Are the genera of Nematomorpha monophyletic taxa?

Twenty-two genera of horsehair worms (Nematomorpha) have been described to date, one (Nectonema) from the marine environment and the remaining ones (taxon Gordiida) from freshwater. The aim of this investigation was to test whether the genera represent monophyletic taxa or are likely to be paraphyletic assemblages. The recognition and delimitation of monophyletic taxa are prerequisites for further systematic analyses. From the 15 nematomorph genera containing more than one species, autapomorphies can be found for five genera (Chordodes, Nectonema, Beatogordius, Acutogordius, Noteochordodes). In three further genera (Paragordius, Parachordodes, Gordionus), the evaluation of characters as autapomorphies is likely, but relies on further investigations. For seven genera ('Gordius', 'Paragordionus', 'Euchordodes', 'Neochordodes', 'Pseudochordodes', 'Spinochordodes', 'Digordius'), no autapomorphy could be found. Five genera ('Digordius', Lanochordodes, Progordius, Pantachordodes, Pseudogordius) are likely to be synonymous with other genera, and one genus ( Chordodiolus ) has been synonymized with Beatogordius recently. The importance of high quality documentation of characters in investigations of nematomorph species is stressed.     

Macrodasyida (Gastrotricha): a cladistic analysis of morphology

Abstract. A cladistic analysis based on 33 morphological characters was performed for the 31 genera currently assigned to the order Macrodasyida (Gastrotricha). Outgroup analysis indicated that the order is monophyletic and that it is defined by the structure of the pharynx and the complex distribution of duo-gland adhesive organs. Of the 6 currently recognized families in Macrodasyida, our analysis confirmed that 4 families are monophyletic: Dactylopodolidae, Macrodasyidae, Thaumastodermatidae and Turbanellidae. Dactylopodolidae was further confirmed as the most basal family within the order based on the retention of several plesiomorphies. The other three families have well-defined autapomorphies but will require further investigation to increase inter- and intrafamilial phylogenetic resolution. Planodasyidae appeared to be a paraphyletic taxon with no obvious autapomorphies; genera clustered among members of a polyphyletic family, Lepidodasyidae. We recommend that future research on macrodasyidan phylogeny focus on issues of comparative morphology and ultrastructure in lesser-known taxa such as the Dactylopodolidae, and on the taxa Lepidodasyidae and Planodasyidae.     

Signal through the noise? Phylogeny of the Tachinidae (Diptera) as inferred from morphological evidence

The oestroid family Tachinidae represents one of the most diverse lineages of insect parasitoids. Despite their broad distribution, diversity and important role as biological control agents, the phylogeny of this family remains poorly known. Here, we review the history of tachinid systematics and present the first quantitative phylogenetic analysis of the family based on morphological data. Cladistic analyses were conducted using 135 morphological characters from 492 species belonging to 180 tachinid genera, including the four currently recognized subfamilies (Dexiinae, Exoristinae, Phasiinae, Tachininae) and all major tribes. We used characters of eggs, first‐instar larvae and adults of both sexes. We examined the effects of implied weighting by reanalysing the data with varying concavity factors. Our analysis generally supports the subfamily groupings Dexiinae + Phasiinae and Tachininae + Exoristinae, with only the Exoristinae and the Phasiinae reconstructed as monophyletic assemblages under a wide range of weighting schemes. Under these conditions, the Dexiinae, which were previously considered a well‐established monophyletic assemblage, are reconstructed as being paraphyletic with respect to the Phasiinae. The Tachininae are reconstructed as a paraphyletic grade from which the monophyletic Exoristinae arose. The Exoristinae are reconstructed as a monophyletic lineage, but phylogenetic relationships within the subfamily are largely unresolved. We further explored the evolution of oviposition strategy and found that the oviparous groups are nested within ovolarviparous assemblages, suggesting that ovipary may have evolved several times independently from ovolarviparous ancestors. This counterintuitive pattern is a novel hypothesis suggested by the results of this analysis. Finally, two major patterns emerge when considering host associations across our phylogeny under equal weights: (i) although more than 60% of tachinids are parasitoids of Lepidoptera larvae, none of the basal clades is unambiguously associated with Lepidoptera as a primitive condition, suggesting that tachinids were slow to colonize these hosts, but then radiated extensively on them; and (ii) there is general agreement between host use and monophyly of the major lineages.     

Cerapeplus gen.n. and the classification of micropeplids (Coleoptera: Micropeplidae)

Cerapeplus siamensis gen.n., sp.n. from Thailand is described and illustrated. The new genus is characterized by filament-based lamellar secretions arising from the lateral body margin, a feature observed for the first time in Coleoptera. Possible autapomorphies for the micropeplid genera are discussed. It is concluded that Cerapeplus and Kalissus are monophyletic; Micropeplus and/Or Peplomicrus are probably polyphyletic but both genera together form a monophyletic group. A number of characters support the family status of the micropeplids recently proposed by some authors. A revised family diagnosis is provided.     

Phylogenetic relationships of the genus Dugesia (Platyhelminthes, Tricladida, Paludicola)

A phylogenetic analysis was performed on the genera and subgenera within the freshwater triclad family Dugesiidae, based on 19 terminal taxa and 17 morphological characters. The phylogenetic tree proposed has length of 27 steps and consistency index of 0.66. This phylogenetic hypothesis implies that the current genus Dugesia is paraphyletic and that its subgenera Girardia, Schmidiea and Dugesia S.S. should be elevated to the rank of genus. The genera Cura, Spathula and Neppia are presumed monophyletic by default because the database was unable to provide autapomorphies for any of these genera. The genera Dugesia S.S. and Neppia share sistergroup relationship. Several characters are discussed which were previously considered to be of phylogenetic importance but were not included in the present analysis. It is emphasized that sensory organs form potentially useful set of phylogenetic characters for the Dugesiidae.     

A phylogenetic analysis of the genus Eunice (Eunicidae, polychaete, Annelida)

Species of Eunice are distributed worldwide, inhabiting soft and hard marine bottoms. Some of these species play significant roles in coral reef communities and others are commercially important. Eunice is the largest and most poorly defined genus in Eunicidae. It has traditionally been subdivided in taxonomically informal groups based on the colour and dentition of subacicular hooks, and branchial distribution. The monophyly of Eunice and of its informal subgroups is tested here using cladistic analyses of 24 ingroup species based on morphological data. In the phylogenetic hypothesis resulting from the present analyses Eunice and its subgroups are paraphyletic; the genus may be divided in at least two monophyletic groups, Eunice s.s. and Leodice , but several species do not fall inside these two groups. Most of the traditional characters used in the taxonomy of Eunice are homoplasies; however, characters used for the first time in this study, such as certain jaw characters and characters derived from a close examination of chaetal variation along the body, are promising sources of phylogenetic signal.  © 2007 The Linnean Society of London, Zoological Journal of the Linnean Society , 2007, 150 , 413–434.     

Morphology,molecules and the phylogeny of Characidae (Teleostei,Characiformes)

This is the most comprehensive phylogenetic analysis of the Characidae to date and the first large-scale hypothesis of the family, combining myriad morphological data with molecular information. A total of 520 morphological characters were analysed herein, of which 98 are newly defined. Among the analysed taxa, 259 species were coded by examining specimens, three fossil species were coded from the literature, one species was coded almost completely from published figures, 122 were partially coded from the literature, and 88 were analysed exclusively from molecular data. The total number of species in the analysed dataset is 473. Analyses were made by parsimony under equal and extended implied weighting with a broad range of parameters. The final hypothesis was selected using a stability criterion that chooses among the most parsimonious trees of all searches. It was found by weighting molecular characters with the average homoplasy of entire partitions (markers). The resulting hypothesis is congruent with previous molecular-based phylogenies of the family. The Characidae are monophyletic, with four main clades: the Spintherobolinae new subfamily; an expanded Stethaprioninae including the Grundulini, Gymnocharacini, Rhoadsiini and Stethaprionini; the Stevardiinae; and a clade composed of the Aphyocharacinae, Characinae, Cheirodontinae, Exodontinae and Tetragonopterinae. Also, a stem Characidae was found, as formed by the Eocene–Oligocene genera †Bryconetes and †Paleotetra as successive sister groups of extant members of the family. A subfamilial classification is proposed, but deep changes in the systematics that are beyond the scope of this study are still needed to classify the Characidae into monophyletic genera.     

Cladistic analysis of the alderflies of America north of Mexico (Megaloptera: Sialidae)

Abstract. A cladistic analysis of thirty-five morphological characters is used to produce a cladogram for the twenty-four American species of Sialis found north of Mexico, two European species, and three species of Protosialis . Three species groups ( americana, califomica and infumata ) proposed by Ross 1937 are shown to be monophyletic. Sialis, Protosialis , and the aequalis species group as currently constituted, are paraphyletic; S.nevadensis is nested within Protosialis and Protosialis is nested within Sialis . Character evolution is evaluated in the light of a phylogenetic hypothesis, and structural modifications are found to correlate with presumed functional constraints.     

A Posteriori and a Priori Methodologies for Testing Hypotheses of Causal Processes in Vicariance Biogeography

Methods used in vicariance biogeography fall into the categories of a posteriori methods (e.g., Component Compatibility Analysis and Brooks Parsimony Analysis) and a priori methods (e.g., Component Analysis, Reconciled Tree Analysis, and Three Area Statement Analysis). Each category corresponds to a particular methodology that arrives at general area cladograms by testing null hypotheses in a particular way. A posteriori methods assume the process of vicariance only (A0) as a common cause of the distribution of different monophyletic groups of taxa under the null hypothesis. Whenever a parsimony analysis of combined data from these monophyletic groups results in a general area cladogram with homoplasy, the null hypothesis is rejected and extinction and dispersal are invoked a posteriori as ad hoc process explanations. A priori methods assume not only vicariance (A0) but also combinations of vicariance with the processes of extinction (A1) and dispersal (A2) as possible causes of the distribution of the taxa of different monophyletic groups. Each assumed set of processes corresponds to a different null hypothesis. Under the assumption of independence and thus additivity of the processes involved, the sets of area cladograms obtained under A0, A1, and A2 from data of each monophyletic group must be inclusive (requirement I). Whenever no congruent area cladograms are found in the intersection of sets of area cladograms derived under the same assumption for different monophyletic groups (II), the corresponding null hypothesis is rejected.     

Regional integration of evidence for evolution in the Silurian Pentamerus-Pentameroides lineage

Under ideal conditions, the stratophenetic test for phyletic gradualism in fossil lineages requires nearly continuous samples of time-specific populations through rock sequences representing geologically significant periods of time. Shifting environments account for one source of 'imperfection' in the local rock record. Few species are sufficiently eurytopic to survive changing environments in a given region over a long time span. Where marine fossils are involved, independently correlated sea-level curves may be used to patch together segments of stratigraphic sections from different regions that collectively encompass the uninterrupted environment of the target species. In the Lower Silurian of Iowa, stratigraphic gaps among test samples for phyletic gradualism in pentamerid brachiopods occur wherever pentamerid communities are succeeded by deeper water stricklandiid communities or by shallower-water coral-algal communities. Correlation of sea-level curves indicates that water depth was consistently shallower in the northern Great Lakes area. Usually when stricklandiid communities replaced a pentamerid community in Iowan seas, contemporaneous pentamerid communities replaced a coral-algal community in Michigan or Ontario seas. Temporal meshing of samples from Iowa and the northern Great Lakes area (based on congruent Stricklandia and Eocoelia lineage zones) supports the hypothesis that Pentamerus oblongus evolved to Pentameroides subrectus through a gradual narrowing and loss of divergence in its outer plates. A transitional morphotype between the two genera (from the Lower Silurian of Alabama) is illustrated. □ Phyletic gradualism, punctuated equilibria, Brachiopoda, Pentamerida, Silurian, North America.     

Marine insects: genital morphology, phylogeny and evolution of sea skaters, genus Halobates (Hemiptera: Gerridae)

Five species of sea skaters, genus Halobates Eschscholtz, are the only insects to have successfully colonized the open ocean. In addition, 36 species are found in sheltered coastal waters throughout tropical Indo-Pacific. The taxonomy of the genus is relatively well known, but reliable hypotheses about phylogenetic relationships are required if the biogeography and evolution of sea skaters is to be discussed in a meaningful way. This work presents the results of a study of new characters from the genital segments, especially those of the male phallus and the female gynatrial complex, and a reinterpretation for several other characters. In total 64 characters were scored for 26 species of Halobates , two species ofAsclepios and one species of Metrocoris. With Asclepios and Metrocoris species as outgroups, the character state sets were analysed cladistically using the computer program Hennig86. After critical evaluations of both characters and clades, a phylogeny for Halobates is presented and its taxonomic implications are discussed. A number of monophyletic species groups are delimited. One genus-level synonymy and three species-level synonymies are suggested. The evolution of Halobates is discussed in the light of the reconstructed phylogeny and present knowledge of the ecology and behaviour of sea skaters. A hypothesis of ecological evolution in halobatine water striders is proposed and tested.     

Species concepts and the recognition of ancestors

Whether or not ancestral species can be recognised depends on the species concept adopted. A “metaspecies”; is a species that completely lacks autapomorphies, and which might (or might not) be ancestral to other species. Such taxa have been identified among both living and fossil organisms. Under the most commonly‐used species concepts (biological, evolutionary, phenetic, phylogenetic, ecological, recognition and cohesion), “metaspecies”; can be assumed to be ancestral. Even if the known members of a metaspecies are not ancestral to anything, parsimony dictates that the (as yet unknown) ancestral lineage is identical to the metaspecies and, under these species concepts, assignable to the same species. Only the cladistic and monophyletic species concepts would deny “metaspecies”; ancestral status, but these species concepts are problematical and have never been used by practising systematists.     

Larval morphology and phylogenetic position of Micromalthus debilis LeConte (Coleoptera: Micromalthidae)

Abstract External and internal structures of the cerambycoid and triungulin larvae of Micromalthus debilis are described and compared to features found in larvae of other groups of Coleoptera. The morphological data are evaluated with respect to the systematic position of Micromalthidae. A cladistic analysis was carried out with fifty characters. Micromalthidae are not closely related to Lymexylidae (Polyphaga: Cucujiformia) but belong to Archostemata, which is confirmed as a monophyletic unit. Micromalthidae are specialized in terms of morphology and life cycle and are characterized by a considerable number of larval autapomorphies. Their sister-group relationship with Cupedidae is supported by several apomorphic features, which are probably correlated with xylobiontic habits: head transverse and strongly rounded laterally, absence of stemmata, shortened antennae, presence of sternal asperities and presence of eversible lobes of segment IX. Cupedidae is monophyletic and Priamca is the sister group of the remaining genera of Cupedidae included in the analysis. A closer relationship between Tenomerga and Rhipsideigma is supported by several larval synapomorphies. The ancestral life style of larvae of Archostemata was probably xylobiontic. This is suggested by derived groundplan features of the suborder, which are also found in larvae of non-related, wood-associated families.     

NULL MODELS FOR THE NUMBER OF EVOLUTIONARY STEPS IN A CHARACTER ON A PHYLOGENETIC TREE

Random trees and random characters can be used in null models for testing phylogenetic hypothesis. We consider three interpretations of random trees: first, that trees are selected from the set of all possible trees with equal probability; second, that trees are formed by random speciation or coalescence (equivalent); and third, that trees are formed by a series of random partitions of the taxa. We consider two interpretations of random characters: first, that the number of taxa with each state is held constant, but the states are randomly reshuffled among the taxa; and second, that the probability each taxon is assigned a particular state is constant from one taxon to the next. Under null models representing various combinations of randomizations of trees and characters, exact recursion equations are given to calculate the probability distribution of the number of character state changes required by a phylogenetic tree. Possible applications of these probability distributions are discussed. They can be used, for example, to test for a panmictic population structure within a species or to test phylogenetic inertia in a character's evolution. Whether and how a null model incorporates tree randomness makes little difference to the probability distribution in many but not all circumstances. The null model's sense of character randomness appears more critical. The difficult issue of choosing a null model is discussed.     

Status of Corynosoma (Acanthocephala: Polymorphidae) based on anatomical, ecological, and phylogenetic evidence, with the erection of Pseudocorynosoma n. gen

The possession of genital spines has been considered as a key taxonomic trait to differentiate Corynosoma from other genera of the Polymorphidae. However, Corynosoma currently consists of 2 groups of species with clear ecological and morphological divergences: the "marine" group (with ca. 30 species) infects mammals and piscivorous birds in the marine realm, whereas the "freshwater" group (with ca. 7 species) infects waterfowl in continental waters. Species from these groups differ in shape of body and neck, trunk spination, lemnisci length and shape, testes arrangement, and number and shape of cement glands. We tested whether species from these 2 groups formed a monophyletic assemblage based on a phylogenetic analysis by using 15 morphological characters. We also included species of Andracantha, Polymorphus, and Hexaglandula with which potential taxonomic conflicts could most likely arise. We obtained 108 equally most parsimonious trees of 32 steps, with a consistency index (CI) = 0.59, and a retention index (RI) = 0.82. The strict consensus tree indicated that the "freshwater" species of Corynosoma form a monophyletic assemblage closely related to some species of Polymorphus, whereas the "marine" species of Corynosoma are grouped together with Andracantha. Accordingly, Corynosoma is not a monophyletic assemblage, and Pseudocorynosoma n. gen. is proposed for the "freshwater" species of Corynosoma. This decision was strongly supported by (1) a functional comparison of foretrunk muscles between species of Polymophus, Andracantha, and Corynosoma; (2) a multivariate morphometric study of proboscis characters and egg size; and (3) an analysis of ecological patterns of host-parasite relationships.     

Molecular phylogeny of the subfamilies in Geometridae (Geometroidea: Lepidoptera).

Molecular sequence data from three gene fragments were used to examine critically a provisional phylogenetic classification based on morphological characters of the Geometridae, one of the most species-rich families of moths. The sister group relationship between Geometridae and Drepanidae gained further support from the molecular analysis, which was based on the ND1 mitochondrial gene and the first and second expansion segments of the 28S ribosomal RNA gene. Although the alignment of the second expansion segment contained regions with many gaps, it provided the most resolution of the gene fragments. Parsimony analysis of the combined data resulted in a cladogram in which species belonging to Drepanidae, Larentiinae, and Sterrhinae formed monophyletic groups. The Ennominae did not form a monophyletic group but rather were contained within a broader monophyletic group including Archiearinae, Geometrinae, and Alsophilinae (represented by only one species per group in the present study). The molecular results were used to explore further the relationship between Sterrhinae and Larentiinae, the question as to whether Ennominae actually represent a monophyletic group, and the relationships between Ennominae and some of the other subfamilies.