Thomas Bartolomaeus

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.

New insights into polychaete phylogeny (Annelida) inferred from 18S rDNA sequences

Annelid systematics and the ingroup relationships of polychaete annelids are matter of ongoing debates in recent analyses. For the investigation of sedentary polychaete relationships a molecular phylogenetic analysis was conducted based on 94 sequences of 18S rDNA, including unpublished sequences of 13 polychaete species. The data set was analyzed with maximum parsimony and maximum likelihood methods, as wells as Bayesian inference. As in previous molecular analyses the monophyly of many traditional polychaete families is confirmed. No evidence has been found for a possible monophyly of Canalipalpata or Scolecida. In all analyses a placement of the Echiura as a derived polychaete ingroup with a close relationship to the Capitellidae is confirmed. The orbiniids appear paraphyletic with regard to Questa. Travisia is transferred from Opheliidae to Scalibregmatidae. The remaining opheliids include a yet undescribed ctenodrilid species from Elba, whereas the other investigated ctenodrilid Ctenodrilus serratus groups with the Cirratulidae and shows a close affinity to the cirratulid genus Dodecaceria. A common ancestry of Branchiomaldane and Arenicola, which has been predicted on morphological data, is confirmed by the analysis and a sistergroup relationship between Arenicolidae and Maldanidae is also recovered. These results support our assumption that on the basis of a broader taxon sampling the phylogenetic position of controversially discussed taxa can be inferred by using 18S rDNA sequence data.

Structure and formation of the uncini in Pectinaria koreni, Pectinaria auricoma (Terebellida) and Spirorbis spirorbis (Sabellida): inplications for annelid phylogeny and the position of the Pogonophora

Setation is an important taxonomic character of the Annelida. Within this taxon, Terebellida and Sabellida both have transverse rows of short, apically toothed setae which are situated inside the neuropodial rim. The apical spines are curved and their tips point anteriorly. These setae are termed uncini. In the terebellidans Pectinaria koreni, Pectinaria auricoma and in the sabellidan Spirorbis spirorbis, these setae arise from a follicle which consists of a chaetoblast and two follicle cells. The special structure of the uncini is a result of temporal modifications of the actin-filament system of the chaetoblast and changing spatial interactions between the chaetoblast and the follicle cells during the formation of these setae. Once the uncinus is formed, the microvilli are withdrawn and electron-dense material is deposited in the remaining canals. The microvilli are replaced by short processes of the chaetoblast, and the actin-filament system is replaced by a system of intermediate filaments which help to mechanically attach the uncinus to the follicle. Such uncini are also described in both pogonophoran groups, the Perviata and the Obturata (Vestimentifera). In several structural details they correspond to those of the species investigated in this paper, so that the hypothesis of a homology of the uncini seems to be justified. This hypothesis leads to the conclusion that uncini evolved in the common stem lineage of Pogonophora, Terebellida and Sabellida, implying a monophyletic origin of these three taxa. The uncini are compared to the hooked setae of the Arenicolida, Maldanida and Psammodrilida, which are also aligned in transverse rows inside the neurophodial rim. Hooked setae and uncini are hypothesized to be homologous. It, therefore, can be concluded that Arenicolida, Maldanida and Psammodrilida are closely related to the monophylum consisting of Terebellida, Sabellida and Pogonophora, and that these six taxa share a common ancestor, which evolved transverse rows of setae with apically curved spines and a formative site lateral to the edge of the neuropodial rim. According to the phylogenetic relationships proposed here, the Pogonophora are a subordinate taxon within the Annelida.

The linguistic problem of morphology: structure versus homology and the standardization of morphological data

The present article discusses the need for standardization in morphology in order to increase comparability and communicability of morphological data. We analyse why only morphological descriptions and not character matrices represent morphological data and why morphological terminology must be free of homology assumptions. We discuss why images only support and substantiate data but are not data themselves. By comparing morphological traits and DNA sequence data we reveal fundamental conceptual shortcomings of the former that result from their high average degree of individuality. We argue that the delimitation of morphological units, of datum units, and of evidence units must be distinguished, each of which involves its own specific problems. We conclude that morphology suffers from the linguistic problem of morphology that results from the lack of (i) a commonly accepted standardized morphological terminology, (ii) a commonly accepted standardized and formalized method of description, and (iii) a rationale for the delimitation of morphological traits. Although this is not problematic for standardizing metadata, it hinders standardizing morphological data. We provide the foundation for a solution to the linguistic problem of morphology, which is based on a morphological structure concept. We argue that this structure concept can be represented with knowledge representation languages such as the resource description framework (RDF) and that it can be applied for morphological descriptions. We conclude with a discussion of how online databases can improve morphological data documentation and how a controlled and formalized morphological vocabulary, i.e. a morphological RDF ontology, if it is based on a structure concept, can provide a possible solution to the linguistic problem of morphology.

Polychaete phylogeny based on morphological data – a comparison of current attempts

Annelid phylogeny is one of the largest unresolved problems within the Metazoa. This is due to the enormous age of this taxon and also strongly influenced by the current discussion on the position of the Arthropoda, which traditionally is hypothesized to be the annelid sister taxon. Within the framework of recent discussions on the position of the Annelida, the ground pattern of this taxon is either a clitellate-like, parapodia-less dwelling organism or an organisms that resembles errant polychaetes in having parapodia and gills and probably being a predator. To solve this problem different attempts have been made in the past, cladistic analysis, scenario based plausibility considerations and a successive search for sister taxa base on isolated characters. These attempts are presented and critically discussed. There is at least strong support for the Annelida as wells as for several of its taxa above the level of traditional families; the monophyly of the Polychaeta, however, remains questionable.

Mitochondrial genome sequence and gene order of Sipunculus nudus give additional support for an inclusion of Sipuncula into Annelida

BackgroundMitochondrial genomes are a valuable source of data for analysing phylogenetic relationships. Besides sequence information, mitochondrial gene order may add phylogenetically useful information, too. Sipuncula are unsegmented marine worms, traditionally placed in their own phylum. Recent molecular and morphological findings suggest a close affinity to the segmented Annelida.ResultsThe first complete mitochondrial genome of a member of Sipuncula, Sipunculus nudus, is presented. All 37 genes characteristic for metazoan mtDNA were detected and are encoded on the same strand. The mitochondrial gene order (protein-coding and ribosomal RNA genes) resembles that of annelids, but shows several derivations so far found only in Sipuncula. Sequence based phylogenetic analysis of mitochondrial protein-coding genes results in significant bootstrap support for Annelida sensu lato, combining Annelida together with Sipuncula, Echiura, Pogonophora and Myzostomida.ConclusionThe mitochondrial sequence data support a close relationship of Annelida and Sipuncula. Also the most parsimonious explanation of changes in gene order favours a derivation from the annelid gene order. These results complement findings from recent phylogenetic analyses of nuclear encoded genes as well as a report of a segmental neural patterning in Sipuncula.

Systematic comparison and reconstruction of sea urchin (Echinoidea) internal anatomy: a novel approach using magnetic resonance imaging

BackgroundTraditional comparative morphological analyses and subsequent three-dimensional reconstructions suffer from a number of drawbacks. This is particularly evident in the case of soft tissue studies that are technically demanding, time-consuming, and often prone to produce artefacts. These problems can partly be overcome by employing non-invasive, destruction-free imaging techniques, in particular micro-computed tomography or magnetic resonance imaging.ResultsHere, we employed high-field magnetic resonance imaging techniques to gather numerous data from members of a major marine invertebrate taxon, the sea urchins (Echinoidea). For this model study, 13 of the 14 currently recognized high-ranking subtaxa (orders) of this group of animals were analyzed. Based on the acquired datasets, interactive three-dimensional models were assembled. Our analyses reveal that selected soft tissue characters can even be used for phylogenetic inferences in sea urchins, as exemplified by differences in the size and shape of the gastric caecum found in the Irregularia.ConclusionThe main focus of our investigation was to explore the possibility to systematically visualize the internal anatomy of echinoids obtained from various museum collections. We show that, in contrast to classical preparative procedures, magnetic resonance imaging can give rapid, destruction-free access to morphological data from numerous specimens, thus extending the range of techniques available for comparative studies of invertebrate morphology.

Phylogeny and mitochondrial gene order variation in Lophotrochozoa in the light of new mitogenomic data from Nemertea

BackgroundThe new animal phylogeny established several taxa which were not identified by morphological analyses, most prominently the Ecdysozoa (arthropods, roundworms, priapulids and others) and Lophotrochozoa (molluscs, annelids, brachiopods and others). Lophotrochozoan interrelationships are under discussion, e.g. regarding the position of Nemertea (ribbon worms), which were discussed to be sister group to e.g. Mollusca, Brachiozoa or Platyhelminthes. Mitochondrial genomes contributed well with sequence data and gene order characters to the deep metazoan phylogeny debate.ResultsIn this study we present the first complete mitochondrial genome record for a member of the Nemertea, Lineus viridis. Except two trnP and trnT, all genes are located on the same strand. While gene order is most similar to that of the brachiopod Terebratulina retusa, sequence based analyses of mitochondrial genes place nemerteans close to molluscs, phoronids and entoprocts without clear preference for one of these taxa as sister group.ConclusionAlmost all recent analyses with large datasets show good support for a taxon comprising Annelida, Mollusca, Brachiopoda, Phoronida and Nemertea. But the relationships among these taxa vary between different studies. The analysis of gene order differences gives evidence for a multiple independent occurrence of a large inversion in the mitochondrial genome of Lophotrochozoa and a re-inversion of the same part in gastropods. We hypothesize that some regions of the genome have a higher chance for intramolecular recombination than others and gene order data have to be analysed carefully to detect convergent rearrangement events.

The phylogenetic position of Brachiopoda—a comparison of morphological and molecular data

Analyses of rRNA and rDNA among Metazoa result in a hypothesis of a sistergroup relationship of Brachiopoda and certain spiralian taxa, whereas analyses of morphological data imply that Brachiopoda show affinities to Deuterostomia within the Radialia. Regarding Brachiopoda as a derived spiralian taxon must be followed by a reinterpretation of the evolution of distinct brachiopod morphological characters—like cleavage pattern, coelom or larva. The experimental insertion of a monophyletic taxon consisting of Brachiopoda and Phoronida into a widely accepted phylogenetic tree of Spiralia leads to the hypothesis that at least trimeric organization, mesosomal tentacular apparatus and heterogeneously assembled metanephridia are products of convergent evolution in Brachiopoda plus Phoronida and Deuterostomia. The hypothesis of a radialian nature of Brachiopoda and Phoronida, as implied by morphological data, remains as the most parsimonious possibility to explain the evolution of seven regarded characters (cleavage pattern, larva, tentacular apparatus, coelom, metameric segmentation, metanephridia and chaetae) in Brachiopoda. Due to the conflicting results of both methods a hitherto undetected systematical problem is discussed possibly hindering data comparability. If the course of evolution can principally be inferred from the information preserved in recent and fossil animals, the results should be congruent in the analyses of both, molecular and morphological data.

Structure and development of nephridia in Annelida and related taxa

Two different kinds of filtration nephridia, protonephridia and metanephridia, are described in Polychaeta. During ontogenesis protonephridia generally precede metanephridia. While the latter are segmentally arranged, protonephridia are characteristic for the larva and are the first nephridial structure formed during ontogenesis. There is strong evidence that both organs depend on the same information and that their specific structure depends on the way in which the coelom is formed and which final expansion it gains. While metanephridia are regarded to be homologous throughout the polychaetes, protonephridia seem to have evolved in several lineages. Some of the protonephridia closely resemble less differentiated stages of metanephridial development, so that protonephridial evolution can be explained by truncation of the metanephridial development. Nevertheless, structural details are large enough to allow us to expect information on the polychaete evolution if the database on polychaete nephridia increases. A comparison of the polychaete metanephridia with those of the Clitellata and Sipuncula reveals some surprising details. In Clitellata the structure of the funnel is quite uniform in microdrilid oligochaetous Clitellata and resembles that of the aeolosomatids. Like the nephridia in the polychaete taxa Sabellida and Terebellida, those of the Sipunucla possess podocytes covering the coelomic side of the duct.