Sperm transfer by spermatophore in Diptera: new results from the Diopsidae

Sperm transfer by spermatophore is reported as a common and possibly primitive feature of the Diopsidae (Diptera, Schizophora). Spermatophores occur in four out of five investigated genera, in Cyrtodiopsis, Diasemopsis, Teleopsis , and in Sphyracephala , which is positioned close to the base of the family. Within the genus Diopsis the spermatophore is subject to reduction. Spermatophore size and copulation duration are positively correlated across the species. This is only partially explained by the time requirement of sperm transfer. Parsimony in time and material seems to be favoured in species with high mating frequency at special aggregation sites, whereas high investment in time and material occurs in species that do not mate preferentially at certain times or locations. The utilization of spermatophores has strong implications on the possibilities of sperm competition and cryptic female choice and may account for the occurrence of a postcopulatory passive phase or prolonged mating in some species. The increasing number of records of spermatophores in Diptera is reviewed.     

Baltic amber fossils reveal early evolution of sexual dimorphism in stalk-eyed flies (Diptera: Diopsidae)

A cladistic analysis of 16 species of extant and amber fossil stalk-eyed flies of the family Diopsidae places the fossil †Prosphyracephala succini (Loew) as the sister group of all other Diopsinae, the subfamily in which eye stalks occur. The study is based on a scoring including five old and 23 new finds of †P. succini from Baltic amber, and for the first time allows a morphometric analysis of eye span and various body size parameters in this species. The data indicate that sexual dimorphism of the eye stalks already existed in †Prosphyracephala, suggesting that this feature evolved early in the Diopsinae. Contrary to recent views that the ancestral condition of diopsine eye stalks was monomorphic, the new results suggest that sexual selection was involved in the evolution of eye stalks from the very beginning of the lineage.     

Big ‘antlers’ are favoured: female choice in stalk-eyed flies (Diptera,Insecta), field collected harems and laboratory experiments

Summary Cyrtodiopsis whitei (Diopsidae) from Malaysia is one of the stalk-eyed flies which show a marked sexual dimorphism ofeye spans. The particularly long eye stalks of male flies are to be regarded as an epigamic feature. Field observations show the males' eye spans to be correlated with the possession of a harem and with that harem's size. The question whether such a harem is acquired by male competition only, or whether female choice plays a part, is solved in female preference tests, offering dummy males of different eye spans for choice. Optical clues are found to be the relevant stimuli.     

The enigmatic monotypic crab plover Dromas ardeola is closely related to pratincoles and coursers (Aves, Charadriiformes, Glareolidae)

The phylogenetic placement of the monotypic crab plover Dromasardeola (Aves, Charadriiformes) remains controversial. Phylogenetic analysis of anatomical and behavioral traits using phenetic and cladistic methods of tree inference have resulted in conflicting tree topologies, suggesting a close association of Dromas to members of different suborders and lineages within Charadriiformes. Here, we revisited the issue by applying Bayesian and parsimony methods of tree inference to 2,012 anatomical and 5,183 molecular characters to a set of 22 shorebird genera (including Turnix). Our results suggest that Bayesian analysis of anatomical characters does not resolve the phylogenetic relationship of shorebirds with strong statistical support. In contrast, Bayesian and parsimony tree inference from molecular data provided much stronger support for the phylogenetic relationships within shorebirds, and support a sister relationship of Dromas to Glareolidae (pratincoles and coursers), in agreement with previously published DNA-DNA hybridization studies.     

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.     

槭树科植物广义形态学性状分支分析

通过45个广义的形态学性状对槭树科(Aceraceae)尤其是槭属(Acer L.)的主要类群做了分支分析,结果显示：1)槭属内由于各类群分布着较多的同塑性状状态,而难以为属下组间关系的解决提供更多有价值的信息;通过对具体的性状状态分布分析显示,对于象槭属这样在形态上分化较大的类群,由于多数分类性状在不同类群间经历了平行和逆转演化,因而在较低分类阶元水平很难选择合适的性状来通过分支分析构建其系统发育;2)鸡爪槭组(section Palmata)作为整个槭属的基部类群,虽然支持率较低,但与其它类群相比在槭属内维持了较多的原始性状;3)金钱槭属(Dipteronia Oliv.)的两个种作为单系得到了100％的靴带支持,且和槭属作为姐妹群也得到了较好的支持。     

Relationships within Siphini (Hemiptera,Aphidoidea: Chaitophorinae) in light of molecular and morphological research

The aim of this paper was to further explore the phylogeny of Siphini by analysing molecular data (two mitochondrial genes and two nuclear markers), together with morphological (29) and ecological (two) characters, for comprehensive analyses concerning the evolution of Siphini, relationships within the tribe, and between Siphini and other Chaitophorinae. Nine Siphini species, which represent all the genera of this tribe, as well as 12 out‐group species (mainly Chaitophorini representatives of the genera Chaitophorus and Periphyllus), were used in the analyses. Molecular phylogenetic trees were reconstructed by the Bayesian inference (BI) phylogenetic analysis and maximum parsimony (MP) criterion. The cladistic analysis was performed using nona . The monophyly of Siphini was confirmed. Species belonging to subgenera Sipha and Rungsia were clustered together, and this clade was a sister with reference to a clade including the genera Atheroides and Chaetosiphella. Monophyly of Atheroides was confirmed by the molecular data; however, in cladistic analysis Atheroides seemed to be paraphyletic because Atheroides hirtellus was placed as sister to Atheroides serrulatus and Chaetosiphella. The monotypic genera Caricosipha and Laingia formed separate lineages, and Laingia was sister to all other Siphini. Chaitophorini was not retrieved by the molecular and combined data: Periphyllus was sister to a clade containing Chaitophorus and Siphini.     

康蚖群的系统发生和生物地理学的研究(原尾目:始蚖科)(英文)

应用HENNIG86系统发生分析程序,选择出成虫的有代表性的16个共同衍征,分析了与康蚖属相关各属的各个种类,得出了该类群的相关系统树。同时对特征的演化以及其扩散途径等也进行了讨论。文中还应用Page 1993的COMPONENT程序进行了生物地理学的分析;得到种类—地区支序图。其中康蚖属表现为典型的印度—太平洋分布型,而其姐妹群新康蚖属则兼有东洋区和全北区的分布型。至于分布地区的相互关系和在这些地区的分衍替代事件的发生等问题也作了初步的假想。     

Phylogeny of the World Anthophorini (Hymenoptera: Apoidea: Apidae)

Abstract.  The phylogenetic relationships of members of the long-tongued bee tribe Anthophorini were investigated. Twenty-six ingroup taxa, representing all known genera and the most common subgenera of the Anthophorini, were included in a cladistic analysis of the tribe based on 51 morphological characters. The analysis yielded the generic relationships: (( Habrophorula , Elaphropoda ) ( Habropoda ( Deltoptila ( Pachymelus ( Amegilla , Anthophora ))))). The monophyly of the Anthophorini and each of its genera was confirmed. A second tribe, Habropodini, as postulated by some authors, is paraphyletic. On the basis of the results of the cladistic analysis and the biogeographical data of Anthophorini, an evolutionary scenario is postulated in which all genera of Anthophorini probably evolved in the Late Cretaceous, except Amegilla and Anthophora , which originated in the Oligocene. The northern part of south-east Asia (India to south-east China) is regarded as the most probable place of origin and radiation of the Anthophorini. The New World was probably colonized three times independently by Habropoda (upper Cretaceous to Tertiary), by an ancestral lineage of Deltoptila (upper Cretaceous to Tertiary) and by Anthophora (Tertiary to Quaternary).     

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。     

Notes on the position of the true freshwater crabs within the brachyrhynchan Eubrachyura (Crustacea: Decapoda: Brachyura)

Cladistic and phenetic relationships of 51 eubrachyuran crab genera, comprising 36 genera of marine crabs and 18 genera of true freshwater crabs from 7 families, were investigated using 121 parsimony-informative adult morphological characters. The data matrix was subjected to four different treatments: (1) a cladistic analysis with a combination of unordered and ordered characters, (2) a cladistic analysis with all characters unordered, (3) neighbour-joining, and (4) UPGMA phenetic analyses. The parsimony analysis conducted with a combination of ordered and unordered characters produced a set of hypotheses which supported monophyly of a Pseudothelphusidae+Potamoidea clade. Furthermore, exemplar genera of the Bythograeidae and Pinnotheridae formed an unresolved polytomy with the Pseudothelphusidae+Potamoidea group, the Thoracotremata. The trichodactylid freshwater crabs were positioned as the sister taxon of the basal portunoid Carcinus, but were unresolved relative to other portunoids and geryonids. Second, the parsimony analysis conducted with all characters unordered resulted in a [bythograeid, pseudothelphusid+potamoid, pinnotherid, thoracotreme] group with no hierarchical resolution, which in turn formed a polytomy with a goneplacid+portunoid clade and a polyphyletic Xanthoidea. And third, phenetic groupings of the eubrachyuran genera invariably placed the pseudothelphusids with the potamoids, and this clustered with a group containing the thoracotremes (either in whole or part). Support was thus found for morphological connections among the nontrichodactylid freshwater crabs, thoracotremes, bythograeids, and pinnotherids, and for the placement of the trichodactylids within the Portunoidea. These two latter findings (that used a range of genera from each family) are broadly congruent with a previous cladistic analysis of selected eubrachyuran familial groundpatterns that used a basal exemplar of each marine and freshwater crab family (Sternberg et al., 1999). However, it is clear that the large scale homoplasy identified here may nullify any reliable hypothesis of brachyrhynchan groupings at this stage.     

Cladistic and phenetic studies of genera of Afrotropical Cryptobiina (Coleoptera,Staphylinidae, Paederinae) II. The phenetic approach

The phenetics of 12 genera of Afrotropical Cryptobiina have been analysed by multivariate statistics. Three analyses (principal component analysis, canonical discriminant analysis and cluster analysis) have been used to determine the relative positions of the taxa in a morphospace. These analyses have allowed us to confirm the generic divisions based on alpha-systematic and phylogenetic grounds. While phenetic systematics and cladistic systematics have often been opposed, this study shows that both approaches are, in the end, complementary. Phenetical analysis allow, in addition to a quantitative definition of the genera, the isolation of ethological or ecological characters, while the cladistic approch gives to them precision and a phylogenetic sense. The three types of analyses point out the importance in Cryptobiina of the size, length of the scapus and width of the neck, all three characters which were also designated as significant characters by the phylogenetic analysis.     

螟黄足盘绒茧蜂复合群支序系统学研究（膜翅目：茧蜂科：小腹茧蜂亚科）

对螟黄足盘绒茧蜂复合群Cotesia flavipes complex (膜翅目：茧蜂科：小腹茧蜂亚科)分布全世界的5个种和外群侧沟茧蜂Microplitis 及荻茧蜂Diolcogaster 的25个性状,以及复合群5个种和外群螟蛉盘绒茧蜂Cotesia ruficrus、粘虫盘绒茧蜂Cotesia kariyai、粉蝶盘绒茧蜂Cotesia glomerata 的24个性状,分别进行比较研究,并运用支序分析的方法探讨该复合群内5个种种间的系统发育关系。支序分析表明螟黄足盘绒茧蜂复合群是一单系群,二化螟盘绒茧蜂C. chilonis 和大螟盘绒茧蜂C. sesamiae 近缘,芦螟盘绒茧蜂C. chiloluteelli 和汉寿盘绒茧蜂C. hanshouensis 近缘,螟黄足盘绒茧蜂C. flavipes相对独立。 以上研究表明无论是以近缘属作外群还是以同属其它种作外群,所得结果基本上都能反映螟黄足盘绒茧蜂复合群各种之间的分类地位。     

Integration of mitogenomic and morphological data disentangles the systematics of Pollenia and establishes a revised phylogenetic hypothesis for the Polleniidae

The Polleniidae (Diptera) are a family of flies best known for species of the genus Pollenia, which overwinter inside human dwellings. Previously divided across the Calliphoridae, Tachinidae and Rhinophoridae, the polleniid genera have only recently been united. Several studies have utilized molecular data to analyse polleniid phylogenetic relationships, although all have suffered from low taxon sampling or insufficient phylogenetic signal in molecular markers. To alleviate these problems, we utilized two automated organellar genome extraction software, GetOrganelle and MitoFinder, to assemble mitogenomes from genome skimming data from 22 representatives of the polleniid genera: Dexopollenia, Melanodexia, Morinia, Pollenia and Xanthotryxus. From these analyses, we provide 14 new mitogenomes for the Polleniidae and perform phylogenetic analyses of 13 protein-coding mitochondrial genes using both maximum likelihood and Bayesian inference. Subfamilial phylogenetic relationships within the Polleniidae are interrogated and Pollenia is found to form a monophyletic clade sister to Melanodexia, Morinia and Dexopollenia, providing no evidence for the synonymisation of any of these genera. Our topology conflicts with previous morphology-based cladistic interpretations, with the amentaria, griseotomentosa, semicinerea and viatica species-groups resolving as non-monophyletic. We provide support for our topology through analysis of adult morphology and male and female terminalia, while identifying new diagnostic characters for some of the clades of the Pollenia. To test the validity of the current diagnostic morphology in the Polleniidae, newly assembled cytochrome C oxidase subunit 1 (COI) data are combined with a polleniid COI barcode reference library and analysed using the species delimitation software ASAP. COI barcodes support the current morphologically defined species within the Pollenia.     

Preliminary morphological analysis of relationships between the spider wasp subfamilies (Hymenoptera: Pompilidae): revisiting an old problem

No qualitative cladistic analysis has been performed previously for the subfamily classification of Pompilidae (Hymenoptera). In 1994 Shimizu proposed six subfamilies, but their validity and relationships remain inconclusive. The objective of this study was to perform a quantitative analysis of phylogenetic relationships of the Pompilidae, with emphasis on testing the validity of proposed subfamilies. Two cladistic analyses were performed based on morphological evidence. First, a maximum-parsimony analysis of Shimizu's original morphological data matrix (72 taxa by 54 characters) was conducted, with the data subjected to a heuristic search for the first time with phylogenetic software. The resulting strict-consensus cladogram yielded a monophyletic Ceropalinae that was sister group to a large polytomy containing members of the remaining five subfamilies. In a second analysis, several of Shimizu's characters were re-examined, and new characters and more taxa were added to the data set. Terminal taxa were coded as species rather than as generic abstractions, and 20 additional morphological characters were introduced. The analysis was based on 77 morphological characters derived from the adults of 84 taxa. This second analysis suggested that Notocyphinae sensu Shimizu (1994) was nested within Pompilinae and that Epipompilinae sensu Shimizu (1994) was nested within Ctenocerinae; neither should retain their status as a separate subfamily. Lastly, Chirodamus s .s., which historically has been a member of the Pepsinae, is placed within the Pompilinae with reservations rather than erecting a new subfamily. After these allowances were made, a strict consensus tree gave the following relationships: (Ceropalinae + (Pepsinae + (Ctenocerinae + Pompilinae))).     

A phylogenetic analysis of the extant Aeshnidae (Odonata: Anisoptera)

Abstract A cladistic analysis of the world Aeshnidae is presented, based on fifty-eight characters of adult and larval anatomy. The ingroup taxa include all the extant genera of Aeshnidae, and the austropetaliid genera Phyllopetalia and Hypopetalia were chosen as the outgroup. The strict consensus tree obtained after successive weighting shows that the subgroups defined traditionally for Aeshnidae are paraphyletic or polyphyletic. The previous reclassification derived from analyses based on wing venation is supported in terms of the monophyly of Aeshnidae, Gomphaeschninae and its sister group comprising the remaining Aeshnidae. Gomphaeschninae is confirmed as sister group of the remaining Aeshnidae (= Aeshnodea Bechly). The sister-group relationships between Gomphaeschna + Sarasaeschna and Linaeschna + Oligoaeschna are corroborated. Within Aeshnodea, three monophyletic groups emerged: Boyeria + ( Petaliaeschna + ( Limnetron + Gynacanthaeschna + Periaeschna )) + (( Cephalaeschna + Caliaeschna ) + ( Allopetalia ( Notoaeschna + Spinaeschna ))); Dendroaeschna + ( Epiaeschna + ( Aeschnophlebia + ( Nasiaeschna + ( Tetracanthagyna + Brachytron )))); and Polycanthagyna + ( Basiaeschna + ( Amphiaeschna + ( Indaeschna + ( Oplonaeschna + ( Racenaeschna + Plattycantha + Agyrtacantha + Triacanthagyna + ( Subaeschna + Austrogynacantha + Gynacantha ) + ( Heliaeschna + ( Neuraeschna + Staurophlebia ))) + (( Castoraeschna + Coryphaeschna + Remartinia ) + ( Oreaeschna + ( Aeshna + ( Anaciaeschna + ((' A .' isosceles + Andaeshna ) + ( Anax + Hemianax )))))). Additional informative characters are required to test the relationships suggested here between the main groups of Aeshnodea and some enigmatic basal taxa ( Antipodophlebia , Austroaeschna , Acanthaeschna , Telephlebia , Austrophlebia and Planaeschna ).     

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.     

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.