Peter Rehm

A transcriptome approach to ecdysozoan phylogeny.

The monophyly of Ecdysozoa, which comprise molting phyla, has received strong support from several lines of evidence. However, the internal relationships of Ecdysozoa are still contended. We generated expressed sequence tags from a priapulid (penis worm), a kinorhynch (mud dragon), a tardigrade (water bear) and five chelicerate taxa by 454 transcriptome sequencing. A multigene alignment was assembled from 63 taxa, which comprised after matrix optimization 24,249 amino acid positions with high data density (2.6% gaps, 19.1% missing data). Phylogenetic analyses employing various models support the monophyly of Ecdysozoa. A clade combining Priapulida and Kinorhyncha (i.e. Scalidophora) was recovered as the earliest branch among Ecdysozoa. We conclude that Cycloneuralia, a taxon erected to combine Priapulida, Kinorhyncha and Nematoda (and others), are paraphyletic. Rather Arthropoda (including Onychophora) are allied with Nematoda and Tardigrada. Within Arthropoda, we found strong support for most clades, including monophyletic Mandibulata and Pancrustacea. The phylogeny within the Euchelicerata remained largely unresolved. There is conflicting evidence on the position of tardigrades: While Bayesian and maximum likelihood analyses of only slowly evolving genes recovered Tardigrada as a sister group to Arthropoda, analyses of the full data set, and of subsets containing genes evolving at fast and intermediate rates identified a clade of Tardigrada and Nematoda. Notably, the latter topology is also supported by the analyses of indel patterns.

The diversity and evolution of chelicerate hemocyanins

BackgroundOxygen transport in the hemolymph of many arthropod species is facilitated by large copper-proteins referred to as hemocyanins. Arthropod hemocyanins are hexamers or oligomers of hexamers, which are characterized by a high O2 transport capacity and a high cooperativity, thereby enhancing O2 supply. Hemocyanin subunit sequences had been available from horseshoe crabs (Xiphosura) and various spiders (Araneae), but not from any other chelicerate taxon. To trace the evolution of hemocyanins and the emergence of the large hemocyanin oligomers, hemocyanin cDNA sequences were obtained from representatives of selected chelicerate classes.ResultsHemocyanin subunits from a sea spider, a scorpion, a whip scorpion and a whip spider were sequenced. Hemocyanin has been lost in Opiliones, Pseudoscorpiones, Solifugae and Acari, which may be explained by the evolution of trachea (i.e., taxon Apulmonata). Bayesian phylogenetic analysis was used to reconstruct the evolution of hemocyanin subunits and a relaxed molecular clock approach was applied to date the major events. While the sea spider has a simple hexameric hemocyanin, four distinct subunit types evolved before Xiphosura and Arachnida diverged around 470 Ma ago, suggesting the existence of a 4 × 6mer at that time. Subsequently, independent gene duplication events gave rise to the other distinct subunits in each of the 8 × 6mer hemocyanin of Xiphosura and the 4 × 6mer of Arachnida. The hemocyanin sequences were used to infer the evolutionary history of chelicerates. The phylogenetic trees support a basal position of Pycnogonida, a sister group relationship of Xiphosura and Arachnida, and a sister group relationship of the whip scorpions and the whip spiders.ConclusionFormation of a complex hemocyanin oligomer commenced early in the evolution of euchelicerates. A 4 × 6mer hemocyanin consisting of seven subunit types is conserved in most arachnids since more than 400 Ma, although some entelegyne spiders display selective subunit loss and independent oligomerization. Hemocyanins also turned out to be a good marker to trace chelicerate evolution, which is, however, limited by the loss of hemocyanin in some taxa. The molecular clock calculations were in excellent agreement with the fossil record, also demonstrating the applicability of hemocyanins for such approach.

Phylogenetic position of Myriapoda revealed by 454 transcriptome sequencing.

Myriapods had been considered closely allied to hexapods (insects and relatives). However, analyses of molecular sequence data have consistently placed Myriapoda either as a sister group of Pancrustacea, comprising crustaceans and hexapods, and thereby supporting the monophyly of Mandibulata, or retrieved Myriapoda as a sister group of Chelicerata (spiders, ticks, mites and allies). In addition, the relationships among the four myriapod groups (Pauropoda, Symphyla, Diplopoda, Chilopoda) are unclear. To resolve the phylogeny of myriapods and their relationship to other main arthropod groups, we collected transcriptome data from the symphylan Symphylella vulgaris, the centipedes Lithobius forficatus and Scolopendra dehaani, and the millipedes Polyxenus lagurus, Glomeris pustulata and Polydesmus angustus by 454 sequencing. We concatenated a multiple sequence alignment that contained 1550 orthologous single copy genes (1,109,847 amino acid positions) from 55 euarthropod and 14 outgroup taxa. The final selected alignment included 181 genes and 37,425 amino acid positions from 55 taxa, with eight myriapods and 33 other euarthropods. Bayesian analyses robustly recovered monophyletic Mandibulata, Pancrustacea and Myriapoda. Most analyses support a sister group relationship of Symphyla in respect to a clade comprising Chilopoda and Diplopoda. Inclusion of additional sequence data from nine myriapod species resulted in an alignment with poor data density, but broader taxon average. With this dataset we inferred Diplopoda+Pauropoda as closest relatives (i.e., Dignatha) and recovered monophyletic Helminthomorpha. Molecular clock calculations suggest an early Cambrian emergence of Myriapoda ∼513 million years ago and a late Cambrian divergence of myriapod classes. This implies a marine origin of the myriapods and independent terrestrialization events during myriapod evolution.

Macrofaunal communities on the continental shelf off Victoria Land (Ross Sea, Antarctica)

Abstract In austral summer 2004 benthic macrofauna was sampled along a latitudinal gradient along the northern Victoria Land coast (Ross Sea). An Agassiz trawl was used for semi-quantitative data collection of macrozoobenthos at depths from 84 to 537 m. Multivariate analysis of abundance of higher taxonomic units discriminated between the four sample sites along the latitudinal gradient. A SIMPROF analysis emphasized these geographical clusters, as the samples showed no significant differences within each cluster. A change in community structure with depth was not observed. The dominant taxonomic groups along the Victoria Land coast were Echinodermata (39%), Arthropoda (24%), Polychaeta (14%), and Mollusca (12%), not accounting for colonial organisms. Thus, the overall structure of the benthic community off the Victoria Land coast is comparable to other Antarctic regions and shows a closer relationship to the eastern Weddell Sea shelf, which may be attributable to the extensive impact of grounded ice affecting both areas.

Description of a new subspecies Diastylis enigmatica rossensis (Crustacea: Peracarida: Cumacea) from the Ross Sea, Antarctica

Diastylis enigmaticarossensis n. ssp. is described and illustrated on the basis of new specimens from the Antarctic shelf in the Ross Sea. The material was collected off the Victoria Land coast during the 19th Antarctic expedition of the Italian research vessel Italica in 2004. Diastylis enigmatica Ledoyer, 1993 was first obtained during the EPOS 3 campaign at Halley Bay in the Weddell Sea. A redescription based on further material from the Weddell Sea was published by Petrescu and Wittmann (Zool Med Leiden 77:557–630, 2003), which shows several differences to the newly collected material from the Ross Sea. The most obvious differences from Diastylis enigmaticarossensis n. ssp. to Diastylis enigmaticaenigmatica Ledoyer, 2003 are the absence of four spines on the merus of the second pereiopod, a lower number of setae on the uropodal endopod, a gap between spines on the peduncle article 1 of antenna 1, and a higher number of articles in both flagella of antenna 1.