The sperm pumps of Strepsiptera and Antliophora (Hexapoda)

Male genital structures of representatives of Strepsiptera, Siphonaptera and Diptera are described in detail, with special emphasis on sperm pumps. The parts involved in the apparatus are evaluated with respect to their homology. Functional interpretations are presented based on the morphological observations. The phylogenetic significance of characters related to the male genital apparatus is discussed. The sperm pumps differ strongly in Strepsiptera and Antliophora (s.s.) and are not homologous. The strepsipteran type, which lacks any sclerotized parts, has evolved independently. Autapomophies of the male genital apparatus are the compact testes, the large balloon‐shaped vesicula seminalis, the strongly developed musculature of the proximal ductus ejaculatorius, the strongly simplified copulatory organ, the unusual muscles of segments VIII and IX, and the complete absence of accessory glands. The median fusion and almost globular shape of the vesicula are potential autapomorphies of Corioxenidae. The absence of the furrow separating the testes from the vesicula seminalis is a derived condition found in Xenos and Myrmecolax. A sperm pump is absent in Boreus (Mecoptera) and Culicomorpha and the functionally relevant parts and their arrangement differ strongly in Siphonaptera, Pistillifera and Diptera (excl. Culicomorpha). The presence of a functional and homologous pumping apparatus does not belong to the groundplan of Antliophora, which implies that this alleged autapomorphy of the clade is invalid. A sperm pump belongs to the groundplan of Diptera and was secondarily reduced in Culicomorpha, many representatives of Bibionomorpha, and in Diopsidae. It was very likely primarily absent in Mecoptera. However, the precise reconstruction of the groundplan depends on the position of Nannochoristidae within Mecoptera and on the possible affinities of Siphonaptera and Boreidae. Sperm pumps should be considered as a functional term and not be used as a character for phylogenetic reconstruction, unless specific similarities are included in the character definition.     

Phylogeny of the holometabolous insect orders: molecular evidence

Phylogenetic relationships among the holometabolous insect orders were reconstructed using 18S ribosomal DNA data drawn from a sample of 182 taxa representing all holometabolous insect orders and multiple outgroups. Parsimony analysis supports the monophyly of all holometabolous insect orders except for Coleoptera and Mecoptera. Mecoptera is paraphyletic with respect to Siphonaptera, which is nested within Mecoptera. Coleoptera is scattered as a paraphyletic assemblage across the tree topology. These data support a monophyletic Halteria (Strepsiptera + Diptera), Amphiesmenoptera (Trichoptera + Lepidoptera), Neuropterida (Neuroptera + (Megaloptera + Raphidioptera)), but Antliophora (Halteria + Mecoptera + Siphonaptera) and Mecopterida (Antliophora + Amphiesmenoptera) are paraphyletic. The limitations of using 18S ribosomal DNA as the sole phylogenetic marker for reconstructing insect ordinal relationships are discussed.     

The evolution of sperm and non-sperm producing organs in male Drosophila

The male ejaculate is made up of two components: sperm and non-sperm. There has been little consideration of how these two basic compartments evolve. If they are subject to trade-offs, theory predicts that when the sperm competition raffle is unfair, when seminal fluid proteins stimulate fecundity and/or when ejaculate components alter fertilization success, there will be differential selection on sperm versus non-sperm ejaculate characteristics. However, the fundamental assumption that there are trade-offs between sperm and non-sperm ejaculate compartments in Drosophila has not yet been tested. To address this, we examined testis (sperm producing) and accessory gland (non-sperm producing) size across 22 species of Drosophila . We also examined how these characters varied with copulation duration, which may represent an additional target for sperm competition. The results showed no evidence of a trade-off between testis length and accessory gland length. Copulation duration correlated negatively with accessory gland length and there was a positive correlation with testis length, but only after correcting for body size. Overall, the results suggest no evidence for gross trade-offs in sperm versus non-sperm compartments across these Drosophila species, and motivate more detailed examination of ejaculate investment patterns.  © 2008 The Linnean Society of London, Biological Journal of the Linnean Society , 2008, 94 , 505–512.     

Revision of Pseudonereis (Polychaeta, Nereididae)

A taxonomic revision of Pseudonereis (Polychaeta, Nereididae) shows that some of the described taxa are very similar in most morphological characteristics. The revision includes all ten taxa considered valid, and are redescribed from type material. Lectotypes are designated for Pseudonereis anomala Gravier, 1901, Pseudonereis noodti (Hartmann-Schröder, 1962) and Pseudonereis trimaculata Horst, 1924. The widely geographically distributed and well-known P. gallapagensis Kinberg, 1865 and P. variegata ( Grube, 1857 ) show striking morphological resemblance to less well-known taxa with similar distribution. Paragnath variation in populations of P. anomala is discussed relating to its geographical distribution. Pseudonereis trimaculata is recorded from Australia for the first time. Taxa belonging to Pseudonereis are predominantly tropical and subtropical. A cladistic analysis using parsimony is included to test for monophyly of Pseudonereis . A monophyletic clade including all Pseudonereis taxa is given low bootstrap support. This clade is supported by the synapomorphies: presence of paragnaths in closely spaced comb-like rows on the maxillary ring on the pharynx, and presence of p-bar paragnaths in Areas II–IV and VII–VIII. Several of the included taxa share the shield-shaped paragnath in Area VI, which serves to distinguish Pseudonereis spp. from Perinereis spp. Paragnaths of the type p-bars and shield-shaped bar is described for the first time; the latter character is different from the smooth bar-shaped paragnaths in Area VI as has previously been described in these taxa.  © 2007 The Linnean Society of London, Zoological Journal of the Linnean Society , 2007, 150 , 145–176.     

The thoracic morphology of Nannochorista (Nannochoristidae) and its implications for the phylogeny of Mecoptera and Antliophora

External and internal thoracic structures of Nannochorista spp. are described in detail. The results are compared with conditions found in other endopterygote taxa, especially in members of Antliophora. Seventy-seven characters potentially useful for phylogenetic reconstruction are discussed, coded, presented as a data matrix and analysed cladistically. The thorax of Nannochorista shows a number of plesiomorphic characters compared with other mecopterans (except for Merope ) and members of the other antliophoran groups (e.g. presence of prospina and associated muscles). No specific affinities of thoracic features of Nannochoristidae and Diptera were found. The cladistic analysis results in strongly supported Antliophora (e.g. intraprofurcal muscle and ventral pleural arms present; bundle of M. mesonoto-pleuralis posterior originates on pleural arm). The thoracic characters do not support the monophyly of Mecoptera. This is possibly an artefact of the analysis. Several potential thoracic autapomorphies of the order are inapplicable in Boreidae, Siphonaptera and Diptera. Boreidae and Siphonaptera share a suite of characters related with flightlessness and are retrieved as sistertaxa when characters associated with wing reduction are predefined as irreversible. Merope appears exceptionally plesiomorphic in its thoracic morphology. Pistillifera (excluding Meropidae) and Panorpoidea (Panorpidae + Panorpodidae) are supported as clades. Due to the strongly modified thoracic morphology of Siphonaptera, the position of this group remains uncertain. The phylogenetic reconstruction using thoracic features alone is clearly impeded by far reaching modifications in Diptera in correlation with an advanced type of anteromotorism, and complex suites of reductional features in the secondary wingless forms.     

Phylogenetic analysis of Nymphaeales using fast-evolving and noncoding chloroplast markers

The Nymphaeales (water-lilies and relatives) represent one of the earliest branching lineages of angiosperms and comprise about 70 aquatic species. Here, we present a comprehensive study of phylogenetic relationships within the Nymphaeales from a dataset containing 24 representatives of the order, including all currently recognized genera and all subgenera of the genus Nymphaea , plus 5 outgroup taxa. Nine different regions of the chloroplast genome − comprising spacers, group II introns, a group I intron, and a protein coding gene − were analysed. This resulted in a character matrix of 6597 positions and an additional 369 characters obtained from coded length mutations. Maximum parsimony and Bayesian analyses of the complete dataset yielded congruent, fully resolved and well-supported trees. Our data confirm the monophyly of the Cabombaceae but do not provide convincing support for the monophyly of Nymphaeaceae with respect to Nuphar . Moreover, the genus Nymphaea is inferred to be paraphyletic with respect to Ondinea , Victoria and Euryale . In fact, the Australian endemic Ondinea forms a highly supported clade with members of the Australian Nymphaea subgenus Anecphya . In addition, Victoria and Euryale are inferred to be closely related to a clade comprising all night-blooming water-lilies ( Nymphaea subgenera Hydrocallis and Lotos ). An experimental approach showed taxon sampling to be of influence on the nodes reconstructed in core Nymphaeaceae. The results underscore that more diverse genera, if not clearly known to be monophyletic, should be represented by all major lineages.  © 2007 The Linnean Society of London, Botanical Journal of the Linnean Society , 2007, 154 , 141–163.     

Phylogenetic relationships among Acestrorhynchus species (Ostariophysi: Characiformes: Acestrorhynchidae)

A total of 104 osteological and external morphological features were examined in 13 species of Acestrorhynchus and 15 outgroup taxa to advance a hypothesis of relationships within the genus. Two most parsimonious hypotheses corroborate the monophyly of Acestrorhynchus but differ in the hypothesized relationships of Acestrorhynchus heterolepis . Three proposed supraspecific assemblages are at least partially correlated with groups of species previously diagnosed on the basis of colour pattern: (1) Acestrorhynchus britskii , Acestrorhynchus grandoculis , Acestrorhynchus microlepis , and Acestrorhynchus minimus ; (2) Acestrorhynchus falcirostris , Acestrorhynchus isalineae , and A. Acestrorhynchus nasutus ; and (3) Acestrorhynchus abbreviatus , Acestrorhynchus altus , Acestrorhynchus falcatus , Acestrorhynchus lacustris , and Acestrorhynchus pantaneiro . In one hypothesis A. heterolepis is proposed as the closest relative of the clade formed by A. falcirostris , A. isalineae , and A. nasutus , and in the alternative hypothesis it is proposed as a sister species of the clade formed by A. abbreviatus , A. altus , A. falcatus , A. lacustris , and A. pantaneiro . Relationships among species of the latter clade remain unresolved. Two independent episodes of reduction of body size are hypothesized to have occurred within the genus: one associated with the clade formed by A. grandoculis and A. minimus , and the other with the clade formed by A. isalineae and A. nasutus . © 2007 The Linnean Society of London, Zoological Journal of the Linnean Society , 2007, 151 , 691–757.     

Phylogenetic analysis of the Camaenidae (Mollusca: Stylommatophora) with special emphasis on the American taxa

The monophyly of the land snail family Camaenidae has been in doubt due to a disjunct bihemispheric distributional pattern and to the lack of morphological synapomorphies. A cladistic analysis is presented using an ingroup composed of representatives of the three subfamilies distributed in Australia and 52 other species with American distribution. Bradybaenidae, Helicidae and Helminthoglyptidae were used as outgroups. Fifty morphological characters were treated as unordered and analysed using Pee-Wee ver. 2.9, a program for parsimony analysis using implied weights. The results of the analysis support Camaenidae as a monophyletic family (synapomorphies: oval genital orifice, absence of penial sheath). Two of the three Australasian subfamilies, Sinumeloninae and Camaeninae, are monophyletic in the strict consensus tree. The American taxa are classified in eight genera and arranged into two main clades. Caracolus is proposed as the sister group of the American Continental Camaenidae. The genus Solaropsis , previously excluded from this family by different authors, is reassigned to Camaenidae. Shell characters proved to be phylogenetically informative in defining Pleurodonte , Caracolus , Solaropsis , Isomeria and Labyrinthus . © 2003 The Linnean Society of London, Zoological Journal of the Linnean Society, 2003, 138, 449–476.     

Historical influences on genital morphology among sympatric species: gonopod evolution and reproductive isolation in the crayfish genus Orconectes (Cambaridae)

Freshwater crayfishes represent an incredibly diverse component of temperate aquatic ecosystems. The genus Orconectes (Cambaridae) comprises approximately 17% of total global crayfish diversity and is native to central and eastern North America. Using both Bayesian and parsimony-based phylogenetic hypotheses from complete cytochrome c oxidase subunit I (COI) mtDNA sequence data (1545 bp) along with information on male gonopod shape and species distribution patterns, we examined the evolution of male genital morphology and its potential influence on species co-occurrence. We found that sympatric species exhibited similar male genital morphology more often than expected based on the frequency of genital shapes within the genus. When evolutionary history was incorporated into this analysis, the result was no longer significant, suggesting that species co-occurrence and the associated genital shapes of sympatric congeners cannot be explained solely by interspecific ecological interactions. Our results also do not support the current subgeneric classifications within Orconectes or the monophyly of the genus, both of which have been based previously on male genital morphology.  © 2006 The Linnean Society of London, Biological Journal of the Linnean Society , 2006, 89 , 1–12.