Phylogenetic reassessment and biogeography of the Ectateus generic group (Coleoptera: Tenebrionidae: Platynotina)

Owing to the reinterpretation of its morphological synapomorphies, the taxonomic composition of the Ectateus generic group had been ambiguous. The present study scrutinized all existing taxonomic concepts of the group based on a cladistic analysis of the adult morphology of all of the Afrotropical platynotoid Platynotina genera. The phylogenetic relationships were reconstructed using parsimony and Bayesian inference. The results show that all previous taxonomic concepts of the Ectateus generic group concerned paraphyletic entities. The cladistic analysis revealed the following synapomorphies for the taxon: (1) presence of basal indentations of the pronotal disc, (2) ratio of prothorax width to its maximal height > 6.0, and (3) ratio of maximal height of the prothorax to total height < 0.3. Moreover, phylogenetic studies revealed the existence of the Upembarus generic group, a sister‐taxon group to the Ectateus generic group, within the Afrotropical platynotoid Platynotina. Autapomorphic and synapomorphic character mapping show that several taxonomic and nomenclatural changes are needed to consider the particular generic‐level entities traditionally assigned to Afrotropical platynotoid Platynotina as monophyletic lineages. The following taxonomic and nomenclatural adjustments are made in this paper: P teroselinus gen. nov. is erected to accommodate a single species that was previously assigned to Zidalus: Pteroselinus insularis comb. nov. Additionally, the following synonymies are proposed: Anchophthalmops (= Platykochius syn. nov. ), Angolositus (= Aberlencus syn. nov. , = Platymedvedevia syn. nov. ), Glyptopteryx (= Microselinus syn. nov. , = Quadrideres syn. nov. , = Synquadrideres syn. nov. ). In addition, Kochogaster is lowered in rank and is treated as one of the subgenera of Anchophthalmus. Moreover, Pseudoselinus is treated as a subgenus of Upembarus. An identification key to all Afrotropical platynotoid Platynotina genera and subgenera is presented. Zoogeographical analyses revealed the following dispersal barriers for the Ectateus generic group: (1) the Sahara (northern barrier); (2) the dry ecosystems of Botswana, Namibia, and South Africa (southern barrier); and (3) the Congolian rainforests (internal distributional gap). The ancestor of the taxon probably originated in East African ecoregions that predominantly contained wattletrees (acacias) and Commiphora Jacq. Moreover, past climate changes seem to have had a great impact on the observed generic distribution. © 2015 The Linnean Society of London     

Male and female genitalia in some Libyan species of Acrididae (Orthoptera: Acridoidea)

A comparative study of male and female genitalia was carried out in thirty‐seven Libyan species representing twenty genera of the family Acrididae. An attempt has been made to describe and illustrate the different structures, namely, epiphallus, aedeagus, subgenital plate, supra‐anal plate and cerci of the male, and spermatheca, ovipositor, subgenital plate, supra‐anal plate and cerci of the female, in Acridids, with an aim to discover their significance in order to make the identification of genera and species, together with other generic characters, more perfect and convenient. Distinct family characters are shield or bridge‐shaped condition of epiphallus; presence or absence of dorso‐lateral appendices, oval sclerites and lophi on epiphallus; divided, undivided or flexured condition of aedeagus; presence or absence of gonopore process on aedeagus; condition of apical and pre‐apical diverticula of spermatheca; presence or absence of glandular pouches of Cornstock and Kellog on female subgenital plate; and rudimentary or well developed condition of egg‐guide. Stable characters for separating the subfamilies are taken to be presence or absence of ancorae on epiphallus, long or short condition of aedeagal sclerites; elongate, slender or short and broad condition of ovipositor valves: presence or absence of Jannone's organs and setae on posterior margin of female subgenital plate; and shape of diverticula of spermatheca. Useful generic characters are shape of male subgenital plate, supra‐anal plate and cerci, broad or narrow condition of bridge, presence or absence of branch of bridge connecting lophi with bridge of epiphallus; mono‐, bi‐ or tri‐lobate condition of lophi of epiphallus, length and upcurved or downcurved condition of apical valve of aedeagus, shape of posterior margin of female subgenital plate, presence of setae on the whole posterior margin or confined to lateral margins only; and toothed, tuberculate or smooth condition of ovipositor valves, length of the lateral apodeme in relation to the dorsal valves. Specific characters are shape of egg‐guide of female subgenital plate, shape of ovipositor valves and apical tips, shape of male supra‐anal plate and cerci, size of anterior and posterior lobes of lophi of epiphallus, size and shape of ancorae, shape of apical valves of aedeagus; and size of apical and pre‐apical diverticula and presence of protuberance on pre‐apical diverticulum.     

Phylogeny of the platygastroid wasps (Hymenoptera) based on sequences from the 18S rRNA, 28S rRNA and cytochrome oxidase I genes: implications for the evolution of the ovipositor system and host relationships

The Platygastroidea are a diverse group of mostly small to tiny wasps, the common biology for which is endoparasitism of insect and spider eggs. No analytically‐based phylogeny exists for the superfamily, and the current suprageneric classification is flawed in part because of its reliance on homoplasious and pleisiomorphic morphological characters. To determine platygastroid relationships as a basis for investigating host and ovipositor evolution, phylogenies of > 70 in‐group species (representing 55 genera) were reconstructed by parsimony and Bayesian methods using three molecular markers; the mitochondrial cytochrome oxidase I and the nuclear genes 28S and 18S rRNA. The results strongly support the monophyly of the superfamily and one of the two families, Platygastridae, but the Scelionidae are most likely polyphyletic. However, within the Scelionidae, there is a well supported ‘main scelionid clade’ that contains the majority of genera assigned to the family. At the subfamilial level, both putative subfamilies of Platygastridae, the Platygastrinae, and Sceliotrachelinae, are likely to be polyphyletic. Within the Scelionidae, both the Teleasinae and Telenominae are monophyletic, but the Scelioninae is clearly not so. The current tribal classification for the Scelionidae is in need of major reassessment because no tribes, with the exception of the Scelionini s.s., were found to be monophyletic. Further illustrating the problems associated with the current classification is the nonmonophyly of a number of genera, namely Opisthacantha Caloteleia, Telenomus, Trimorus, Teleas and Idris. Analysis of ovipositor evolution in the superfamily revealed that the Ceratobaeus‐type ovipositor system is ancestral; however, this trait was lost prior to the evolution of the main scelionid clade, for which the Scelio‐type ovipositor system is ancestral and defines a mostly paraphyletic assemblage. Ancestral state analysis indicates that the Ceratobaeus‐type ovipositor was subsequently re‐evolved in the main scelionid clade, representing a possible contradiction of Dollo’s law. Previously, the tribal placement has been used to predict the host associations of genera for which host data were unavailable. However, the fact that most tribes are not monophyletic throws into doubt any such speculation based on the current classification. © 2007 The Linnean Society of London, Biological Journal of the Linnean Society, 2007, 91 , 653–669.     

A MORPHOLOGICAL STUDY AND TAXONOMIC REASSESSMENT OF THE GENERITYPE SPECIES IN THE GRACILARIACEAE1

We investigated the reproductive morphology of representative material corresponding to the type species of each of the described genera presently placed in synonymy under Gracilaria. From these observations and published studies of recognized genera, 10 species groups are identified in the Gracilariaceae based on spermatangial type and cystocarp development. Actual or potential generic names are given in brackets after each group: 1) abscissa group (Melanthalia), 2) flabellata group (Curdiea), 3) lemaneiformis group (Gracilariopsis), 4) chilensis group, 5) edulis group (Plocaria/Polycavernosa), 6) urvillei group (Hydropuntia), 7) crassissima group, 8) salicornia group (Corallopsis), 9) gracilis group, and 10) bursa‐pastoris group (Gracilaria). Tyleiophora was shown to belong to the bursa‐pastoris group. The type species of the parasitic genera Gracilariophila and Congracilaria are closely related to their host species. Species assemblages recognized here based on morphological evidence received moderate to strong bootstrap support in recently published molecular phylogenies. Further studies may show that some groups correspond to genera, whereas others do not merit generic status.     

Morphological characterization of infra‐generic lineages in Deparia (Athyriaceae: Polypodiales)

Deparia, including the previously recognized genera Lunathyrium, Dryoathyrium (=Parathyrium), Athyriopsis, Triblemma, and Dictyodroma, is a fern genus comprising about 70 species in Athyriaceae. In this study, we inferred a robust Deparia phylogeny based on a comprehensive taxon sampling (~81% of species) that captures the morphological diversity displayed in the genus. All Deparia species formed a highly supported monophyletic group. Within Deparia, seven major clades were identified, and most of them were characterized by inferring synapomorphies using 14 morphological characters including leaf architecture, petiole base, rhizome type, soral characters, spore perine, and leaf indument. These results provided the morphological basis for an infra‐generic taxonomic revision of Deparia.     

A chemically defined medium for rearing Epiphyas postvittana (Lepidoptera: Tortricidae)

Abstract

The endemic New Zealand genus Neolimnia is redescribed, and a key is given to the 14 species now included. Four new species are described - Neolimnia raiti, N. pepekeiti, N. ura, and N. repo. Two new synonymies are proposed; N. ocellata Tonnoir &; Malloch with N. castanea (Hutton); and N. dubiosa Tonnoir &; Malloch with N. irrorata Tonnoir &; Malloch. Lectotypes are designated for N. tranquilla (Hutton), N. castanea (Hutton), N. obscura (Hutton), and N. striata (Hutton). Male postabdomens of all species are illustrated. Character variation and generic, subgeneric, and species relationships are discussed briefly.     

Phylogeny and evolution of phytophagous ladybird beetles (Coleoptera: Coccinellidae: Epilachnini), with recognition of new genera

Phytophagous ladybird beetles of the tribe Epilachnini are a cosmopolitan, species‐rich group of significant economic importance as pests of agricultural crops. The tribe is well characterized morphologically and clearly monophyletic, but very little is known about its internal phylogenetic relationships and their genus‐level taxonomy. In order to infer the evolutionary history of Epilachnini, test its monophyly and provide a phylogeny‐based classification, we assembled a comprehensive dataset, consisting of four DNA markers (18S and 28S rRNA and 16S, COI mtDNA) and a matrix of 104 morphological characters for 153 species of Epilachnini representing all previously recognised genera, ～11% of the known species, and 14 outgroup taxa. Molecular, morphological and combined datasets were analysed using maximum likelihood, parsimony and Bayesian inference. Bayes factors and Approximately Unbiased tests (AU) were used to compare alternative phylogenetic hypotheses of unconstrained and backbone‐constrained analysis. Only 14 of the 25 included genera were recovered monophyletic, as originally defined. Afidentula Kapur, Afidenta Dieke, Afissula Kapur, Epilachna Chevrolat, Henosepilachna Li Toxotoma Weise and Mada Mulsant are shown to be poly‐ or paraphyletic; Chnootriba Chevrolat, Subafissa Bielawski, Lalokia Szawaryn & Tomaszewska and Papuaepilachna Szawaryn & Tomaszewska form monophyletic groups within larger clades of genus level. All of these genera are redefined here. The two largest genera of Epilachnini, Epilachna Chevrolat and Henosepilachna Li were represented by multiple monophyletic clades, which we described as new genera: Chazeauiana Tomaszewska & Szawaryn gen.n. ; Diekeana Tomaszewska & Szawaryn gen.n .; Fuerschia Tomaszewska & Szawaryn gen.n. and Ryszardia Tomaszewska & Szawaryn gen.n . The following new synonyms are proposed: Afissa Dieke (=Afissula Kapur); Henosepilachna Li in Li & Cook (=Subafissa Bielawski); Papuaepilachna Szawaryn & Tomaszewska (=Lalokia Szawaryn & Tomaszewska). This published work has been registered in ZooBank, http://zoobank.org/urn:lsid:zoobank.org:pub:440E7FA4‐C859‐47E0‐8335‐30D478CBA8FA .     

Phylogeny and taxonomy of the Prenolepis genus‐group of ants (Hymenoptera: Formicidae)

Abstract. We investigated the phylogeny and taxonomy of the Prenolepis genus‐group, a clade of ants we define within the subfamily Formicinae comprising the genera Euprenolepis, Nylanderia, gen. rev. , Paraparatrechina, gen. rev. & stat. nov. , Paratrechina, Prenolepis and Pseudolasius. We inferred a phylogeny of the Prenolepis genus‐group using DNA sequence data from five genes (CAD, EF1αF1, EF1αF2, wingless and COI) sampled from 50 taxa. Based on the results of this phylogeny the taxonomy of the Prenolepis genus‐group was re‐examined. Paratrechina (broad sense) species segregated into three distinct, robust clades. Paratrechina longicornis represents a distinct lineage, a result consistent with morphological evidence; because this is the type species for the genus, Paratrechina is redefined as a monotypic genus. Two formerly synonymized subgenera, Nylanderia and Paraparatrechina, are raised to generic status in order to provide names for the other two clades. The majority of taxa formerly placed in Paratrechina, 133 species and subspecies, are transferred to Nylanderia, and 28 species and subspecies are transferred to Paraparatrechina. In addition, two species are transferred from Pseudolasius to Paraparatrechina and one species of Pseudolasius is transferred to Nylanderia. A morphological diagnosis for the worker caste of all six genera is provided, with a discussion of the morphological characters used to define each genus. Two genera, Prenolepis and Pseudolasius, were not recovered as monophyletic by the molecular data, and the implications of this result are discussed. A worker‐based key to the genera of the Prenolepis genus‐group is provided.     

Phylogeny of the tribe Empoascini (Hemiptera: Cicadellidae: Typhlocybinae) based on morphological characteristics,with reclassification of the Empoasca generic group

A morphology‐based phylogenetic analysis of the tribe Empoascini (Hemiptera: Cicadellidae: Typhlocybinae) is presented for 58 of 83 formerly recognized genera based on 99 morphological characters of adults. The results support excluding the New World Beamerana generic group from Empoascini. The remaining genera of Empoascini were recovered as a monophyletic sister group of Dikraneurini. Previously recognized tribes Jorumini and Helionini are derived from within Empoascini and are considered synonyms of the latter tribe. Three previously recognized informal generic groups, the Empoasca group, Alebroides group and Usharia group were paraphyletic but the Ficiana group was recovered as monophyletic based on five synapomorphies. Genera previously placed in the Alebroides group represent at least six independent lineages, indicating that the hind wing character separating this group from the Empoasca group (CuA and MP veins free) is highly homoplasious. Empoasca (sensu lato) is also paraphyletic. Thus, twelve previously recognized subgenera of Empoasca are elevated to genus status and five species groups of Empoasca from the New World are recognized as separate new genera. Sikkimasca Dworakowska, 1993 is treated as synonym of Marolda Dworakowska, 1977 based on the phylogeny. Biogeographic analysis suggests that Empoascini most likely first evolved in the Oriental region and spread to other biogeographic realms more recently by multiple independent invasions.     

Reassessment of Paleotachina Townsend and Electrotachina Townsend and their removal from?the?Tachinidae (Diptera)

The monotypic genera Paleotachina Townsend, 1921 and Electrotachina Townsend, 1938 were originally described as fossils in amber but were later discovered to be inclusions in copal. Both taxa were originally assigned to the Tachinidae (Diptera) and this placement has continued to the present day. The holotypes of the two type species, P. smithii Townsend and E. smithii Townsend, were examined and the following taxonomic and nomenclatural changes are proposed: Paleotachina is transferred to the Muscidae and placed in synonymy with Aethiopomyia Malloch, 1921, syn. n.; P. smithii Townsend, type species of Paleotachina, is synonymized with Aethiopomyia gigas (Stein, 1906), syn. n.; Electrotachina is transferred to the Sarcophagidae and placed in synonymy with Dolichotachina Villeneuve, 1913, syn. n.; E. smithii Townsend, type species of Electrotachina, is recognized as a valid species of Dolichotachina comb. n. Images of the holotypes of P. smithii and E. smithii are provided and features that have helped place these copal inclusions in their new combinations are discussed.     

Ericandersonia sagamia, a new genus and species of deep-water eelpouts (Perciformes: Zoarcidae) from Japan

A new genus and species of deep-water zoarcid fish, Ericandersonia sagamia, is described on the basis of four specimens collected from Sagami Bay, Japan, at depths of 880–930 m. This species is placed in the subfamily Gymnelinae and is distinguished from all genera of gymnelines by the following characters: pseudosubmental crest present; frontals partially fused dorsally; parietals meeting in midline; pelvic fins absent; ventral ramus of posttemporal weak; postorbital head pores 5; pectoral-fin rays 14. The phylogenetic analysis (based on 42 morphological characters) indicates that its position is outside the highly modified (= advanced) genera Seleniolycus, Melanostigma, Puzanovia, Nalbantichthys, Opaeophacus, and Andriashevia.     

Phylogenetic relationships in the Marcetia alliance (Melastomeae,Melastomataceae) and implications for generic circumscription

The Marcetia alliance of Melastomataceae is an exclusively Neotropical group that includes at least 12 genera of mostly herbs and subshrubs, occurring in the cerrado of central Brazil and savannas of the Amazon region and Guayana highlands. This study aimed to test the monophyly of genera in the Marcetia alliance, evaluate their phylogenetic relationships and generic boundaries, and investigate morphological characters as potential synapomorphies for delimiting clades or genera. We used nuclear (ITS, ETS) and plastid (accD‐psaI, atpH‐atpF, trnS‐trnG) DNA sequences of 107 terminals in 12 genera from the alliance. Aciotis, Fritzschia, Marcetia and Siphanthera were shown to be monophyletic and supported by molecular and morphological characters. Other genera with variable morphology and wider distributions, such as Acisanthera, Comolia, Ernestia and Macairea, were recovered as paraphyletic or polyphyletic. Most morphological characters analysed were found to be homoplastic, but when combined they are potentially useful for the diagnosis of genera and infrageneric groups. This study represents a major step in understanding internal relationships and provides the basis for a revision of the generic classification in the Marcetia alliance.     

The ovipositing female of Ooencyrtus telenomicida relies on physiological mechanisms to mediate intrinsic competition with Trissolcus basalis

Ongoing studies by our group showed that the outcome of the intrinsic competition between two solitary egg parasitoids, Trissolcus basalis (Wollaston) (Hymenoptera: Scelionidae) and Ooencyrtus telenomicida (Vassiliev) (Hymenoptera: Encyrtidae), is dominated by O. telenomicida. In this article we investigated the role played by the ovipositing O. telenomicida female in the suppression of a T. basalis competitor. Laboratory experiments were conducted by allowing an O. telenomicida female to puncture the eggs of Nezara viridula (L.) (Heteroptera: Pentatomidae) with her ovipositor (= no oviposition) or to parasitize them. The results show that O. telenomicida relies on some physiological mechanisms to mediate its interspecific intrinsic competition with T. basalis. In fact, the emergence of T. basalis was strongly reduced in host eggs that were parasitized either before or after being punctured by O. telenomicida at fixed time intervals (5, 15, 30, or 45 h). The low percentage of emergence of T. basalis (ranging from approximately 4–20%) was a consequence of the delay and growth rate reduction of larval development. Furthermore, the percentage of eclosion of N. viridula nymphs was negatively affected by the O. telenomicida female’s punctures (96% from healthy host eggs, 4% from punctured host eggs). Host eggs punctured or oviposited in by O. telenomicida showed alterations in the ooplasm including some melanized‐like areas near the hole made with the ovipositor; such alterations indicate that the adult parasitoid releases substances that affect the host eggs survival. These results suggest that the O. telenomicida female influences both the physiological interspecific parasitoid‐parasitoid interaction, as well as the host‐parasitoid interaction, providing, for the first time in egg parasitoids, evidence that physiological suppression of some competitive egg parasitoids is mediated by the ovipositing female.     

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.     

The suprageneric groups ofthe Pimplinae (Hymenoptera: Ichneumonidae): a cladistic re-evaluationand evolutionary biological study

The phylogenetic relationships of the suprageneric groupsof the ichneumonid subfamily Pimplinae (Hymenoptera) are re‐assessedusing 166 morphological and biological characters for 162 species,representing all of the available described genera and subgenera.The cladistic analysis was repeated using abstracted genera, re‐codedfrom the ­initial set of species, as terminal taxa. Thetopology of the resulting cladograms was similar. In the first (primary) analysisseveral genera (including Neotheronia, Itoplectis, Dolichomitus, Dreisbachia, Polysphincta, Oxyrrhexis and Zonopimpla)were not retrieved as monophyletic groups; however, all except thelast were found to be monophyletic in the second analysis. Theseresults suggest that using abstracted taxa may force a ‘falsemonophyly’ on the preselected groups. Thus we reject theuse of such abstractions, preferring instead to use exemplar speciesthat together show much of the variation that occurs within a hypothesizedgenus. Within the Pimplinae three major groupings were recognized,the Delomeristini (including the Perithoini syn. nov.) , thePimplini and the Ephialtini. Within the Pimplini, two generic groupswere recovered, the Xanthopimpla and Pimpla genus‐groups,but a third postulated group, the Theronia genus‐group, wasfound to be paraphyletic. Within the Ephialtini five groups wererecognized, the Pseudopimpla, Alophosternum, Camptotypus, Ephialtes and Sericopimpla genus‐groups.The spider parasitizing complex of genera (the Polysphincta genus‐complex)was found to nest within the Sericopimpla genus‐group confirmingthe placement of Polysphinctini as a synonym of Ephialtini. Problemswith the status of some existing genera are highlighted, but formalnomenclatural changes are not proposed. The ancestral Pimplinaeare hypothesized to have been solitary ectoparasitic idiobiontson weakly concealed immature Hymenoptera. The major radiations withinthe Pimplinae are shown as: (1) a progressive exploitation of cocooned,then weakly cocooned, lepidopterous pupae in the Pimplini leadingto idiobiont endoparasitism; (2) increasing specialization to attackhosts deeply concealed in wood in the Ephialtes genus‐group,and (3) specialization on a variety of cocooned hosts, includingspider egg sacs, leading to koinobiont ectoparasitism of spiders.A brief synopsis of the distribution of the group is given, and somebiogeographical inferences drawn. The group is presumed to haveoriginated and radiated on Laurasia; no evidence for trans‐Antarcticrelationships can be found. © 2002 The LinneanSociety of London, Zoological Journal of the Linnean Society,2002, 136 , 421?485