Phenetic and cladistic relationships among tenebrionid beetles (Coleoptera)

Abstract. The higher classification of Tenebrionidae is analysed using numerical phenetic, numerical cladistic and traditional Hennigian methods. In all, eighty characters are examined for about 335 taxa; definitive analyses are made on combinations of eighteen to seventy characters for thirty-three OTUs. At lower levels of relationship (genera and closely related tribes) phenetic and cladistic classifications are shown to be congruent, but at higher levels (tribes and subfamilies) there is marked discordance with phenetic results being more stable. A consensus classification is more similar to the Hennigian cladogram than is any single computer generated cladogram. Two main tribal groups – the Lagrioid and Tenebrionoid groups – are suggested which differ in defensive glands, female anatomy, wing and mouthpart morphology, larval characters and other features. The Tenebrionoid group consists of three main subdivisions – the tenebrionine, coelometopine and diaperine lineages. Changes in classificatory position are recommended for eighty-seven genera and tribes (listed in Appendix E) and implied for numerous others.     

A cladistic analysis of the nomadine bees (Hymenoptera: Apoidea)

Abstract. This study compares the results of Rozen's cladistic analysis of the larvae of fifteen genera of cleptoparasitic bees in the subfamily Nomadinae with an independent data set of adult characters for the same genera. Adult characters exhibited considerably higher levels of homoplasy and poorer resolution of cladistic relationships, with multiple equally parsimonious cladograms. However, comparison of a Nelson consensus tree based on adult characters with the cladogram based on larval characters reveals three components consistently supported in both analyses (the tribes Epeolini and Ammobatini, and Neopasites + Neolarra) , one component supported only by adult characters (Isepeolus + Protepeolus) , and one terminal component supported only by larval characters (Nomada + Ammobatini), as well as several more inclusive groupings based on larval characters that are difficult to compare with the adult consensus tree because it shows so much less resolution. When adult and larval characters are combined in a single data matrix, the resulting cladogram closely resembles the cladogram based on larval characters alone, although levels of homoplasy are considerably higher than in the larval analysis.
A preliminary analysis of adult characters for thirty-four genera in the Nomadinae also exhibited high levels of homoplasy and very large numbers of equally parsimonious cladograms. Nevertheless, certain consistent monophyletic groupings, most notably the Epeolini and Ammobatini, were also supported in this analysis. The one currently recognized tribe whose monophyly has received no support from any analysis is the Nomadini.
The relevance of these phylogenetic hypotheses to our understanding of host associations and variable features of egg morphology and oviposition behaviour in nomadine bees is briefly discussed.     

The phylogeny and systematics of European predacious Syrphidae (Diptera) based on larval and puparial stages

The external larval morphology of predacious Syrphidae is examined and compared with that of other groups of syrphids. Predacious larvae form a distinctive, monophyletic group. Relationships among 29 of the 34 European genera and subgenera are analysed using cladistic methods based on larval and puparial characters. Interpretation of the cladogram suggests that, within Syrphinae, a system of five tribes can be recognized and that a major trend has been a shift from ground layer to arboreal larval habitats, with an increasing degree of obligate dependence on soft-bodied Homoptera as prey. Generic diagnoses with biological and taxonomic notes are given and a key to genera based on larvae and puparia is provided.     

Monophyly of Euaesthetinae (Coleoptera: Staphylinidae): phylogenetic evidence from adults and larvae,review of austral genera,and new larval descriptions

Abstract We develop a morphological dataset for the rove beetle subfamily Euaesthetinae comprising 167 morphological characters (135 adult and 32 larval) scored from 30 terminal taxa including 25 ingroup terminals (from subfamilies Euaesthetinae and Steninae) and five outgroups. Four maximum parsimony analyses using different sets of terminals and character sets were run to test the monophyly of (1) Euaesthetinae, (2) Steninae, (3) Euaesthetinae + Steninae, (4) euaesthetine tribes Austroesthetini, Alzadaesthetini, Euaesthetini, Fenderiini and Stenaesthetini, and (5) the ten currently known austral endemic genera together. Analyses of adult and larval character sets separately and in combination recovered the monophyly of Euaesthetinae, Steninae, and both subfamilies together, with strong support. Analysis of 13 ingroup terminals for which complete data were available suggests that monophyly of Euaesthetinae is supported by 19 synapomorphies (13 adult, six larval), of Steninae by 23 synapomorphies (14 adult, nine larval), and of both subfamilies together by 24 synapomorphies (21 adult, three larval). Within Euaesthetinae, only the tribe Stenaesthetini was recovered as monophyletic based on adult characters, and in no analyses were the ten austral endemic genera recovered as a monophyletic group. Phylogenetic relationships among euaesthetine genera were weakly supported, although analyses including adult characters supported monophyly of Octavius and Protopristus separately, and of Octavius + Protopristus, Austroesthetus + Chilioesthetus and Edaphus + Euaesthetus. Steninae may include a third genus comprising two undescribed species probably possessing a ‘stick–capture’ method of prey capture, similar to that in Stenus. These two species formed a strongly supported clade recovered as the sister group of Stenus based on adult characters. Diagnoses and a key to adults are provided for the 15 euaesthetine genera currently known from the austral region (Australia, New Zealand, South Africa and southern South America). Euaesthetine larvae previously were known only for Euaesthetus, and we describe the larvae of nine more genera and provide the first larval identification key for genera of Euaesthetinae.     

Phylogeny and classification of Corylophidae (Coleoptera: Cucujoidea) with descriptions of new genera and larvae

Abstract Phylogenetic relationships within the family Corylophidae were investigated. Twenty ingroup taxa and six outgroups were included in a cladistic analysis, based on 48 characters derived from adult and larval morphology. Phylogenetic analysis confirms that Corylophidae are monophyletic within the superfamily Cucujoidea and may be subdivided into two subfamilies: the Australian Periptycinae and the cosmopolitan Corylophinae containing 10 tribes: Foadiini trib.n. , Cleidostethini, Aenigmaticini, Parmulini, Sericoderini, Peltinodini, Orthoperini, Corylophini, Teplinini and Rypobiini. All currently recognized family‐group taxa are thoroughly diagnosed, and keys to their identification based on adults and larvae are provided. Two new genera and three species are described: Weirus gen.n ., containing only W. tozer sp . n . (Australia: Queensland), and Stanus gen.n. , with the two species S. bowesteadi sp.n . (New Zealand) and S. tasmanicus sp.n. (Tasmania). The larvae of Pakalukodes bimaculatus?lipiński et al. from Queensland and of Stanus bowesteadi sp.n. from New Zealand are described and illustrated for the first time.     

Phylogenetic analysis of the Pinnotheridae (Crustacea, Brachyura) based on larval morphology, with emphasis on the Dissodactylus species complex

Comparative larval morphology was used to elucidate phylogenetic relationships within the Pinnotheridae and the Dissodactylus species complex. Within the family, seven zoeal and six megalopal characters suggested two equally parsimonious phylogenetic hypotheses for pinnotherid larvae, both with Ostracotheres tridacnae representing the sister group for the Dissodactylus complex. Results indicated that the genus Pinnotheres is a polyphyletic taxon, and that the traditional subfamilial arrangement comprises paraphyletic taxa within the subfamilies Pinnotherinae and Pinnothereliinae. Certain evidence has suggested that Fabia and Juxtafubia should be excluded from the Pinnotherinae and placed into the Pinnothereliinae. Larval and adult morphology suggested that Pinnotheres politus should be included within Tumidotheres. The phylogenetic analysis within the Dissodactylus complex involved one zoeal and 16 megalopal characters. Results suggested a single phylogenetic hypothesis based on larval morphology. Combining adult morphology with larval evidence resulted in two equally parsimonious phylogenetic hypotheses, one of which agreed with a previously suggested hypothesis based only on adult characters.     

Higher Level Phylogeny of Curculionidae (Coleoptera: Curculionoidea) based mainly on Larval Characters, with Special Reference to Broad-Nosed Weevils

A cladistic analysis of Curculionidae was performed using 49 characters (41 from larvae, three from pupae, and five from adults). Illustrations of characters of immatures are provided. The analysis involved 19 terminal units and a hypothetical ancestor determined by the outgroup comparison method used to root the tree. One most parsimonious cladogram was obtained based on the complete data set and the following phylogenetic hypothesis is proposed: Ithycerinae, Microcerinae, and Brachycrinae sensu stricto are broad-nosed weevils placed sequentially at the base of the cladogram. The remaining weevil subfamilies form two major natural groups: one constituted by the sister taxa Rhynchophorinae—Platypodinae; the other with Erirhininae at the base, as sister taxon of the "Curculionidae sensu stricto " which show an unresolved trichotomy involving Curculioninae, Cossoninae—Scolytinae, and the clade including the Entiminae and allied subfamilies. This latter clade of broad-nosed weevils has Thecesterninae at the base; the next branch is Amycterinae, the sister taxon of the clade comprising two groups: one constituted by Aterpinae, Rhytirrhininae, and Gonipterinae; the other is Entiminae whose units form two main clades: one constituted by the sister tribes Pachyrhynchini—Ectemnorhinini, and the other by Alophini, Sitonini, and Entimini. When the analysis was done using only immature characters, results congruent with those based on the complete data set were obtained, except for the placement of Erirhininae. According to the results the hypothesis of monophyly of broad-nosed weevils is not accepted; the Entiminae are justified as monophyletic and their natural classification into tribes is proposed and the phylogenetic position and relationships of higher taxa of Curculionidae are discussed. This paper shows the importance of immature characters in recognition of natural groups and relationships in Curculionidae.     

A revised subfamily classification of Tenebrionidae (Coleoptera)

Existing classifications of Tenebrionidae are reviewed briefly. The inclusion of the families Alleculidae, Lagriidae, and Nilionidae in Tenebrionidae is confirmed. The splitting off from this complex of a family, Tentyriidae, by Doyen is discussed and rejected. Various taxa which had been included in Tenebrionidae are excluded, amongst which Syrphetodes, Brouniphylax, Exohadrus, Arthopus, Cotulades, Docalis, and Latometus (=Elascus) have not previously been formally excluded. A new family, Archeocrypticidae, is established and defined briefly for Archeocrypticus, Sivacrypticus, and Enneboeus.

Data from matrices based on adult and larval characters comparing Tenebrionidae with most other families of Tenebrionoidea (=Heteromera) are presented for derived characters in common, and for overall similarity. The families most closely related to Tenebrionidae according to these data are Zopheridae, Chalcodryidae, Merycidae, Archeocrypticidae, Synchroidae, Colydiidae, and Monommatidae; none is very close to Tenebrionidae, which has had a long independent history.

Characters of the subfamilies recognised are tabulated, and interpreted in a phylo‐genetic dendrogram. Phylogeny is discussed in relation to adaptive changes in the biology of the various subfamilies, which are Zolodininae new subfamily, Pimeliinae new sense (including Tentyriinae), Toxicinae new sense, Phrenapatinae new sense (including Archeoglenini new tribe), Diaperinae new sense, Gnathidiinae, Tenebrioninae new sense, Alleculinae, Nilioninae, Lagriinae new sense, Cossyphinae, and Cossyphodinae new status.

Biology, economic importance, copulation, orientation of the aedeagus, and distribution are discussed briefly.

Definitions of the family and subfamilies and a key to subfamilies are given, and keys to tribes are included for the smaller subfamilies. The previously unknown larvae of the genera Zolodinus, Menimus, Archeoglenes, Lepispilus, and Nyctoporis are described in detail. Pupae of Zolodinus and Nyctoporis are described. Keys to larvae include many other genera which were hitherto unknown or poorly known.     

Phylogeny of the bees of the family Apidae based on larval characters with focus on the origin of cleptoparasitism (Hymenoptera: Apiformes)

Abstract.  Fifty-four genera of the bee family Apidae comprising almost all tribes were analysed based on 77 traditional and one new character of the mature larvae. Nine, especially cleptoparasitic species, were newly added. Analyses were performed by maximum parsimony and Bayesian inference. Trees inferred from the analysis of the complete dataset were rooted by taxa from the families Melittidae and Megachilidae. Unrooted trees inferred from the analysis of the partial dataset (excluding outgroup taxa) are also presented to preclude possible negative effects of the outgroup on the topology of the ingroup. Only the subfamily Nomadinae was statistically well supported. The monophyly of the subfamilies Xylocopinae and Apinae was not topologically recovered. The monophyly of the tribe Tetrapediini was supported, and this tribe was found to be related to xylocopine taxa. At the very least, larval morphology suggests that Tetrapedia is not a member of the subfamily Apinae. Our analyses support the monophyly of the Eucerine line (Emphorini, Eucerini, Exomalopsini, Tapinotaspidini) and of the Apine line (Anthophorini, Apini, Bombini, Centridini, Euglossini, Meliponini). All analyses support the monophyly of totally cleptoparasitic tribes of the subfamily Apinae. We named this group the Melectine line (Ericrocidini, Isepeolini, Melectini, Osirini, Protepeolini, Rhathymini). In previous studies all these cleptoparasitic tribes were considered independent evolutionary lineages. Our results suggest that their similarities with hosts in morphology and pattern are probably the result of convergence and host–parasite co-evolution than phylogenetic affinity. According to the present analysis, the cleptoparasitism has evolved independently only six times within the family Apidae.     

A CLADISTIC ANALYSIS OF THE FAMILY TRISTIRIDAE (ORTHOPTERA, ACRIDOIDEA)

Abstract— A cladistic analysis of the endemic South American family Tristiridae was performed using 29 characters from external morphology and the genitalia. Polarity of characters was based on the outgroup comparison method. One most parsimonious cladogram of 54 steps was obtained, from which a classification of the family Tristiridae was constructed. The analysis of phylogenetic relationships showed that the different kinds of characters define taxa at different levels in the cladogram. Those mostly from the phallic complex define suprageneric taxa while those from external morphology characterize genera. It is hypothesized that in Tristiridae differentiation of the phallic complex preceded differentiation of external morphology and that characters from the phallic complex arc less conservative than those from the external morphology.     

Phylogeny and classification of Leptophlebiidae (Ephemeroptera) with an emphasis on Neotropical fauna

Mayflies from the family Leptophlebiidae are cosmopolitan and highly diverse morphologically; they are also the largest family in numbers of genera and the second in number of species in the order Ephemeroptera. In spite of their broad diversity and the efforts employed to understand the evolution of this group, the internal classification of Leptophlebiidae remains controversial at all levels. More recently, important changes have been incorporated into the systematics of the family, increasing the number of subfamilies (from two to six) and recognizing several tribes. We present a phylogeny of the family based on 153 taxa (53 genera) and two molecular markers, representing 1655 bp, and verify the taxonomic status of the subfamilies, tribes and complexes. Based on these results, the number of subfamilies has been increased from six to eight and one new tribes and two new subtribes have been added. In addition, new ranks are proposed and the concept of Atalophlebiinae revised, including genera with distributions in the Australasian and Neotropical regions.     

Phylogenetic studies of Meloidae (Coleoptera), with emphasis on the evolution of phoresy

One of the most recent classifications of Meloidae is based on the assumption that phoretic first‐instar larvae evolved twice in the family, once in Meloinae and again in Nemognathinae. Within Meloinae, this scheme places all presumed phoretic taxa in Meloini regardless of other characteristics. This paper challenges this classification with a cladistic analysis of all meloid genera whose first‐instar larvae were available for study. It concludes that phoresy evolved several times in Meloinae alone and that Meloini, when defined to include all phoretic genera, is polyphyletic. Cladistic support also is presented for four subfamilies of Meloidae and for several of the traditional tribes recognized in recent classifications.     

First phylogeny of predatory flower flies (Diptera, Syrphidae, Syrphinae) using mitochondrial COI and nuclear 28S rRNA genes: conflict and congruence with the current tribal classification

The family Syrphidae (Diptera) is traditionally divided into three subfamilies. The aim of this study was to address the monophyly of the tribes within the subfamily Syrphinae (virtually all with predaceous habits), as well as the phylogenetic placement of particular genera using molecular characters. Sequence data from the mitochondrial protein-coding gene cytochrome c oxidase subunit I ( COI ) and the nuclear 28S ribosomal RNA gene of 98 Syrphinae taxa were analyzed using optimization alignment to explore phylogenetic relationships among included taxa. Volucella pellucens was used as outgroup, and representatives of the tribe Pipizini (Eristalinae), with similar larval feeding mode, were also included. Congruence of our results with current tribal classification of Syrphinae is discussed. Our results include the tribe Toxomerini resolved as monophyletic but placed in a clade with genera Ocyptamus and Eosalpingogaster . Some genera traditionally placed into Syrphini were resolved outside of this tribe, as the sister groups to other tribes or genera. The tribe Bacchini was resolved into several different clades. We recovered Paragini as a monophyletic group, and sister group of the genus Allobaccha . The present results highlight the need of a reclassification of Syrphinae.
© The Willi Hennig Society 2008.     

Larval morphology enhances phylogenetic reconstruction in Cetoniidae (Coleoptera: Scarabaeoidea) and allows the interpretation of the evolution of larval feeding habits

Abstract.  The Cetoniidae, the showiest of scarabs, comprises some 3900 species in 515 genera, distributed worldwide except for subpolar areas and some offshore New Zealand islands. Parsimony analysis of 76 larval and adult characters and 42 terminal taxa supports the monophyly of Cetoniidae (sensu Krikken, 1984 ), but not of the traditionally considered subfamilies Cetoniinae and Trichiinae (sensu Krikken, 1984 ). In the study taxon, larval characters are shown to be more informative than those of adults for deeper phylogeny. The evolution of some larval characters (head and legs) in relation to feeding habits is discussed on the basis of phylogenetic analysis. The results show an evolutionary shift from wood associations to a higher ecological plasticity that allows the larvae to feed on wide sources of organic matter (including compost, dung, gopher burrows, packrat middens, ant debris piles, etc.).     

Phylogeny and tribal classification of Sterrhinae with emphasis on delimiting Scopulini (Lepidoptera: Geometridae)

Abstract. The phylogenetic relationships of tribes of the geometrid subfamily Sterrhinae (Lepidoptera) were studied, with special emphasis on finding delimiting characters for the tribe Scopulini. Two cladistic analyses were conducted for fifty‐nine species representing all previously recognized Sterrhinae tribes and covering the geographical range of the subfamily. In the first analysis, twelve putative synapomorphies of Scopulini, taken from the literature, were coded for actual specimens in order to test their ability to support the monophyly of the group. The resulting strict consensus cladogram was totally unresolved. In the second analysis, the twelve characters were combined with additional information from the morphology and ecology of adults and immature stages. Analysis of these ninety‐six characters resulted in a well‐resolved cladogram. The tribes were found to be monophyletic, except Cosymbiini and Rhodostrophiini. There are two main lineages within Sterrhinae: Cosymbiini + Rhodometrini + Timandrini and Rhodostrophiini + Cyllopodini + Sterrhini + Scopulini. Aletini and Problepsini lay within the concept of Scopulini. The association of the included Larentiinae taxa with the Cosymbiini + Rhodometrini + Timandrini lineage questions the monophyly of Sterrhinae. A majority of the recovered synapomorphic characters had been recognized previously, but several new phylogenetically informative characters were found, especially from the thorax. No unique characters diagnosing the tribe Scopulini were found, but many homoplastic synapomorphic features were found which diagnose parts of it. All recognized Sterrhinae genera are assigned tentatively to tribes and problematic cases are discussed.     

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.     

Comparative morphology,biology and phylogeny of terminal‐instar larvae of the European species of Toryminae (Hym., Chalcidoidea,Torymidae) parasitoids of gall wasps (Hym. Cynipidae)

We present a phylogenetic and taxonomic study of the morphology and biology of the terminal‐instar larval stage of 19 species representing all the genera of Torymidae parasitoids of gall wasps in Europe, with the single exception of Megastigmus. The genera studied include Adontomerus Nikol'skaya, Idiomacromerus Crawford, Chalcimerus Steffan & Andriescu, Glyphomerus Förster, Pseudotorymus Masi and Torymus Dalman. We primarily used chaetotaxy and some head structures. The terminal‐instar larvae of all studied species are thoroughly described for the first time and illustrated with SEM images. We provide diagnostic characters for the family and the genera studied, and keys to genera and species for the identification of torymid larvae associated with cynipid galls. The majority of the torymid larvae studied are solitary monophagous parasitoids. Finally, to assess the potential use of larval characters in systematic studies of the family, a phylogenetic analysis of the studied taxa based on 42 larval morphological characters is proposed and compared with the current taxonomy of Torymidae. Our results suggest that body chaetotaxy, and characters of the head and mouthparts could be used for genera and species discrimination. © 2008 The Linnean Society of London, Zoological Journal of the Linnean Society, 2008, 154 , 676–721.     

Phylogeny of split‐footed lacewings (Neuroptera,Nymphidae), with descriptions of new Cretaceous fossil species from China

A phylogeny of the lacewing family Nymphidae based on morphology and DNA sequences is presented including representatives of all living genera and selected fossil genera. Widely distributed Jurassic and Cretaceous genera gave rise to recent taxa now restricted to Australasia. Two previously defined clades (i.e. Nymphinae and Myiodactylinae) were recovered and reflect the diverging adult and larval morphology of members of these two subfamilies. From Chinese Cretaceous deposits, a new genus (Spilonymphes gen. nov.) is described with one new species, as well as new species described in the genera Baissoleon Makarkin and Sialium Westwood.     

Phylogeny of Myrmeleontiformia based on larval morphology (Neuropterida: Neuroptera)

The suborder Myrmeleontiformia is a derived lineage of lacewings (Insecta: Neuroptera) including the families Psychopsidae, Nemopteridae, Nymphidae, Ascalaphidae and Myrmeleontidae. In particular, Myrmeleontidae (antlions) are the most diverse neuropteran family, representing a conspicuous component of the insect fauna of xeric environments. We present the first detailed quantitative phylogenetic analysis of Myrmeleontiformia, based on 107 larval morphological and behavioural characters for 36 genera whose larvae are known (including at least one representative of all the subfamilies of the suborder). Four related families were used as outgroups to polarize character states. Phylogenetic analyses were conducted using both parsimony and Bayesian methods. The reconstructions resulting from our analyses corroborate the monophyly of Myrmeleontiformia. Within this clade, Psychopsidae are recovered as the sister family to all the remaining taxa. Nemopteridae (including both subfamilies Nemopterinae and Crocinae) are recovered as monophyletic and sister to the clade comprising Nymphidae + (Myrmeleontidae + Ascalaphidae). Nymphidae consist of two well‐supported clades corresponding to the subfamilies Nymphinae and Myiodactylinae. Our results suggest that Ascalaphidae may not be monophyletic, as they collapse into an unresolved polytomy under the Bayesian analysis. In addition, the recovered phylogenetic relationships diverge from the traditional classification scheme for ascalaphids. Myrmeleontidae are reconstructed as monophyletic, with the subfamilies Stilbopteryginae, Palparinae and Myrmeleontinae. We retrieved a strongly supported clade comprising taxa with a fossorial habit of the preimaginal instars, which represents a major antlion radiation, also including the monophyletic pit‐trap building species.