Molecules, development and fossils in the study of metazoan evolution; Articulata versus Ecdysozoa revisited

Two conflicting hypotheses of protostome relationships, Articulata and Ecdysozoa, are reviewed by evaluating the evidence in favor and against each one of them. Understanding early embryonic development and segmentation in non-arthropod non-annelid protostomes seems crucial to the debate. New ways of coding metazoan matrices, avoiding ground-patterns and higher taxa, and incorporating fossil evidence seems the best way to avoid circular debates. Molecular data served as the catalyzer for the Ecdysozoa hypothesis, although morphological support had been implicitly suggested. Most molecular analyses published so far have shown some support for Ecdysozoa, whereas none has ever supported Articulata. Here, new analyses of up to four nuclear loci, including 18S rRNA, myosin heavy chain II, histone H3 and elongation factor 1- are conducted to test the molecular support for Ecdysozoa, and, at least under some parameter sets, most data sets show a clade formed by the molting animals. In contrast, support for Articulata is not found under any analytical conditions.     

The position of the Arthropoda in the phylogenetic system

Traditionally, Panarthropoda (Euarthropoda, Onychophora, Tardigrada) are regarded as being closely related to Annelida in a taxon Articulata, but this is not supported by molecular analyses. Comparisons of gene sequences suggest that all molting taxa (Panarthropoda, Nematoda, Nematomorpha, Priapulida, Kinorhyncha, Loricifera) are related in a monophyletic taxon Ecdysozoa. An examination of the characters supporting Articulata reveals that only segmentation with a teloblastic segment formation and the existence of segmental coelomic cavities with nephridia support the Articulata, whereas all other characters are modified or reduced in the panarthropod lineage. Another set of characters is presented that supports the monophyly of Ecdysozoa: molting under influence of ecdysteroid hormones, loss of locomotory cilia, trilayered cuticle and the formation of the epicuticle from the tips of epidermal microvilli. Comparative morphology suggests Gastrotricha as the sister group of Ecdysozoa with the synapomorphies: triradiate muscular sucking pharynx and terminal mouth opening. Thus there are morphological characters that support Articulata, but molecular as well as morphological data advocate Ecdysozoa. Comparison of both hypotheses should prompt further thorough and targeted investigations. J. Morphol. 238:263–285, 1998. © 1998 Wiley-Liss, Inc.     

Metaves,Mirandornithes, Strisores and other novelties – a critical review of the higher‐level phylogeny of neornithine birds

Recent hypotheses on the higher‐level phylogeny of modern birds are reviewed, and areas of agreement and major conflict are detailed, with emphasis being put on congruence among independent molecular and morphological data sets. Although molecular data significantly contributed to a better understanding of avian phylogeny, they do not seem to be free of homoplasy and caution is warranted in the interpretation of some results. The recently proposed ‘Metaves’ clade is likely to be an artefact of the β‐fibrinogen gene, and current molecular data do not yield well‐supported phylogenies for some groups whose interrelationships can be resolved with morphological evidence. There exists, however, congruent and strong molecular evidence for several novel clades that were not recognized by morphologists before, and to ease future discussions the terms Picocoraciae (non‐leptosomid ‘Coraciiformes’ and Piciformes) and Aequornithes (‘waterbird assemblage’) are introduced. Molecular studies further congruently recover some clades, which have not yet been adequately appreciated and are outlined in the present review.     

Molecular phylogeny of the Forcipulatacea (Asteroidea: Echinodermata): systematics and biogeography

We present a comprehensively sampled three‐gene phylogeny of the monophyletic Forcipulatacea, one of three major lineages within the crown‐group Asteroidea. We present substantially more Southern Hemisphere and deep‐sea taxa than were sampled in previous molecular studies of this group. Morphologically distinct groups, such as the Brisingida and the Zoroasteridae, are upheld as monophyletic. Brisingida is supported as the derived sister group to the Asteriidae (restricted), rather than as a basal taxon. The Asteriidae is paraphyletic, and is broken up into the Stichasteridae and four primary asteriid clades: (1) a highly diverse boreal clade, containing members from the Arctic and sub‐Arctic in the Northern Hemisphere; (2) the genus Sclerasterias; (3) and (4) two sister clades that contain asteriids from the Antarctic and pantropical regions. The Stichasteridae, which was regarded as a synonym of the Asteriidae, is resurrected by our results, and represents the most diverse Southern Hemisphere forcipulatacean clade (although two deep‐sea stichasterid genera occur in the Northern Hemisphere). The Labidiasteridae is artificial, and should be synonymized into the Heliasteridae. The Pedicellasteridae is paraphyletic, with three separate clades containing pedicellasterid taxa emerging among the basal Forcipulatacea. Fossils and timing estimates from species‐level phylogeographic studies are consistent with prior phylogenetic hypotheses for the Forcipulatacea, suggesting diversification of basal taxa in the early Mesozoic, with some evidence for more widely distributed ranges from Cretacous taxa. Our analysis suggests a hypothesis of an older fauna present in the Antarctic during the Eocene, which was succeeded by a modern Antarctic fauna that is represented by the recently derived Antarctic Asteriidae and other forcipulatacean lineages. © 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 162 , 646–660.     

A multidisciplinary approach to the fine-systematics within Tisbe - an evaluation of morphological and molecular methods

Six species from the species-rich taxon Tisbe (Copepoda, Harpacticoida) were selected that could be reared in the laboratory as mass cultures. Phylogenetic relationships among these species were assessed by morphological studies of adults and larvae, DNA restriction site polymorphisms, allozymic, immunological distance, and lipid composition. Limits of scope and practicability of these analyses became apparent, as well as their potential and importance for future work in zoological systematics.     

Molecular phylogeny of Gastrotricha on the basis of a comparison of the 18S rRNA genes: Rejection of the hypothesis of a relationship between Gastrotricha and Nematoda

Gastrotricha are the small meiobenthic acoelomate worms whose phylogenetic relationships between themselves and other invertebrates remain unclear, despite all attempts to clarify them on the basis of both morphological and molecular analyses. The complete sequences of the 18S rRNA genes (8 new and 7 known) were analyzed in 15 Gastrotricha species to test different hypotheses on the phylogeny of this taxon and to determine the reasons for the contradictions in earlier results. The data were analyzed using both maximum likelihood and Bayesian methods. Based on the results, it was assumed that gastrotrichs form a monophyletic group within the Spiralia clade, which also includes Gnathostomulida, Plathelminthes, Syndermata (Rotifera + Acanthocephala), Nemertea, and Lophotrochozoa. Statistical tests rejected a phylogenetic hypotheses considering Gastrotricha to be closely related to Nematoda and other Ecdysozoa or placing them at the base of the Bilateria tree, close to Acoela or Nemertodermatida. Among gastrotrichs, species belonging to the orders Chaetonotida and Macrodasyida form two well-supported clades. The analysis confirmed monophyly of the families Chaetonotidae and Xenotrichulidae from the order Chaetonida, as well as the families Turbanellidae and Thaumastodermatidae from the order Macrodasyida. Lepidodasyidae is a polyphyletic family, because the genus Mesodasys forms a sister group for Turbanellidae; genus Cephalodasys forms a separate branch at the base of Macrodasyida; and Lepidodasys groups with Neodasys between Thaumastodermatidae and Turbanellidae. To confirm these conclusions and to get an authentic view of the phylogeny of Gastrotricha, it is necessary to study more Gastrotricha species and to analyze some other genes.     

Molecular phylogeny of the Paleogene fungus gnat tribe Exechiini (Diptera: Mycetophilidae) revisited: Monophyly of genera established and rapid radiation confirmed

The phylogeny of the fungus gnat tribe Exechiini (Diptera: Mycetophilidae) is reconstructed based on the combined analysis of five nuclear (18S, two parts of 28S, CAD, EF1α) and two mitochondrial (12S, COI) gene markers. According to known fossil record, and recent higher‐level phylogenies, the tribe constitutes the most apomorphic, distinctly monophyletic clade of the family Mycetophilidae. The tribe originated in the Paleogene and apparently quickly diversified in the Neogene with an unusual rapid radiation of complex male terminalia. Earlier attempts to reconstruct the phylogeny of the tribe, based on both morphology and molecular methods, have not yielded reliable hypotheses, neither in terms of resolution nor in terms of support for major clades. Increased taxon sampling and wider gene sampling have been suggested to achieve better phylogenetic resolution. Aiming at this, we present new phylogenies, for the first time with all known genera and subgenera of Exechiini represented. While many terminal intergeneric relationships are well supported, both in maximum likelihood and in Bayesian analyses, most of the major, deeper clades remain poorly supported. We suggest that a rapid radiation event close to the root may be causing the low resolution at this level in the phylogeny. This contrasts parallel phylogenies of the older subfamilies and tribes of the family Mycetophilidae, where traditional clades have usually been recovered with high support. Further in‐depth studies into the evolutionary history of the tribe are needed to enlighten and coalesce the specific phenomena driving their unique morphological, genetic and phylogeographic histories.     

Systematics and phylogeny of the tribe Paragini (Diptera: Syrphidae) based on molecular and morphological characters

We studied the phylogeny and systematics of the tribe Paragini (Diptera: Syrphidae) using morphological and molecular data. The paper presents separate parsimony analyses of both adult morphological characters and partial DNA sequence data from mitochondrial cytochrome c oxidase I and nuclear ribosomal 28S rRNA gene, as well as a combined analysis of all the data. The data set of morphological characters included some features of the male terminalia (i.e. shape of the ejaculatory apodeme; relative position of elements of the aedeagal complex; shape of surstylar apodeme; shape of the aedeagal apodeme) not previously used in the systematics of the Paragini. The trees obtained from separate parsimony analyses of molecular and morphological data produced almost identical topologies. Four lineages are supported by the combined data set, and we establish two new subgenera, i.e. Serratoparagus Vujić et Radenković subgen. nov., and Afroparagus Vujić et Radenković subgen. nov., and redefine Pandasyopthalmus Stuckenberg, 1954 stat. rev. and Paragus Latreille, 1804, stat. rev. The monophyly of the Pandasyopthalmus clade, including the species fitting neither of the current species groups ( jozanus -group) of Paragini, is established. Diagnoses of all known species groups are presented, including a new arrangement of almost all valid species of Paragini.  © 2008 The Linnean Society of London, Zoological Journal of the Linnean Society , 2008, 152 , 507–536.     

Molecular evidence for the phylogeny of Australian gekkonoid lizards

Partial sequences of the mitochondrial 12S rRNA and nuclear c-mos genes were determined for 12 species of gekkonoid lizards representing the four major taxa of the Australian region, the Diplodactyhni and Carphodactylini (forming the subfamily Diplodactylinae), the Pygopodidae and the Gekkoninae. One further species represented a non-Australian gekkonoid lineage, the Eublepharinae. The combined sequence data were used to reconstruct the underlying molecular phylogeny. We used the molecular phylogeny to test the monophyly of the diplodactyline tribes and conflicting hypotheses of relationships of die pygopods and of the genus Oedura. Monophyly of the Diplodactylinae is supported, while pygopods form a monophyletic sister lineage to all Diplodactylinae. The molecular data support the monophyly of the Diplodactyhni, with Oedura firmly placed as a diplodactylin. Monophyly of the Carphodactylini is not supported. The four carphodactylin genera form a paraphyletic cluster at the base of the Diplodactyhni. Pygopods are nested within the traditional Gekkonidae and pygopods plus diplodactylines form a well-supported monophyletic group with respect to the remaining gekkonoids, the gekkonines and eublepharines.     

On the information content of characters in comparative morphology and molecular systematics

A review of the fundamental difference between single molecular-sequence positions, or numerical characters, and complex morphological characters is the subject of this study. It has been found that transformation series of single complex structures contain enough information to allow a priori determination of character order and that rooting of a dendrogram is possible without out-group comparison, while trees based on less-informative characters can usually only be rooted with out-group comparison. Furthermore, the quality of total information used is decisive in discriminating between hypotheses of relationships. Numerical methods for the inference of phylogenies have been found to be useful for high numbers of characters that have only a low information content, while the Hennigian procedure seems to be preferable for complex characters.     

High-level phylogeny of early Tertiary rodents: dental evidence

Major crown‐groups of rodents were well established in the early Tertiary, and fossils provide an invaluable window into their evolutionary history. The main focus of this project was to perform a cladistic assessment of the dental evidence for early Tertiary rodent cladogenesis – the masticatory apparatus and teeth are the most frequently preserved anatomical features in the fossil record. We focused on groups that existed in a period corresponding to their early history, combining fossils belonging to extinct lineages and to stem‐groups leading to modern lineages. While the monophyly of some groups is not systematically explored, our results have important implications for high‐level rodent relationships and systematics. These results are consistent with those of recent molecular phylogenies and reliably congruent with the stratigraphic record, thus enhancing the pertinence of dental characters for phylogenetic inference. Our approach provides evidence of a fundamental dichotomy in early rodent history. Two major clades have been identified: (1) the earliest ‘ctenodactyloid’ (Ctenodactylidae, Chapattimyidae, Yuomyidae, Diatomyidae) and hystricognathous (Tsaganomyidae, Baluchimyinae, ‘phiomorphs’, ‘caviomorphs’) rodents, and (2) the earliest ‘ischyromyoid’ rodents with their closest relatives (Muroidea + Dipodoidea + Geomyoidea + Anomaluroidea + Castoroidea + Sciuravidae + Gliroidea, and Sciuroidea + Aplodontoidea + Theridomorpha). This topology has led us to endorse Ctenohystrica as the first clade and propose a new taxon, Ischyromyiformes, for the second. Although minimized in our working hypothesis, the homoplasy in dental characters remains significant. However, a number of homoplasic characters reveal structuring in their internal distribution, allowing us to discern evolutionary morphological patterns, notably the pentalophodonty of molars, zygomasseteric complex and incisor enamel microstructure. © 2004 The Linnean Society of London, Zoological Journal of the Linnean Society, 2004, 142 , 105–134.     

On the information content of characters in comparative morphology and molecular systematics

A review of the fundamental difference between single molecular-sequence positions, or numerical characters, and complex morphological characters is the subject of this study. It has been found that transformation series of single complex structures contain enough information to allow a priori determination of character order and that rooting of a dendrogram is possible without out-group comparison, while trees based on less-informative characters can usually only be rooted with out-group comparison. Furthermore, the quality of total information used is decisive in discriminating between hypotheses of relationships. Numerical methods for the inference of phylogenies have been found to be useful for high numbers of characters that have only a low information content, while the Hennigian procedure seems to be preferable for complex characters.

Zusammenfassung

Ein fundamentaler Unterschied zwischen der einzelnen Sequenzposition oder auch numerischen Merkmalen und komplexen morphologischen Merkmalen ist ihr Informationsgehalt. Merkmalsreihen komplexer Strukturen enthalten meist genügend Information, um a priori die Bestimmung der Lesrichtung zu ermöglichen. Die Feststellung des Ursprunges eines Dendrogramms ist somit ohne kladistischen Außengruppenvergleich möglich, während Bäume (Topologien), die auf wenig informativen Merkmalen beruhen, allgemein nur mit dem kladistischen Außengruppenvergleich 'gewurzelt' werden können. Die Qualität der insgesamt verwendeten Information ist entscheidend für die Wahl zwischen alternativen Verwandtschaftshypothesen. Numerische Methoden der Rekonstruktion der Phylogenese sind nützlich bei Verwendung einer gro β en Zahl informationsarmer Merkmale; das Hennigsche Verfahren ist für komplexe Merkmale vorzuziehen.     

Intergeneric phylogenetic relationships in catfishes of the Loricariinae (Siluriformes: Loricariidae), with the description of Fonchiiloricaria nanodon: a new genus and species from Peru

Recent investigations in the upper Río Huallaga in Peru revealed the presence of an intriguing species of the Loricariinae. To characterize and place this species within the evolutionary tree of the subfamily, a molecular phylogeny of this group was inferred based on the 12S and 16S mitochondrial genes and the nuclear gene F-reticulon4. The phylogeny indicated that this distinctive species was a member of the subtribe Loricariina. Given its phylogenetic placement, and its unusual morphology, this species is described as a new genus and new species of Loricariinae: Fonchiiloricaria nanodon. This new taxon is diagnosed by usually possessing one to three premaxillary teeth that are greatly reduced; lips with globular papillae on the surface; the distal margin of lower lip bearing short, triangular filaments; the premaxilla greatly reduced; the abdomen completely covered by plates, with the plates between lateral abdominal plates small and rhombic; a caudal fin with 14 rays; the orbital notch absent; five lateral series of plates; dorsal-fin spinelet absent; preanal plate present, large and solid, and of irregular, polygonal shape, the caudal peduncle becoming more compressed posteriorly for the last seven to 10 plates.     

Revisiting the phylogeny of papilionoid legumes: New insights from comprehensively sampled early-branching lineages

• Premise of study: Phylogenetic relationships of the papilionoid legumes (Papilionoideae) reveal that the early branches are more highly diverse in floral morphology than are other clades of Papilionoideae. This study attempts for the first time to comprehensively sample the early-branching clades of this economically and ecologically important legume subfamily and thus to resolve relationships among them. • Methods: Parsimony and Bayesian phylogenetic analyses of the plastid matK and trnL intron sequences included 29 genera not yet sampled in matK phylogenies of the Papilionoideae, 11 of which were sampled for DNA sequence data for the first time. • Key results: The comprehensively sampled matK phylogeny better resolved the deep-branching relationships and increased support for many clades within Papilionoideae. The potentially earliest-branching papilionoid clade does not include any genus traditionally assigned to tribe Swartzieae. Dipterygeae is monophyletic with the inclusion of Monopteryx. The genera Aldina and Amphimas represent two of the nine main but as yet unresolved lineages comprising the large 50-kb inversion clade within papilionoids. The quinolizidine-alkaloid-accumulating genistoid clade is expanded to include a strongly supported subclade containing Ormosia and the previously unplaced Clathrotropis s.s., Panurea, and Spirotropis. Camoensia is the first-branching genus of the core genistoids. • Conclusions: The well-resolved phylogeny of the earliest-branching papilionoids generated in this study will greatly facilitate the efforts to redefine and stabilize the classification of this legume subfamily. Many key floral traits did not often predict phylogenetic relationships, so comparative studies on floral evolution and plant–animal interactions, for example, should also benefit from this study.     

The ethmoidal region of Dibamus taylori (Squamata: Dibamidae), with a phylogenetic hypothesis on dibamid relationships within Squamata

The nasal capsule and nasal sac of Dibamus taylori are described based on a plate reconstruction of serially sectioned material and are compared with several other squamate taxa. The ethmoidal region of D. taylori is characterized by a short nasal vestibule, a well developed concha, a much reduced and nearly functionless Jacobson's organ, an incomplete nasal roof, and a true nasopharyngeal duct with a complicated secondary palate. Cartilaginous and epithelial structures of the ethmoidal region provide many informative and useful characters for a phylogenetic analysis. Different anatomical features of D. taylori are compared with other burrowing forms ( Acontias meleagris, Annietta, pulchra Voetzkowia mira, Feylinia curron ) and discussed functionally and phylogenetically. Phylogenetic analysis of all higher squamate taxa using 144 available data derived from many regions of the body indicates relative relationships as (((Dibamidae, Amphisbaenia) Serpentes) Scleroglossa).     

The anostracan genus Branchinella (Crustacea: Branchiopoda), in need of a taxonomic revision; evidence from penile morphology

Three morphologically different penile types discovered in branchinellids from Botswana are compared with literature information on congeners. The striking differences among penile structures in Branchinella ondonguae, B. ornata and the halobiontic B. spinosa raise doubts about their congeneric status. Penis morphology in B. ondonguae corresponds with the most common configuration in branchinellids, but it is largely deviant in the other two. Branchinella ornata shares penile structures with Dendrocephalus , while basal parts in B. spinosa reflect affinity with chirocephalids. A considerable general morphological variability in Branchinella , not met in any other anostracan genus, may reflect the antiquity of the group, or else poor taxonomy. Large intra-branchinellid variability and vague generic boundaries, even when considering the usually conservative penile structures, stress the need of a thorough revision of the entire family of the Thamnocephalidae.     

Evolutionary morphology of whip spiders: towards a phylogenetic system (Chelicerata: Arachnida: Amblypygi)*

The aim of this study is to present a cladogram and phylogenetic system and to use this to discuss the phylogeny and biogeography of the Amblypygi. A total of 29 morphological structures were studied, their plesiomorphic and apomorphic characters or character states were identified, and the resulting data matrix was analysed. As a result, the ‘old’Charontidae or Pulvillata emerge as a paraphyletic group; the genus Paracharon is the sister group of all other amblypygids, which are now termed Euamblypygi. The ‘new’Charontidae (sensu Quintero: the genera Stygophrynus and Charon) are the sister group of the Phrynida or Apulvillata; together they form the Neoamblypygi. The relationships of the genera of the Charinidae cannot be resolved with the available data. They may be a paraphyletic group. The genus Catageus is a possible candidate for being the sister group of the Neoamblypygi. The new system allows a discussion of the phylogeny and biogeography of whip spiders. It also points to unresolved taxa and thus indicates the questions future research should address.