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.     

叶甲亚科口器比较形态学研究(鞘翅目,叶甲科)

叶甲亚科成虫为植食性昆虫,是典型的咀嚼式口器.对叶甲亚科8属8种叶甲(Agrosteomela chinensis (Weise),Ambrostoma fortunei(Baly),Asiparopsis convexa (Weise),Chrysolina aurichalcea(Mannerheim),Chrysomela populi Linnaeus,Gonioctena tredecimmaculata(Jacoby),Parambrostoma mahesa(Hope),Paropsides soriculata(Swartz))的口器进行了比较形态学研究,并对其演化进行了讨论.     

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.     

A description and classification of some final—instar larvae of the Mesochorinae (Hymenoptera, Ichneumonidae)

The characters of the final–instar larvae of the Mesochorinae are reviewed on knowledge of species belonging to four genera. Relationships indicated by larval characters, within the Mesochorinae, and between the Mesochorinae and other groups of rchneumonidae, are outlined. Evidence is given that the larval characters of Cidaphus indicate this genus to be the least specialized of the Mesochorinae.     

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 systematic position of Antirrhea and Caerois, with comments on the classification of the Nymphalidae (Lepidoptera)

Based on a Wagner tree analysis of ninety-two characters (eighty-five larval, one egg, six adult), the nymphalid butterfly genera Antirrhea and Caerois are demonstrated to be the closest relatives of the genus Morpho . Accordingly, Antirrhea and Caerois are formally transferred from the Satyrinae to the Morphinae. Without these two genera, the Morphinae ( sensu Ehrlich) is at best a paraphyletic group. During the study, fourteen nymphalid genera were treated as potential outgroups. The analysis suggests that many currently accepted higher taxa within the Nymphalidae are probably untenable: the Satyrinae and Nymphalinae ( sensu Ehrlich) are both probably polyphyletic; the Biinae ( sensu Miller) must be abandoned, being polyphyletic; the Charaxidae ( sensu Rydon), although probably monophyletic, appear to form a group subordinate to part of the 'Satyrinae'; and Apatura does not cluster with the 'Nymphalinae', but appears to form the sister-group of the 'Satyrinae' (less Antirrhea and Caerois ) plus the Charaxinae. Re-analyses of reduced data sets, in which potentially homoplasious larval head-horn and adult wing venational characters were eliminated, leaves these conclusions essentially unaltered. The authors suggest that a solution to the seemingly intractable problem posed by nymphalid higher classification can be sought by the application of cladistic analysis to a large data set gathered from all developmental stages, with special emphasis on detailed comparative larval morphology.     

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.     

The genera of Chrysomelinae (Coleoptera: Chrysomelidae) in Costa Rica

Keys in Spanish and English are given for the genera of Chrysomelinae known from Costa Rica. For each genus, a list of species compiled from collections in the University of Costa Rica, the National Biodiversity Institute, and the entomological literature is presented. The genus Planagetes Chevrolat 1843 is recorded for the first time from Central America, and the genus Leptinotarsa St?l 1858 is synonymized with Stilodes Chevrolat 1843.     

叶甲亚科三种蛹的形态学研究

对叶甲亚科 3属 3种蛹的形态特征进行了详细描述 ,分别为小猿叶甲PhaedonbrassicaeBaly ,菜无缘叶甲ColaphellusbowringiiBaly和柳圆叶甲Plagioderaversicolor(Laicharting) ,其中菜无缘叶甲为首次描述。同时对蛹的形态特征在叶甲亚科系统演化方面的作用进行了一定的探讨     

Homoplasy and the delineation of holophyletic genera in some insect groups

Abstract. Some groups of insects, such as certain Ichneumonidae and fungus-feeding Phlaeothripidae, appear to have undergone frequent reversal or parallelism (homoplasy) of characters during evolution so that extant species present almost every imaginable permutation and combination of characters. Recognition of holophyletic genera in such groups is difficult. Large monothetically defined genera are often not holophyletic, whilst small genera need to be defined by a large and invariable character-suite. Any classification produced by adopting small genera will not possess one of the valuable attributes of the classical Linnaean system, its predictive ability; a large proportion of new species will require new genera. For groups exhibiting a high degree of homoplasy it is suggested that a polythetic classification be erected. Polythetic genera can be holophyletic groups and are not merely phenetic assemblages. The probability of correctly assigning a species (either a new one or when making an identification) is shown to be higher for a polythetic classification. A simple key device, the polyclave, is given to enable practical separation of two polythetic taxa. It is suggested that homoplasy, and its associated problems in classification, may be associated with a particular type of biology in which population size is not limited by direct competition.     

Chemical defences in leaf beetles and their larvae: The ecological, evolutionary and taxonomic significance

The chemical defences of the Chrysomelinae are reviewed. Defensive glandular secretions have evolved independently in larvae and adults, and faster than the morphology of the glands. Both characters are used in a phylogenetic study of the Chrysomelini, disclosing suprageneric affinities. First, a close relationship between the Chrysomelina and Phratora is proposed. Secondly, Leptinotarsa and Gonioctena are probably more closely related to the Chrysolinina than to the Chrysomelina and Phratora. The qualitative and quantitative composition of larval and adult secretions sometimes differs between geographically isolated populations of the same species, which are therefore probably genetically distinct. The host plant shows an influence which explains discrepancies between the current classification and the chemical nature of the secretions. Some beetles sequester secondary plant metabolites or use them as precursors for their own defences. The original biosynthetic pathway has been lost in these species. Other factors which could affect the chemical diversity of the secretions are discussed: random events such as neutral mutations and genetic drift, conditioning or adaptation of predators to specific defensive mixtures, and the diversity of potential predators. The latter could help to explain the divergence in the chemical nature of the larval and adult secretions. The volatile irritants produced by the larvae are well suited to repel small arthropods, such as ants, whereas poisons associated with aposematic coloration in the adults could be directed against small vertebrates, such as birds.     

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.     

Canegrubs and cladistics: what story do adult, larval and ecological characters tell?

Abstract  Melolonthine scarabs, canegrubs, are the most important pests of Australian sugarcane. The 19 species are sometimes viewed as one pest; however, these species are quite diverse, are placed in four genera and a number of species-groupings in one of those genera, show different life histories and adult behaviours, and some have distinct soil-type preferences. Three questions concerning relationships between the canegrubs were addressed here. First, the relationships suggested by a taxonomic revision of the whole tribe based on adult characters were generally confirmed through phylogenetic analysis. Second, neither the larval characters nor the ecological characters show similar distinct phylogenetic patterns; more characters are needed to resolve the relationships. Third, the analysis of a combination of adult, larval and ecological characters, the 'total evidence' approach, does give a more robust phylogenetic arrangement than that derived from only adult characters. This analysis strengthens the understanding of the relationships among canegrubs and provides the basis for improved management systems for these pests.     

Cladistic analysis of the Sepsidae (Cyclorrhapha: Diptera) based on a comparative scanning electron microscopic study of larvae

The result of a phylogenetic analysis of the Sepsidae based on larval characters is presented. It is shown that cyclorrhaphan larvae can be as rich a source of characters as Nematocera immatures when investigated using an SEM. The cladistic analysis comprised fifty-two species in sixteen genera of the Sepsidae and five outgroup species and used fifty-seven morphological characters. It found seven parsimonious trees which only differed with respect to the arrangement of some species within the genus Themira. The basal dichotomies of the phylogenetic trees are particularly well supported, indicating the conservative nature of larval characters. Orygma is confirmed as the sister group of all the remaining sepsids, the Sepsinae. There is good larval evidence that Ortalischema is the sister group of all remaining Sepsinae and that the Toxopodinae constitute an early radiation within the Sepsidae. According to larval data, some genera are paraphyletic ( Themira, Palaeosepsis ), but adult characters appear to contradict these findings. Among the traditionally recognized higher taxa within the Sepsidae, Hennig's Themira species-group and Steysbal's Sepsini have to be rejected as polyphyletic. However, Hennig's Sepsis species-group is confirmed as monophyletic and will probably constitute one major element of a future phylogenetic system of the Sepsidae. States of the strongly modified fore-legs of some adult sepsid males are mapped onto the phylogenetic tree, largely confirming Šulc's ideas about the evolution of these features. The origin and evolution of male sternites with brushes and a gland on the tibiae of the males ('osmeterium') are discussed. Whereas adult characters point to a sister-group relationship between the Sepsidae and the Ropalomeridae, larval characters appear to indicate a sister-group relationship between the Coelopidae and the Sepsidae. The evidence for both hypotheses is critically evaluated.     

The Phylogeny of the Lophopidae and the Impact of Sexual Selection and Coevolutionary Sexual Conflict

A cladistic analysis of Lophopidae was performed, using 73 observed morphological characters and 41 taxa. This analysis involved 36 genera belonging to the Lophopidae family and 5 outgroups. For a better understanding of the selected characters some illustrations are provided. The most parsimonious cladograms obtained show that the Lophopidae cannot be considered as a monophyletic lineage unless two genera are withdrawn from this family: Hesticus Walker, 1862, and Silvanana Metcalf, 1947. The systematic position of these two genera remains uncertain. They cannot yet be included in another family of Fulgoromorpha. A cladistic analysis of each of the 19 remaining Fulgoromorphan families must be performed first. A new family could be created for these two genera, but not before we are sure that these two genera are in no way members of an existing family. The outgroup problem is discussed. The monophyletic lineage represented by the Lophopidae can be divided into four natural groups: Carriona⁺, Makota⁺, Sarebasa⁺, and Bisma⁺. When a cladistic analysis is completed using a data matrix without characters linked to females, the cladogram obtained presents a disrupted basal resolution. Female characters appear to bring a phylogenetic signal important basally in the evolution of the Lophopidae but also apically, directly between the relationships of some genera. A similar analysis, using a matrix without characters linked to males, provides a phylogeny disrupted between the groups that form the Lophopidae and in the basal resolution in these groups. The respective impacts of the genitalic characters are discussed in relation to sexual selection conflict.     

Reassessment of the enigmatic Lepidopteran family Lypusidae (Lepidoptera: Tineoidea; Gelechioidea)

Abstract. The genus Lypusa Zeller, 1852 has been assigned to Tineoidea (Lepidoptera) with varying family positions. The systematic affinities of this genus, currently placed in its own family Lypusidae, were studied using extensive data derived from larval, pupal and adult morphology and certain behavioural traits. In total, 193 characters were considered. On the basis of the results of a parsimony analysis, Lypusa is transferred to the superfamily Gelechioidea, in a monophyletic, exclusively Palaearctic assemblage with the genera Amphisbatis and Pseudatemelia. Several phylogenetically relevant characters support this position, including similarly constructed larval cases, densely porose larval head, and a modification of the pupal abdominal segment 8. The composition of the family Amphisbatidae is discussed and it is proposed that it be delimited as comprising solely the Palaearctic genera Pseudatemelia, Amphisbatis and Lypusa. A detailed diagnosis of the genus Lypusa is given.