Phylogenetic hypotheses for the peracarid order Cumacea are scarce and have not provided a solution to the full extent. In the present study, a fragment of the mitochondrial 16S rDNA was used to erect a phylogenetic hypothesis for three cumacean families, Diastylidae, Bodotriidae and Leuconidae, along with intra-family relationships of the latter. The Cumacea resolved monophyletic with tanaids and isopods as outgroup taxa. The Diastylidae were the only family with good support for monophyly. The genus Leucon resolved paraphyletic, whereas the subgenus Crymoleucon was monophyletic. Furthermore, the genetic structure was analysed for two leuconid species, Leucon antarcticus Zimmer, 1907 and L. intermedius Mühlenhardt-Siegel, 1996, from the Weddell Sea and the Ross Sea. The two species showed different patterns of intraspecific genetic variability. In contrast to L. intermedius, a bimodal distribution of pairwise genetic distances was observed for L. antarcticus, which is correlated with geographical and depth distributions between the Ross Sea and the Weddell Sea. Although a clear evaluation of cryptic speciation in these species requires additional work on more specimens from more geographic regions and broader depth ranges, differences shown in the sequences of 16S rDNA can only be explained by genetic separation of populations between the Weddell Sea and the Ross Sea for an extended period of time.

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.

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.

BACKGROUND:Oxygen 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.RESULTS:Hemocyanin 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.CONCLUSION:Formation 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.

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.

Molecular sequences do not only allow the reconstruction of phylogenetic relationships among species, but also provide information on the approximate divergence times. Whereas the fossil record dates the origin of most multicellular animal phyla during the Cambrian explosion less than 540 million years ago (mya), molecular clock calculations usually suggest much older dates. Here we used a large multiple sequence alignment derived from Expressed Sequence Tags and genomes comprising 129 genes (37,476 amino acid positions) and 117 taxa, including 101 arthropods. We obtained consistent divergence time estimates applying relaxed Bayesian clock models with different priors and multiple calibration points. While the influence of substitution rates, missing data, and model priors were negligible, the clock model had significant effect. A log–normal autocorrelated model was selected on basis of cross-validation. We calculated that arthropods emerged ∼600 mya. Onychophorans (velvet worms) and euarthropods split ∼590 mya, Pancrustacea and Myriochelata ∼560 mya, Myriapoda and Chelicerata ∼555 mya, and 'Crustacea' and Hexapoda ∼510 mya. Endopterygote insects appeared ∼390 mya. These dates are considerably younger than most previous molecular clock estimates and in better agreement with the fossil record. Nevertheless, a Precambrian origin of arthropods and other metazoan phyla is still supported. Our results also demonstrate the applicability of large datasets of random nuclear sequences for approximating the timing of multicellular animal evolution.

During the last years, molecular studies revealed significant population differentiation and cryptic species within various benthic and pelagic marine Antarctic taxa. This is unexpected due to the lack of obvious barriers to gene flow and strong current systems. Using mitochondrial (COI, 16S rDNA) and nuclear (28S rDNA: D2) gene fragments, we tested whether two circum-Antarctic benthic shrimps with planktotrophic larvae, Chorismus antarcticus and Nematocarcinus lanceopes, show patterns of regional differentiation. For both species, the 16S and the 28S fragment were invariant. However, for COI we found 24 different haplotypes for Chorismus antarcticus and 54 for Nematocarcinus lanceopes. No significant differentiation was observed among populations or regions. Furthermore, we found signatures of a population expansion in the late Pleistocene hinting at an impact of large-scale glaciations in particular on the shallow-water shrimp Chorismus antarcticus, supporting a (re)colonization and demographic expansion of this shrimp species in response to climate oscillation.

Diastylis enigmatica rossensis 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 enigmatica rossensis n. ssp. to Diastylis enigmatica enigmatica 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.

A new leuconid cumacean, Leucon (Crymoleucon) rossi n. sp., is described from depths of 84 to 458 m in the Ross Sea off the Victoria Land coast.Leucon rossi n. sp. is distinguished from other members of the genus by a combination of characters including 1) a blunt, horizontally directed pseudorostrum without a distinctly protruding siphon; 2) strongly developed antennal notch; 3) entire dorsomedian margin of carapace appearing serrate, armed with 14 to 19 anteriorly curved spines in female (up to 21 in subadult males); 4) a small, but distinct, spine behind the frontal lobe; and 5) the uropodal peduncle slightly shorter than the exopod.After Leucon antarcticus Zimmer, 1907, L. rossi was the second most frequently occurring cumacean in the samples collected off Victoria Land.Statistical analyses showed significant differences in the proportion of carapace length and height of adult (all incubating in the present study) and immature females compared to immature males; no adult males were available for study.

Macrozoobenthic community structure was studied in two surveys along a transect of 13 stations following the submersed Pleistocene Elbe valley in the south-eastern North Sea during May to June 2000 and March 2001. Two replicates of bottom samples were taken with a van Veen grab of 0.1 m 2 sampling size. In order to analyse the benthic macrofauna communities, the animals obtained were identified and counted, and MDS and cluster analysis were performed. Out of 200 taxa identified, 84 were polychaetes, 46 molluscs, 40 crustaceans, 15 echinoderms, and 15 belonged to other groups. Mean abundance was 4,860 individuals per m 2 , mean biomass 32.9 g ash free dry mass per m 2 . Mean diversity was 1.76 and mean evenness 0.54. The macrofauna of the Pleistocene Elbe valley is composed of three associations according to the cluster analysis. Each association is described by a combination of characterizing and discriminating species. An Amphiura-brachiata ― Tellimya-ferruginosa -association was found in the south-eastern part of the depression, whereas a transitional association with elements of both assemblages lead to an Amphiura - filiformis ― Galathowenia - oculata ― Nuculoma - tenuis -association in the north-western part of the valley. In the context of the entire southern North Sea, both associations are small-scale substructures, and as such are contained in the Nucula - nitidosa -community and the Amphiura - filiformis -community, respectively. A north-westward shift of the community of the southern Elbe valley was found and discussed as a possible consequence of warm winters of the last decades.

The following study was the first to describe composition and structure of the peracarid fauna systematically along a latitudinal transect off Victoria Land (Ross Sea, Antarctica). During the 19th Antarctic expedition of the Italian research vessel "Italica" in February 2004, macrobenthic samples were collected by means of a Rauschert dredge with a mesh size of 500 μm at depths between 85 and 515 m. The composition of peracarid crustaceans, especially Cumacea was investigated. Peracarida contributed 63% to the total abundance of the fauna. The peracarid samples were dominated by amphipods (66%), whereas cumaceans were represented with 7%. Previously, only 13 cumacean species were known, now the number of species recorded from the Ross Sea increased to 34. Thus, the cumacean fauna of the Ross Sea, which was regarded as the poorest in terms of species richness, has to be considered as equivalent to that of other high Antarctic areas. Most important cumacean families concerning abundance and species richness were Leuconidae, Nannastacidae, and Diastylidae. Cumacean diversity was lowest at the northernmost area (Cape Adare). At the area off Coulman Island, which is characterized by muddy sediment, diversity was highest. Diversity and species number were higher at the deeper stations and abundance increased with latitude. A review of the bathymetric distribution of the Cumacea from the Ross Sea reveals that most species distribute across the Antarctic continental shelf and slope. So far, only few deep-sea records justify the assumption of a shallow-water–deep-sea relationship in some species of Ross Sea Cumacea, which is discussed from an evolutionary point of view.

The Victoria-Land Transect project onboard the Italian research vessel "Italica" in February 2004, was a large-scale attempt to obtain benthic samples of smaller macrozoobenthic specimens systematically along a latitudinal and a depth transect along the Victoria-Land coast. Data presented from this survey are based on Rauschert dredge samples, which were taken at four areas at depth ranging from 84 to 515 m. A cluster analysis based on relative numbers of abundance was performed and demonstrated a change in community structure depending on the location along the latitudinal transect. A change in community structure with depth was not recorded. Dominant taxa of the Ross Sea fauna along the Victoria-Land coast were the Arthropoda (65.7%), followed by Annelida (20.7%), Mollusca (9.6%) and Echinodermata (2.5%). Total number of abundance decreased with depth with an exception at Cape Russell, whereas a trend in biomass was not documented. Abundance and biomass proportions of major taxa changed gradually along the latitudinal transect.

Until now the genus Abyssianira Menzies, 1956 included two Atlantic species from abyssal depths (A. dentifrons Menzies, 1956; A. argentenensis Menzies, 1962) and two Australian species from upper to mid-bathyal depths (A. bathyalis Just, 1990; A. tasmaniensis Just, 1990). Except for Abyssianira dentifrons, which has been found in both hemispheres, the remaining ones are only known from the southern hemisphere. In the present contribution the Atlantic species of Abyssianira are redescribed, and two new species are described: A. acutilobi n. sp. from off Argentina and A. lingula n. sp. from off Brazil, raising the total number of species known for this genus to six.

The Victoria Land Transect Cruise with the Italian research vessel Italica, carried out in February 2004, was the first large-scale attempt to obtain benthic samples systematically along a latitudinal transect along the Antarctic Ross Sea shelf. Along the transect, samples were taken in five areas (Cape Russell, Terra Nova Bay, Coulman Island, Cape Hallet and Cape Adare) along a depth gradient ranging from 80m to 515m. Sampling was carried out using a Rauschert dredge with a mesh size of 500 µm, as the main emphasis was on smaller peracarid crustaceans. Distribution of higher taxonomic groups based on total abundance was found as follows: crustaceans (71-83%), polychaetes (12-16%), molluscs (3-7%), echinoderms (0-2%). Exceptions were the areas Cape Adare with an increased occurrence of molluscs (38%) and Coulmann Island with a mass occurrence of polychaetes (63%). The crustaceans were dominated by amphipods (32%-66%), isopods (13-30%), tanaids (4-28%), and cumaceans (<1-35%). Especially in the southernmost areas Terra Nova Bay and Cape Russel, an increase in abundance of cumaceans with increasing latitude was found. Within the cumacean fauna of the Ross Sea, the most abundant families were Leuconidae (8%-85%), Nanastacidae (10-92%) and Diastylidae (3-16%). The systematic collection of smaller crustaceans yielded a high amount of species new to the area under investigation, especially for the Peracarida. A more detailed faunal analysis of the crustaceans is presently under way.