A survey for biochemical synapomorphies to reveal phylogenetic relationships of halichondrid demosponges (Metazoa: Porifera)

Biochemical compounds are a suggested alternative for morphological characters in sponge systematics. The monophyly of the crucial demosponge order Halichondrida is still largely unresolved. Here, we performed a search for synapomorphies, which might aid to define the order and its five families Axinellidae, Bubaridae, Dictyonellidae, Desmoxyidae and Halichondriidae. We also investigated possible links of the order Halichondrida with other demosponge taxa such as suberitids (Hadromerida). Our survey of published compounds revealed hydroxy-nor-sterols as a potential synapomorphy for axinellid taxa, 2,3,6 sulfated sterol nuclei as possible connection between Agelasida and Halichondrida, besides the known pyrrol-2-carboxylic acids and galactosylceramides, furthermore carbonimidic dichlorides, pupukeananes, diterpene isocyanides for the order Halichondrida and linear and cyclic diterpenes for the family Desmoxyidae. The suitability of biochemical characters in systematics is discussed.     

Are monophyly and synapomorphy the same or different? Revisiting the role of morphology in phylogenetics

Species are groups of organisms, marked out by reproductive (replicative) properties. Monophyletic taxa are groups of species, marked out by synapomorphies. In Nelson’s analysis, monophyly and synapomorphy are identical relations. Monophyly and synapomorphy, however, are not equivalent relations. Monophyly is epistemically not accessible, whereas synapomorphy is epistemically accessible through character analysis. Monophyly originates with speciation, the two sister‐species that come into being through the splitting of the ancestral species lineage forming a monophyletic taxon at the lowest level of inclusiveness. Synapomorphy provides the empirical evidence for monophyly, inferred from character analysis in the context of a three‐taxon statement. If synapomorphy and monophyly were equivalent, phylogenetic systematists should find a single tree, instead of multiple equally parsimonious trees. Understanding synapomorphy as the relevant evidence for phylogenetic inference reveals a category mistake in contemporary phylogenetics: the treatment of morphological characters mapped onto molecular trees as synapomorphies and homoplasies. The mapping of morphological characters onto nodes of a molecular tree results in an empirically empty procedure for synapomorphy discovery. Morphological synapomorphies and homoplasies can only be discovered by morphological and combined analyses. The use of morphology in phylogenetic inference in general is defended by examples from Laurales and Squamata in particular. To make empirical evidence scientifically relevant in order to search for concordance, or dis‐concordance, of phylogenetic signal, is certainly more fruitful for phylogenetics than the uncritical mapping of morphological traits on a molecular scaffold. © The Willi Hennig Society 2010.     

Implications from a 28S rRNA gene fragment for the phylogenetic relationships of halichondrid sponges (Porifera: Demospongiae)

Halichondrida is a pivotal demosponge order of which the classification underwent major changes in the recent history. The monophyly of this order and its intra-ordinal phylogeny cannot be reliably determined on the basis of morphology. Here we present a 28Sr RNA gene tree of selected halichondrids, which supports the hypothesis of halichondrid non-monophyly and elucidates further inter-ordinal relationships. We enlarged the analysis by previously published sequences, discuss how previous analyses suffer from taxon bias and analyse the resulting phylogenetic implications. Most halichondrid families (in particular Axinellidae und Dictyonellidae) cluster polyphyletic and the molecular classification of several genera does not agree with the current (morphological) system.     

Molecular phylogeny of the brachyuran crab superfamily Majoidea indicates close congruence with trees based on larval morphology

In this study, we constructed the first molecular phylogeny of the diverse crab superfamily Majoidea (Decapoda: Pleocyemata: Brachyura), using three loci (16S, COI, and 28S) from 37 majoid species. We used this molecular phylogeny to evaluate evidence for phylogenetic hypotheses based on larval and adult morphology. Our study supports several relationships predicted from larval morphology. These include a monophyletic Oregoniidae family branching close to the base of the tree; a close phylogenetic association among the Epialtidae, Pisidae, Tychidae, and Mithracidae families; and some support for the monophyly of the Inachidae and Majidae families. However, not all majoid families were monophyletic in our molecular tree, providing weaker support for phylogenetic hypotheses inferred strictly from adult morphology (i.e., monophyly of individual families). This suggests the adult morphological characters traditionally used to classify majoids into different families may be subject to convergence. Furthermore, trees constructed with data from any single locus were more poorly resolved than trees constructed from the combined dataset, suggesting that utilization of multiple loci are necessary to reconstruct relationships in this group.     

Phylogenetic relationships inCrossostylis (Rhizophoraceae) inferred from restriction site variation of chloroplast DNA

Phylogenetic relationships among the nine species ofCrossostylis (Rhizophoraceae) were elucidated using cladistic analysis of restriction site variations of chloroplast DNA. As a result, this genus was found to comprise two pronounced monophyletic groups as follows:C. biflora, C. grandiflora, C. multiflora andC. sebertii; andC. cominsii, C. pachyantha, C. parksii, C. richii andC. seemannii. Moreover, the monophyly ofC. biflora, C. grandiflora andC. sebertii in the former group and the monophyly ofC. pachyantha, C. parksii, C. richii andC. seemannii in the latter group were also suggested. The molecular tree corresponded well with that inferred from morphological data and no discrepancy was recognized. Many of the floral morphological characters reflected lineage, but all seed coat characters were homoplasious. Evolutionary trends in some morphological characters were optimized on the cpDNA tree obtained. Species from New Caledonia and Polynesia were monophyletic, as were those from the Solomon Islands, Vanuatu and the Fiji Islands. All species endemic to the Fiji Islands made a cluster, and this suggests that speciation occurred from a single ancestral species on the Islands.     

Testing the monophyly of the New Zealand and Australian endemic family Conoesucidae Ross based on combined molecular and morphological data (Insecta: Trichoptera: Sericostomatoidea)

Conoesucidae (Trichoptera, Insecta) are restricted to SE Australia, Tasmania and New Zealand. The family includes 42 described species in 12 genera, and each genus is endemic to either New Zealand or Australia. Although monophyly has been previously assumed, no morphological characters have been proposed to represent synapomorphies for the group. We collected molecular data from two mitochondrial genes (16S and cytochrome oxidase I), one nuclear gene (elongation factor 1-α) (2237–2277 bp in total), and 12 morphological characters to produce the first phylogeny of the family. We combined the molecular and morphological characters and performed both a maximum parsimony analysis and a Bayesian analysis to test the monophyly of the family, and to hypothesize the phylogeny among its genera. The parsimony analysis revealed a single most parsimonious tree with Conoesucidae being a monophyletic taxon and sistergroup to the Calocidae. The Bayesian inference produced a distribution of trees, the consensus of which is supported with posterior probabilities of 100% for 15 out of 22 possible ingroup clades including the most basal branch of the family, indicating strong support for a monophyletic Conoesucidae. The most parsimonious tree and the tree from the Bayesian analysis were identical except that the ingroup genus Pycnocentria changed position by jumping to a neighbouring clade. Based on the assumption that the ancestral conoesucid species was present on both New Zealand and Australia, a biogeographical analysis using the dispersal-vicariance criteria demonstrated that one or two (depending on which of the two phylogenetic reconstructions were applied) sympatric speciation events took place on New Zealand prior to a single, late dispersal from New Zealand to Australia.     

Phylogeny of New Zealand hepialid moths (Lepidoptera: Hepialidae) inferred from a cladistic analysis of morphological data

The phylogeny of the New Zealand hepialid moths was estimated from a cladistic analysis of sixty‐three morphological characters, from all life cycle stages. One hundred and sixteen maximum parsimony trees were produced. The phylogenetic reconstruction indicated that the currently recognized generic concepts, and the four informal lineages hypothesized in a previous morphological taxonomic revision, were monophyletic. The relationships of species within genus Wiseana were not fully resolved. Analysis of a data set of thirty‐nine adult male characters from the New Zealand taxa and the Australian genera Jeana, Oxycanus and Trictena supported the monophyly of the New Zealand ‘Oxycanus’ s.s lineage.     

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

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

A multiple‐gene phylogeny reveals polyphyly among eastern North American Aphaenogaster species (Hymenoptera: Formicidae)

Twenty‐three Aphaenogaster species (Hymenoptera: Formicidae) occur in North America. While morphology and ecology define most species, the species limits of a group in the Eastern United States are unclear. In particular, the morphological and behavioural characters of A. carolinensis, A. picea and A. rudis overlap. These observations suggest that these three species are not monophyletic. We therefore tested the monophyly of Aphaenogaster in the context of molecular phylogenetic analyses. We used DNA data from five genes: CO1, CAD, EF1αF2, long‐wavelength rhodopsin and wingless, to reconstruct phylogenies for 44 Aphaenogaster and outgroup species. In the resulting trees, reconstructed using parsimony and Bayesian inference, species boundaries associated with well‐supported monophyletic clades of individuals in most of the 23 North American Aphaenogaster collected from multiple locations. However, some clades were unresolved, and both A. picea and A. rudis were not monophyletic. Although this may indicate that clades of multiple species represent fewer but morphologically varied species, given the short branch lengths, the lack of resolution may reflect the fact that these ants have recently radiated, and a lack of gene lineage sorting explains the non‐monophyly of species. Additional biological information concerning pre‐ and postmating barriers is needed before a complete revision of species boundaries for Aphaenogaster.     

Morphology agrees with molecular data: phylogenetic affinities of Euptychiina butterflies (Nymphalidae: Satyrinae)

Euptychiina is the most species‐rich subtribe of Neotropical Satyrinae, with over 450 known species in 47 genera (14 monotypic). Here, we use morphological characters to examine the phylogenetic relationships within Euptychiina. Taxonomic sampling included 105 species representing the majority of the genera, as well as five outgroups. A total of 103 characters were obtained: 45 from wing pattern, 48 from genitalia and 10 from wing venation. The data matrix was analysed using maximum parsimony under both equal and extended implied weights. Euptychiina was recovered as monophyletic with ten monophyletic genera, contrasting previous DNA sequence‐based phylogenies that did not recover the monophyly of the group. In agreement with sequence‐based hypotheses, however, three main clades were recognized: the ‘Megisto clade’ with six monophyletic and three polyphyletic genera, the ‘Taygetis clade’ with nine genera of which three were monophyletic, and the ‘Pareuptyhia clade’ with four monophyletic and two polyphyletic genera. This is the first morphology‐based phylogenetic hypothesis for Euptychiina and the results will be used to complement molecular data in a combined analysis and to provide critical synapomorphies for clades and genera in this taxonomically confused group.     

Phylogenetic analysis of the family Ariidae (Ostariophysi: Siluriformes), with a hypothesis on the monophyly and relationships of the genera

Ariid monophyly and intrafamilial relationships are investigated based on cladistic analysis of 230 morphological characters. Terminal taxa examined include whenever possible type‐species, or the most morphologically similar species to the type‐species of the nominal genera, and the largest possible number of species, including cleared and stained specimens, available in zoological collections. Previous hypotheses about monophyly of the Ariidae are strongly corroborated by new synapomorphies discovered in the present study. The subfamily Galeichthyinae and the remaining ariids are strongly supported by new morphological characters. The monotypic subfamily Bagreinae is recognized as the sister group to all nongaleichthyin ariids, supported by a large series of exclusive synapomorphies. A new concept of Ariinae is presented: the subfamily is found to be unequivocally monophyletic and includes all ariid genera, except Galeichthys and Bagre. New data supporting the monophyly of the genera included in the Ariinae are introduced and previous hypotheses of monophyly, species composition, morphological definition, and relationships are reviewed and discussed.     

Morphology,molecules, and the phylogenetics of cetaceans

Recent phylogenetic analyses of cetacean relationships based on DNA sequence data have challenged the traditional view that baleen whales (Mysticeti) and toothed whales (Odontoceti) are each monophyletic, arguing instead that baleen whales are the sister group of the odontocete family Physeteridae (sperm whales). We reexamined this issue in light of a morphological data set composed of 207 characters and molecular data sets of published 12S, 16S, and cytochrome b mitochondrial DNA sequences. We reach four primary conclusions: (1) Our morphological data set strongly supports the traditional view of odontocete monophyly; (2) the unrooted molecular and morphological trees are very similar, and most of the conflict results from alternative rooting positions; (3) the rooting position of the molecular tree is sensitive to choice of artiodactyls outgroup taxa and the treatment of two small but ambiguously aligned regions of the 12S and 16S sequences, whereas the morphological root is strongly supported; and (4) combined analyses of the morphological and molecular data provide a well-supported phylogenetic estimate consistent with that based on the morphological data alone (and the traditional view of toothed-whale monophyly) but with increased bootstrap support at nearly every node of the tree.     

Phylogeny of the Pantomorus–Naupactus complex based on morphological and molecular data (Coleoptera: Curculionidae)

The Pantomorus–Naupactus complex is a Neotropical group of broad‐nosed weevils (Coleoptera: Curculionidae) including several parthenogenetic species usually assigned to the genera Naupactus Dejean, Pantomorus Schoenherr, Asynonychus Crotch, Aramigus Horn, Eurymetopus Schoenherr and Graphognathus Buchanan. Sixteen species were studied to test hypotheses on the monophyly of these genera, and on the origin of the parthenogenetic lineages. A matrix of 30 morphological characters and 999 positions of the Cytochrome Oxidase I gene, was analyzed with separate partitions and simultaneously, under equal and implied weights, and with different transversion/transitions costs. The ILD test indicates that the incongruence between the molecular and morphological data is not significant. Under equal weights, the molecular data resulted in a single tree and morphology in 34 trees; under implied weights morphology gave a different tree, and under TV:TS ≥ 4:1 molecular and combined analyses resulted in the same optimal tree. According to the latter, Naupactus includes Graphognathus, and is thus paraphyletic and basal regarding remaining genera, Pantomorus is polyphyletic and includes Aramigus and Asynonychus, and Eurymetopus is monophyletic. The species in which apomictic parthenogenesis has been verified (Aramigus tessellatus, Asynonychus cervinus and Graphognathus lecuoloma), belong to different clades of the Pantomorus‐Naupactus complex, with basal sexual relatives. © The Willi Hennig Society 2005.     

Cladistic analysis of the Cirripedia Thoracica

We present a cladistic analysis of the Cirripedia Thoracica using morphological characters and the Acrothoracica and Ascothoracida as outgroups. The list of characters comprised 32 shell and soft body features. The operational taxonomic units (OTUs) comprised 26 well-studied fossil and extant taxa, principally genera, since uncertainty about monophyly exists for most higher ranking taxonomic units. Parsimony analyses using PAUP 3.1.1 and Hennig86 produced 189 trees of assured minimal length. We also examined character evolution in the consensus trees using MacClade and Clados. The monophyly of the Balanomorpha and the Verrucomorpha sensu stricto is confirmed, and all trees featured a sister group relationship between the ‘living fossil Neoverruca and me Brachylepadomorpha. In the consensus trees the sequential progression of ‘pedunculate‘sister groups up to a node containing Neolepas also conforms to current views, but certain well-established taxa based solely on plesiomorphies stand out as paraphyletic, such as Pedunculata (= Lepadomorpha); Eolepadinae, Scalpellomorpha and Chthamaloidea. The 189 trees differed principally in the position of shell-less pedunculates, Neoverruca, the scalpelloid Capitulum, and the interrelationships within the Balanomorpha, although the 50% majority rule consensus tree almost fully resolved the latter. A monophyletic Sessilia comprising both Verrucomorpha and Balanomorpha appeared among the shortest trees, but not in the consensus. A tree with a monophyletic Verrucomorpha including Neoverruca had a tree length two steps longer than the consensus trees. Deletion of all extinct OTUs produced a radically different tree, which highlights the importance of fossils in estimating cirripede phylogeny. Mapping of our character set onto a manually constructed cladogram reflecting die most recent scenario of cirripede evolution resulted in a tree length five steps longer than any of our shortest trees. Our analysis reveals that several key questions in cirripede phylogeny remain unsolved, notably the position of shell-less forms and the transition from ‘pedunculate‘to ‘sessile‘barnacles. The inclusion of more fossil species at this point in our understanding of cirripede phylogeny will only result in even greater levels of uncertainty. When constructing the character list we also identified numerous uncertainties in the homology of traits commonly used in discussing cirripede evolution. Our study highlights larval ultrastructure, detailed studies of early ontogeny, and molecular data as the most promising areas for future research.     

Phylogeny of haplo–diploid,fungus‐growing ambrosia beetles (Curculionidae: Scolytinae: Xyleborini) inferred from molecular and morphological data

Cognato, A. I., Hulcr, J., Dole, S. A. & Jordal, B. H. (2010). Phylogeny of haplo‐diploid, fungus‐growing ambrosia beetles (Curculionidae: Scolytinae: Xyleborini) inferred from molecular and morphological data. —Zoologica Scripta, 40, 174–186. The ambrosia beetle tribe Xyleborini currently contains 30 genera and approximately 1200 species which are distributed throughout worldwide forests with most diversity located in the tropics. They also represent the most invasive scolytines in North America. Despite economic concerns and biological curiosity with this group, a comprehensive understanding of generic boundaries and the evolutionary relationship among species is lacking. In this study, we include 155 xyleborine species representing 23 genera in parsimony and Bayesian analyses using 3925 nucleotides from mitochondrial (COI) and nuclear genomes (28S, ArgK, CAD, EF‐1α) and 39 morphological characters. The phylogenies resulting from the parsimony analyses, which treated gap positions either as missing or fifth character states, and the Bayesian analysis were generally similar. Clades with high support or posterior probabilities were found in all trees, while those with low support were not recovered by all analyses. Fourteen of the 23 genera were monophyletic although not all relationships among the genera were resolved. We show monophyly of several species groups associated with particular morphological and biological characters and suggest recognition of these groups as genera. Most interesting was the monophyly of South and Central American species representing several genera. This finding suggests recent and fast radiation of xyleborines in the New World accompanied by morphological and biological diversification.     

Molecular phylogenetics of the genus Ceratitis (Diptera: Tephritidae)

The Afrotropical fruit fly genus Ceratitis MacLeay is an economically important group that comprises over 89 species, subdivided into six subgenera. Cladistic analyses of morphological and host use characters have produced several phylogenetic hypotheses for the genus. Only monophyly of the subgenera Pardalaspis and Ceratitis (sensu stricto) and polyphyly of the subgenus Ceratalaspis are common to all of these phylogenies. In this study, the hypotheses developed from morphological and host use characters are tested using gene trees produced from DNA sequence data of two mitochondrial genes (cytochrome oxidase I and NADH-dehydrogenase subunit 6) and a nuclear gene (period). Comparison of gene trees indicates the following relationships: the subgenus Pardalaspis is monophyletic, subsection A of the subgenus Pterandrus is monophyletic, the subgenus Pterandrus may be either paraphyletic or polyphyletic, the subgenus Ceratalaspis is polyphyletic, and the subgenus Ceratitis s. s. might not be monophyletic. In addition, the genera Ceratitis and Trirhithrum do not form reciprocally monophyletic clades in the gene trees. Although the data statistically reject monophyly for Trirhithrum under the Shimodaira-Hasegawa test, they do not reject monophyly of Ceratitis.     

Phylogeny of West AfricanCaryedon(Coleoptera: Bruchidae): Congruence between Molecular and Morphological Data

Seed beetles belonging to the Old World genusCaryedonfeed in the seeds of various Caesalpinioideae, Mimosoideae, and Combretaceae. In an attempt to resolve broad phylogenetic relationships within the genus, we obtained 332 base pair sequences of mitochondrial 12S ribosomal DNA and morphological data for the 16 West AfricanCaryedonspecies. Morphological characters were analyzed under maximum parsimony and sequences were compared under maximum parsimony, maximum likelihood, and neighbor joining. Using a partition homogeneity test, we determined that morphological and molecular data sets were combinable. Combined data were analyzed under maximum parsimony. Morphological and molecular trees were congruent at the species group level and total evidence analyses yielded the same topologies as molecular data with each of the three outgroups used. Four main terminal clades are recognized, each corresponding with a group of species generally feeding on the same host plant family, subfamily, genus, or species. The monophyly of legume feedingCaryedonis supported by both data sets, and Combretaceae feeders split in two monophyletic assemblages.     

Phylogenetic study of heliconiine butterflies based on morphology and restriction analysis of ribosomal RNA genes

Relationships of ten heliconiine butterflies (genera Dryas and Heliconius , family Nymphalidae) were elucidated by phylogenetic analysis of characters based on ribosomal DNA restriction site variation and morphology. Agraulis vanillae , also a heliconiine, was used as the outgroup species. Although neither the morphological nor the molecular data unambiguously resolve relationships among the heliconiines, a combined analysis of both data sets results in a tree that is similar to traditional systematic arrangements and previous views of radiation in the group. Both pupal-mating and nonpupal-mating species group as clades in the combined analysis. However, the restriction site data alone do not support the monophyly of the pupal-mating clade, and the morphological data alone do not support the monophyly of the non-pupal-mating clade. Furthermore, relationships of H. melpomene, H. cydno and the silvaniform species depart from traditional arrangements based on morphology and reproductive compatibility experiments. All trees support the independent evolution of similar wing patterns of species previously suggested to be members of mimicry complexes. Several mimicry complexes appear to have a member in each of the two major monophyletic groups (pupal-mating and non-pupal-mating clades).     

Molecular phylogeny of the Oriental butterfly genus Arhopala (Lycaenidae,Theclinae) inferred from mitochondrial and nuclear genes

Abstract. We present a phylogeny for a selection of species of the butterfly genus Arhopala Boisduval, 1832 based on molecular characters. We sequenced 1778 bases of the mitochondrial genes Cytochrome Oxidase 1 and 2 including tRNA^Leu, and a 393‐bp fragment of the nuclear wingless gene for a total of 42 specimens of 33 species, representing all major species groups. Analyses of mtDNA and wingless genes show congruent phylogenetic signal. The phylogeny presented here confirms the monophyly of the centaurus, eumolphus, camdeo and epimuta groups and the amphimuta subgroup. It confirms close relationships between species within the agelastus group, that together with the amphimuta subgroup, centaurus and camdeo groups form a monophyletic group. However, incongruencies with previous taxonomic studies also occur; the amphimuta and silhetensis groups are not monophyletic, as is the genus Arhopala itself. One enigmatic species, A. kinabala, was evaluated further for topology and the support for basal placement of this species is due mainly to the wingless gene. However, in the Parsimony analysis, and subsequent Maximum Likelihood evaluations, certain nodes could not be resolved due to insufficient support. The mtDNA shows extreme AT bias with compositional heterogeneity at 3rd codon positions, which may result in saturation. By contrast, the wingless gene does not show compositional bias, suggesting that poor support is not due solely to saturation. The evaluation of morphological characters used in previous studies on Arhopala systematics on the molecular tree indicates that the macular pattern and the absence of tails at the hind wings show extensive homoplasy. A significant phylogenetic signal (as indicated by T‐PTP tests) is present in several of these morphological characters, which are nevertheless of limited use in phylogenetic studies due to their labile nature.     

Mitochondrial and nuclear markers support the monophyly of Dolichopodidae and suggest a rapid origin of the subfamilies (Diptera: Empidoidea)

Abstract. The Dolichopodidae is a species‐rich dipteran group with almost 7000 described species. The monophyly of the subfamilies and their relationships remain largely unknown because the polarities of key morphological characters are unclear and molecular data are available only for 9 of the 19 proposed subfamilies. Here we test whether molecular data from two nuclear (18S, 28S) and four mitochondrial (12S, 16S, Cytb, COI) genes can resolve the higher‐level relationships within the family. Our study is based on 76 Oriental species from 12 dolichopodid subfamilies and uses eight species of Empididae and Hybotidae as outgroups. Parsimony and likelihood analyses confirm the monophyly of the Dolichopodidae, as well as the monophyly of five of the ten subfamilies represented by more than two species [Sympycninae, Sciapodinae, Dolichopodinae, Hydrophorinae (excluding tribe Aphrosylini), Neurigoninae]. There is strong support for restoring the tribe Aphrosylini as a separate subfamily Aphrosylinae. The monophyly of Medeterinae, Peloropeodinae and Diaphorinae is dependent on which tree reconstruction technique is used, how indels are coded, and whether the fast‐evolving sites are excluded. Overall, we find that our sample of Oriental species is largely compatible with the subfamily concepts that were developed for the northern temperate fauna. However, our data provide little support for relationships between the subfamilies. Branch lengths, saturation, and distance plots suggest that this is probably the result of the rapid origin of dolichopodid subfamilies over a relatively short time. We find that genera that are difficult to place into subfamilies based on morphological characters are generally also difficult to place using molecular data. We predict that a dense, balanced taxon sample and protein‐encoding nuclear genes will be needed to resolve the higher‐level relationships in the Dolichopodidae.