A phylogeny of aphid parasitoids (Hymenoptera: Braconidae: Aphidiinae) inferred from mitochondrial NADH 1 dehydrogenase gene sequence

Wasps of the braconid subfamily Aphidiinae are solitary endoparasitoids of aphids. Several aspects of their biology have been the focus of intuitive evolutionary hypotheses which could be tested with a robust phylogeny. Phylogenetic hypotheses have been proposed previously for aphidiines based on morphology, embryology, and DNA sequences. However, many of them are based on a limited number of characters and/or taxa and lack congruence. In addition, many of the inferred phylogenies have not been based upon cladistic analysis. Therefore, a phylogenetic study of Aphidiinae was undertaken, utilizing 465 bp of DNA sequence of the mitochondrial NADH1 dehydrogenase gene. DNA sequences were obtained from 40 taxa, including 14 genera and three outgroups. It is suggested that in agreement with most of the previously proposed phylogenies, the aphidiines, each of the three recognized tribes (Praini, Ephedrini, Aphidiini), and most genera are monophyletic. In contrast to previously proposed phylogenies, the clade of Praon + Dyscritulus (=Praini), rather than Ephedrini, is basal among the aphidiines.     

Phylogenetic relationships among genera of Aphidiinae (Hymenoptera: Braconidae) based on DNA sequence of the mitochondrial 16S rRNA gene

Phylogenetic relationships among forty‐nine taxa representing twenty‐four genera of Aphidiinae (Hymenoptera: Braconidae) were investigated using DNA sequence of a portion of the mitochondrial 16S rRNA gene and parsimony analysis. Seven species in six other subfamilies of Braconidae were used as outgroup. The results suggested that members of Aphidiinae are monophyletic. The basal lineage of Aphidiinae was Aclitus in weighted and unweighted parsimony analyses and Praini was basal relative to Ephedrini. With the exception of Pauesia and Aphidius, all genera were monophyletic. The results support generic status for Euaphidius, but not for Lysaphidus. Diaeretus leucopterus was internal to a clade composed of three Pauesia species, suggesting that the latter genus may be paraphyletic. A combined analysis that included DNA sequence of 16S rRNA, NADH1 dehydrogenase and 28S rRNA resulted in more robust cladograms with topologies similar to those inferred from the 16S rRNA gene sequence alone. The results are compared to previously proposed phylogenies of Aphidiinae based on morphological and molecular characters.     

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.     

Grazers, shredders and filtering carnivores--the evolution of feeding ecology in Drusinae (Trichoptera: Limnephilidae): insights from a molecular phylogeny

We examined the phylogenetic relationships between species and genera within the caddisfly subfamily Drusinae (Trichoptera: Limnephilidae) using sequence data from two mitochondrial loci (cytochrome oxidase 1, large subunit rRNA) and one nuclear gene (wingless). Sequence data were analysed for 28 species from five genera from the subfamily. We analysed individual and combined data sets using a Bayesian Markov Chain Monte Carlo and a maximum parsimony approach and compared the performance of each partition for resolving phylogenetic relationships at this level. In terms of resolution and phylogenetic utility wingless outperformed the two mitochondrial gene partitions. Using both Shimodaira-Hasegawa and expected likelihood weights tests we tested several hypotheses of relationships previously inferred based on adult morphological characters. The data did not support the generic concept, or many previously proposed species groupings, based on adult morphology. In contrast, the molecular data correlated with the morphology and feeding ecology of larvae. Using Bayesian ancestral character state reconstructions we inferred the evolution of feeding ecology and relevant larval morphological characters. Our analyses showed that within the subfamily Drusinae two derived feeding types evolved. One of these--grazing epilithic algae--is otherwise unusual in the Limnephilidae and may have promoted the high degree of diversity in the Drusinae.     

Evolutionary relationships of wing venation and wing size and shape in Aphidiinae (Hymenoptera: Braconidae)

We explored evolutionary changes in wing venation and wing size and shape in Aphidiinae, one of the well-known groups of parasitic wasps from the family Braconidae. Forewings of 53 species from 12 genera were examined, for which a molecular phylogeny was constructed on the basis of the mitochondrial barcoding gene COI. By covering all types of wing venation within the subfamily Aphidiinae and by using landmark-based geometric morphometrics and phylogenetic comparative methods, we tested whether evolutionary changes in wing shape correlate to the changes in wing venation and if both changes relate to wing size. The relationship between wing morphology and host specificity has been also investigated. We found that six types of wing venation, with different degree of vein reduction, could be recognized. Wing venation type is largely genus specific, except in the case of maximal reduction of wing venation which could be found across examined Aphidiinae taxa. The reconstruction of evolutionary changes in wing venation indicates that evolutionary changes in wing shape are related to the changes in wing size, indicating that miniaturization play a role in evolution of wing morphology while host specialization does not affect the wing shape within the subfamily Aphidiinae.     

Morphology and chaetotaxy of the first instar larvae of six species of the Satyrus (s.l.) series (Lepidoptera: Nymphalidae)

Abstract. The morphology and chaetotaxy of the first instar larvae of six species belonging to the genera Hipparchia, Kanetisa and Chazara are described. Specific characters are stated, drawn mainly from size, setal length and morphology, and the shape of the suranal plate. Several characters, other than chaetotaxy, that are of potential use in nymphalid systematics are discussed. The larval chaetotaxy is briefly compared with that of both heliconiine and danaine first instar larvae.     

Characterization and mapping of sensilla on the head appendages of noterid larvae (Coleoptera: Noteridae), and development of a preliminary biometric method for taxa delimitation

Larvae of the burrowing water beetle family Noteridae are distributed worldwide and are often abundant in a broad range of aquatic habitats, playing an important role in structuring freshwater communities, yet they have remained among the most poorly studied groups of aquatic beetles. Studies on sensillar equipment of aquatic insect larvae are largely lacking, despite their potential use in phylogeny and biometric identification methods. In this article, the external morphology and distribution of sensilla on the head appendages of first instar larvae of selected genera of Noteridae were examined using scanning electron microscopy. Seven main types were distinguished based on their morphological structure: basiconica (3 subtypes), campaniformia (2 subtypes), chaetica (7 subtypes), coeloconica (6 subtypes), coniform complex (2 subtypes), placodea, and styloconica (3 subtypes). The apex of the labial palpus was found to be the most variable and informative region in regard to the number, relative position, and topology of sensilla. Fingerprint models were, therefore, generated for this region in each of the studied genera, allowing their identification.     

Larval morphology and biology of oxycorynine weevils and the higher phylogeny of Belidae (Coleoptera, Curculionoidea)

Phylogenetic relationships among members of the family Belidae (Curculionoidea) were reconstructed through cladistic analysis using 58 characters and 17 terminals. The characters were from larval morphology (30), adult morphology (25) and biology regarding larval host-plants and feeding habits (three). They were scored for exemplar taxa in 17 genera, representing different belid subfamilies and tribes, plus two outgroup taxa in Megalopodidae and Nemonychidae. The sampled genera included all those for which larval and adult information is available, and two known only from adults. New information on the larvae and biology of two oxycorynines is provided. These are the Chilean Oxycraspedus cribricollis , whose larvae live in decayed female strobili of the gymnosperm Araucaria araucana , and Hydnorobius hydnorae from Argentina, whose larvae, described and illustrated in the present paper, develop inside the flower and fruit bodies of Prosopanche americana (Hydnoraceae), a root-parasitic angiosperm. The relationships proposed by the single optimal cladogram resulting from simultaneous analysis of all taxa and characters are recovered by one of three optimal cladograms based on the larval data set alone. The cladogram justifies a revised classification of Belidae in two sister subfamilies: Belinae (with tribes Pachyurini, Agnesiotidini and Belini) and Oxycoryninae (with tribes Oxycorynini and Aglycyderini). It summarizes larval and adult synapomorphies defining the family Belidae, subfamilies and tribes. Based on the phylogenetic tree, the evolution of biological traits is traced. Larval development in vegetative organs of conifers is ancestral in Belidae. A shift to reproductive structures characterizes the Oxycorynini, a habit which was conserved while several shifts to distantly related host-plant groups occurred.     

An evaluation of the systematic value of skull morphology in the Trimeresurus radiation (Serpentes: Viperidae: Crotalinae) of Asian pitvipers

Trimeresurus (in its widest sense) is a very diverse and widespread radiation of Asian pitvipers, which has been subject to numerous taxonomic revisions, some of which have been based on characteristics of the skull. In order to evaluate the taxonomic utility of such characters, we conducted a comparison of the skulls of 57 specimens representing nine genera, and two currently unassigned species that are very closely related to each other. A canonical variate analysis reveals three distinct phenetic groups: the Protobothrops group, a group containing Ovophis monticola , ' Ovophis ' okinavensis , and ' Trimeresurus ' gracilis , and finally a group comprising the remaining species and characterized by considerable overlap between most genera with the exception of the monotypic Peltopelor and Himalayophis . Agreement between phenetic similarity based on skull characteristics and phylogenetic relationships based on molecular evidence varies between different groups: the morphological similarity of the skull of Protobothrops sieversorum to the other Protobothrops species is congruent with their recent synonymization while the phenetic similarity among the species within the second group does not reflect current molecular phylogenetic relationships and indicates that convergent or parallel evolution may be responsible for at least some of the phenetic similarity detected among skulls of the Asian pit vipers examined. A test of phylogenetic independence, however, indicates that there is still a significant phylogenetic signal that can be recovered from several skull characteristics. Thus, we conclude that skull morphology can contribute to an overall understanding of pitviper taxonomy, but that it would be unwise to rely on skull characteristics alone.     

Immature stages of Hipparchia Fabricius and the systematics of the 'Satyrus series>> (Lepidoptera: Nymphalidae: Satyrinae)

Abstract. Characters of potential use from the eggs and larvae of four species of Hipparchia (subgenera Hipparchia, Parahipparchia, Neohipparchia and Pseudotergumia ) are described. These characters, together with previously published information on larval, pupal and adult morphology, are used to construct a seventy-one character matrix for Hipparchia and four other genera of the 'Satyrus series' (Miller, 1968): Arethusana, Kanetisa, Chazara and Satyrus . The cladistic relationships among these genera and the relative contribution of adult and immature data sets are discussed. A list of possible synapomorphies of the immature stages of the 'Satyrus series' is proposed.     

Cladistic and phenetic estimates of relationship among genera of eupomphine blister beetles (Coleoptera: Meloidae)

Abstract. The Eupomphina includes seven distinctive genera of Meloidae from the arid and semiarid zones of North America. Cladistic and phenetic estimates of relationship among genera are analysed and discussed. Both adult and larval characters are used, and the relative contribution of each to the classification of the subtribe is examined. Generic keys to adults and first instar larvae are included.     

Phylogeny and classification of the Scopulini moths (Lepidoptera: Geometridae, Sterrhinae)

The phylogenetic relationships of the genera in the geometrid tribe Scopulini (Lepidoptera: Sterrhinae) were examined using 141 characters of adult morphology and ecology. The study material included 92 species, representing all previously recognized genera and covering the morphological variation and full geographical range of the tribe. The cladistic analysis resulted in 20 equally parsimonious trees and a strict consensus cladogram based on these was well resolved. A majority of the recovered synapomorphic characters have been used previously in the taxonomy of the tribe. However, many novel characters were found in the sclerotized structures of the thorax. Many previously recognized genera were found to be nonmonophyletic and based on the present revised, synapomorphy-based classification, the number of recovered genera is reduced considerably. Twenty new generic synonyms and 90 new or revived species combinations are proposed. Seven genera are considered valid, with the large genus Scopula Schrank including over 85% of all species in the tribe. The taxonomic history of the tribe is reviewed and the problems of earlier classifications are discussed. A key to the genera is presented, although an informal diagnosis is preferred. All recognized genera are illustrated and a revised world checklist of the Scopulini is presented.  © 2005 The Linnean Society of London, Zoological Journal of the Linnean Society , 2005, 143 , 473−530.     

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.     

Description of a Polynoncus Burmeister larva, with implications for phylogeny of the Trogidae (Coleoptera: Scarabaeoidea)

Abstract. The larva of Polynoncus seymourensis Mutchler is described from the Galapagos Archipelago, Ecuador. It is the first larva recorded of any species in the genus and has a suite of characters not found in the other trogid genera. Phylogenetic analysis of characters of larvae of all the genera supports the current phylogenetic system which was based on adults.     

The evolution of hymenopteran wings: the importance of size

The allometric relationships between body size and several aspects of wing morphology in the insect order Hymenoptera were investigated using multivariate morphometric techniques. The study focused primarily on wing allometry in five monophyletic genera of bees ( Perdila, Halictus, Ceratina, Trigona and Apis ), but the patterns of size-related evolutionary change found within each of these genera are also found to exist in numerous other hymenopteran taxa. Increased body size in hymenopteran lineages is correlated with the following changes in wing morphology: (1) decreased relative stigma area, (2) distal extension of wing vein elements, (3) increased aspect ratio and (4) proximal shift in the centroid of wing area. The reverse is true for decreased size. The widespread allometric trends most likely result from adaptive change in wing morphology due to size-related changes in the physical properties impinging on the organism–principally the quality and magnitude of drag. The fact that similar wing morphologies among distantly related species can result from similarity in body size has important implications for the study of hymenopteran phylogeny, especially at lower taxonomic levels and when a high proportion of wing characters are employed.     

国剑角蝗科部分属种雌性下生殖板的形态及演化（直翅目：蝗总科）

从20世纪60年代至今的一些研究已基本肯定了蝗虫雌性下生殖板的分类价值。本文选择了中国剑角蝗科6属10种蝗虫对其雌性下生殖板进行了形态学和演化分析。结果表明雌性下生殖板的形态在研究的6属内都是稳定的,属间差异明显。这6个属的系统发育关系为：（卡蝗属（（佛蝗属,戛蝗属）,（菊蝗属（蝗属,剑角蝗属））））,这一结果和形态分类是一致的,支持长腹蝗亚科从剑角蝗科分开。文中还分析了剑角蝗科雌性下生殖板各性状的演化方向。     

Cladistics and the comparative morphology of linyphiid spiders and their relatives (Araneae, Araneoidea, Linyphiidae)

This paper provides the first quantitative cladistic analysis of linyphiid morphology. Classical and novel homology hypotheses for a variety of character systems (male and female genitalia, somatic morphology, spinneret silk spigot morphology, etc.) are critically examined and studied within a phylogenetic context. Critical characters have been illustrated. A sample of linyphiid taxa (nine genera in four subfamilies), five species of Pimoa (Pimoidae), and two other araneoid families (Tetragnathidae and Araneidae, represented by Tetragnatha and Zygiella , respectively) were used to study the implications of the phylogeny of Pimoidae for the systematics of linyphiids. The phylogenetic relationships of these 16 exemplar taxa, as coded for the 47 characters studied, were analysed using numerical cladistic methods. In the preferred cladogram Pimoidae and Linyphiidae are sister groups, Stemonyphantinae are sister group to the remaining linyphiids, and Mynogleninae are sister group to the clade composed of Erigoninae plus Linyphiinae. These results agree with the relationships recently proposed by Wunderlich, except by finding erigonines as the sister group to linyphiines rather than to mynoglenines.