. A single most parsimonious tree was found (length = 38, CI = 71, RI = 70). The resulting tree failed to recover the monophyly of Wockia chewbacca and a clade Wockia sensu stricto, including four congeners from North America, temperate East Asia and Southeast Asia. Wockia sensu stricto was resolved as monophyletic and was divided into two subgroups, one including W. asperipunctella and W. magna, and the other including W. koreana and W. variata. The sister-group relationship of W. asperipunctella and W. magna was moderately supported. Synapomorphies of Wockia sensu stricto are provided from the resulting phylogeny. Systematic definitions of Wockia and other allied genera are revised. Optimal ancestral area reconstruction implemented in DIVA resulted in different hypotheses for Wockia, depending on how to constrain the maximum number of areas. The maximum area number set as two yielded a more likely scenario suggesting that Wockia sensu stricto originated in temperate East Asia and then dispersed into the Oriental region, North America, and Europe. A working hypothesis and other equally possible alternative explanations for the biogeography of Wockia are provided.