The great variety of geological and hydrological conditions in the deep sea generates many different habitats. Some are only recently explored, although their true extent and geographical coverage are still not fully established. Both continental margins and mid-oceanic seafloors are much more complex ecologically, geologically, chemically and hydrodynamically than originally thought. As a result, fundamental patterns of species distribution first observed and explained in the context of relatively monotonous slopes and abyssal plains must now be re-evaluated in the light of this newly recognized habitat heterogeneity. Based on a global database of nematode genus composition, collected as part of the Census of Marine Life, we show that macrohabitat heterogeneity contributes significantly to total deep-sea nematode diversity on a global scale. Different deep-sea settings harbour specific nematode assemblages. Some of them, like coral rubble zones or nodule areas, are very diverse habitats. Factors such as increased substrate complexity in the case of nodules and corals seem to facilitate the co-existence of a large number of genera with different modes of life, ranging from sediment dwelling to epifaunal. Furthermore, strong biochemical gradients in the case of vents or seeps are responsible for the success of particular genera, which are not prominent in more typical soft sediments. Many

Correct taxonomy is a prerequisite for biological research, but currently it is undergoing a serious crisis, resulting in the neglect of many highly diverse groups of organisms. In nematodes, species delimitation remains problematic due to their high morphological plasticity. Evolutionary approaches using DNA sequences can potentially overcome the problems caused by morphology, but they are also affected by theoretical flaws. A holistic approach with a combination of morphological and molecular methods can therefore produce a straightforward delimitation of species. The present study investigates the taxonomic status of some highly divergent mitochondrial haplotypes in the Rhabditis (Pellioditis) marina species complex by using a combination of molecular and morphological tools. We used three molecular markers (COI, ITS, D2D3) and performed phylogenetic analyses. Subsequently, morphometric data from nearly all lineages were analysed with multivariate techniques. We included R. (P.) mediterranea and R. (R.) nidrosiensis to infer species status of the observed lineages. The results showed that highly divergent genotypic clusters were accompanied by morphological differences, and we created a graphical polytomous key for future identifications. This study indisputably demonstrates that R. (P.) marina and R. (P.) mediterranea belong to a huge species complex and that biodiversity in free-living marine nematodes may be seriously underestimated.

Integrative taxonomy considers species boundaries from multiple, complementary perspectives, with the main objective being to compare the observed data against the predictions of the methodologies used. In the present study we used three methods for delineating species boundaries within the cosmopolitan nematode species Rhabditis (Pellioditis) marina and Halomonhystera disjuncta. First, phylogenetic relationships among molecular sequences from the mitochondrial cytochrome oxidase c subunit 1 gene (COI), and from two nuclear regions, internal transcribed spacer (ITS) and D2D3, were analysed. Subsequently, multivariate morphometric analysis was used to investigate whether concordant molecular lineages were also morphologically distinct. When morphological differences were found, typological taxonomy was performed to identify fixed or non-overlapping characters between lineages. Interbreeding experiments were conducted between the two closest related lineages of R. (P.) marina to investigate potential reproductive isolation. This integrative approach confirmed the presence of several species within each nominal species: molecular lineages were concordant across two independent loci (COI and ITS), and were characterized by significant morphological divergence. Most lineages were also detectable in the D2D3 region, but were less resolved. The two lineages investigated in our study did not produce offspring. Our results highlight that classical taxonomy grossly underestimates species diversity within the phylum Nematoda.

An on-board experiment was performed during a research cruise to investigate the ability of deep-sea nematode species to actively colonise defaunated sediments. Small cylinders of 500 µm wire mesh filled with defaunated sediment were inserted into microcosms containing sediment with indigenous meiofauna collected from 1300 m depth in the Arctic Ocean. The defaunated sediments were either enriched with the diatom Thalassiosira weissflogii, or remained unenriched. Samples from the defaunated sediment were taken after 9 and 17 d. As controls, microcosms with sediment containing the indigenous meiofauna but without an internal cylinder were also sampled at each time interval. Nematodes colonised both enriched and unenriched sediments with abundances of up to 20% of the controls. Irrespective of the time of sampling, abundance and number of species were significantly higher in the enriched treatment, suggesting that the presence of food enhances colonisation and resilience. Nematode assemblages in the defaunated sediments were species-rich and differed from the controls. The majority of colonising species were rare or undetectable in the controls, suggesting that episodic disturbances may be necessary for their persistence in deep sea sediments. Colonisation was in part determined by species characteristics such as size and motility. At the same time, a large number of different species colonised the empty cylinders, resulting in poor similarity in community composition between replicates, particularly between samples enriched with diatoms. Our results indicate a poor predictability of community composition of recently disturbed sediments despite highly reproducible abundance and diversity patterns and lends experimental support to the existence of a spatio-temporal mosaic that emerges from highly localised colonisation patterns.