Jesper Guldberg Hansen

Molecular phylogeny of Arthrotardigrada (Tardigrada).

Tardigrades are microscopic ecdysozoans with a worldwide distribution covering marine, limnic and terrestrial habitats. They are regarded as a neglected phylum with regard to studies of their phylogeny. During the last decade molecular data have been included in the investigation of tardigrades. However, the marine arthrotardigrades are still poorly sampled due to their relative rarity, difficult identification and minute size even for tardigrades. In the present study, we have sampled various arthrotardigrades and sequenced the 18S and partial 28S ribosomal subunits. The phylogenetic analyses based on Bayesian inference and maximum parsimony inferred Heterotardigrada (Arthrotardigrada+Echiniscoidea) and Eutardigrada to be monophyletic. Arthrotardigrada was inferred to be paraphyletic as the monophyletic Echiniscoidea is included within the arthrotardigrades. The phylogenetic positions of Stygarctidae and Batillipedidae are poorly resolved with low branch support. The Halechiniscidae is inferred to be polyphyletic as the currently recognized Styraconyxinae is not part of the family. Archechiniscus is the sister-group to the Halechiniscidae and Orzeliscus is placed as one of the basal halechiniscids. The phylogeny of the included eutardigrade taxa resembles the current molecular phylogenies. The genetic diversity within Arthrotardigrada is much larger (18S 15.1-26.5%, 28S 7.2-20.7%) than within Eutardigrada (18S 1.0-12.6%, 28S 1.3-8.2%). This can be explained by higher substitution rates in the arthrotardigrades or by a much younger evolutionary age of the sampled eutardigrades.

A comparative morphological study of the kinorhynch genera Antygomonas and Semnoderes (Kinorhyncha: Cyclorhagida)

Detailed information revealed through combined use of light- and scanning electron microscopy, is given for two species of kinorhynchs, representing the cyclorhagid genera Semnoderes and Antygomonas. The two species have not previously been examined using SEM, and the new observations point out several similarities between species of the two genera, which could indicate a potential close relationship. The generated data is meant to be incorporated in a future phylogenetic analysis in order to clarify the phylogenetic relationships among kinorhynchs.

Preliminary Studies of the Tardigrade Fauna of the Faroe Bank

Abstract Four cruises to the Faroe Bank have collected bottom samples for qualitative analysis of the meiofauna. The preliminary results show a very rich tardigrade fauna, with a large proportion of species new to science. At present 35 species of tardigrades belonging to 4 families (6 sub-families) have been found, of these are 22 new to science (63%). The 35 species comprise more than 20% of all known marine tardigrades. Halechiniscidae is represented by 30 species (1 Euclavarctinae, 11 Styraconyxinae, 12 Tanarctinae, 3 Halechiniscinae, 2 Florarctinae and 1Dipodarctinae). This family comprises 87.87% of the specimens sorted out so far. Specimens from the subfamilies Tanarctinae (46.63%) and Styraconyxinae (31.54%) are dominating. Batillipedidae is represented with 3species (8.63% of the specimens) and Coronarctidae (1.89%) and Stygarctidae (1.61%) with a single species each. Samples with similar sediment from 104—260 m depth have similar species distribution in the families. This implicates that the sediment is the key factor involved in the species distribution and that depth is less important. The calcareous sediment is a unique substrate and the tardigrade fauna of the Faroe Bank can be compared with that of sub-tropical and tropical coralline sand. The composition of species exhibits a strong taxonomic affinity with the tardigrade fauna from more southern latitudes, i.e. the Mediterranean Sea and the south-eastern coast of USA.

Geonemy, ecology, reproductive biology and morphology of the tardigrade Hypsibius zetlandicus (Eutardigrada: Hypsibiidae) with erection of Borealibius gen. n.

The morphology, biology and geographic distribution of Hypsibius zetlandicus (Murray 1907) are considered. Scanning electron microscopy (SEM) and/or light microscopy (LM) analyses have been carried out on H. zetlandicus and the type species of Hypsibius: Hypsibius dujardini (Doyère 1840), with particular emphasis on the buccal–pharyngeal apparatuses. Some unusual characteristics of this apparatus in H. zetlandicus lead us to the erection of the new genus Borealibius, to which H. zetlandicus (Borealibius zetlandicus comb. n.) is transferred. In the light of new discoveries of this species from polar, sub-polar and alpine regions, and based on the available bibliographic references, we hypothesize a boreo–alpine distribution for this species. The presence of traits that are unusual and rare (in other tardigrade species) have been observed whilst analyzing the reproductive biology and ecology of B. zetlandicus (i.e., the presence of hermaphroditism, parental care and the colonization of very different substrates).

Postembryonic development of Antygomonas incomitata (Kinorhyncha: Cyclorhagida)

Postembryonic development in the kinorhynch species Antygomonas incomitata was examined using scanning electron microscopy. The morphology of the six juvenile stages, J‐1 to J‐6, varies at numerous details, but they can also be distinguished by a few key characters. Juvenile stage 1 by its composition of only nine trunk segments; J‐2 by the combination of possessing 10 trunk segments, but no cuspidate spines on segment 9; J‐3 by the presence of cuspidate spines on segment 9, but only one pair of cuspidate spines on segment 8; J‐4 by the combination of 10 trunk segments only, but having two pairs of cuspidate spines on segment 8; J‐5 by possessing 11 trunk segments and same spine compositions as adults but is still maintaining postmarginal spiculae; J‐6 specimens closely resemble adults and are most easily identified by their reduced trunk lengths. New segments are formed in a growth zone in the anterior part of the terminal segment. The complete number of segments is reached in J‐5. Development of cuticular head and trunk structures are described through all postembryonic stages and following developmental patterns could be outlined: the mouth cone possesses outer oral styles from J‐1, but in J‐1 to J‐3, the styles alternate in size. Scalids of the introvert are added after each molt, and scalids appear earliest in the anterior rings, whereas scalids in more posterior rings are added in older postembryonic stages. The early J‐1 stage is poor in spines and sensory spots and both structures increase in number after each molt. The complete spine composition is reached in J‐4, whereas new sensory spots appear after all molts, inclusive the final one from J‐6 to adult. Sensory spots in the paraventral positions often appear as Type 3 sensory spots but are through development transformed to Type 2. This transformation happens earliest on the anterior segments. J. Morphol., 2010.

A new species of freshwater tardigrades from Disko Island (greenland) increases an unsolved paradox in tardigrade systematics

During the “Workshop on Arctic tardigrades” at the Danish Arctic Station (Qeqertarsuaq, Disko Island, Greenland) an undescribed species of Dactylobiotus was found in freshwater sediments of the Isunngua spring. We have the honour and pleasure to describe this new taxon that we dedicate to all participants of that symposium, naming the species Dactylobiotus octavi sp. n. The animals appear similar to Dactylobiotus dispar and Dactylobiotus haplonyx with the presence of a very short secondary branch in the claws of the first three pairs of legs, but they differ from these species in the size of claw and buccal tube width. This new species also has peculiar ornamented eggs. The eggshell consists of bowl-like processes, each one surrounded by a band of fine pores. A morphological parsimony analysis to identify phylogenetic relationships among D. octavi sp. n. and the other Dactylobiotus species was performed, obtaining inconclusive results. The discovery of this new species increases an unsolved paradox in tardigrade systematics related to the presence of closely related species which share a very similar morphology of the animals but clearly differ in their egg morphology, while, conversely, there are species belonging to different evolutionary lines that have similar eggs, but very different adult morphology. The finding of D. octavi sp. n. increases the already high number of species found in Disko Island and once again underlines the importance of tardigrades in the biodiversity of the Arctic area.

A Comparison of Morphological Characters Within the Genus Rhomboarctus (Tardigrada: Heterotardigrada) with the Description of Two New Species

Two new species of marine arthrotardigrades:Rhomboarctus aslaki sp. n. and Rhomboarctus duplicicaudatus sp. n. (Tardigrada: Halechiniscidae) belonging to the genus Rhomboarctus Renaud-Mornant, 1984 are described from the North Atlantic Sea and the Mediterranean Sea, respectively. New features for the formerly monotypic genus are presented and a complete description of the buccal apparatus is given for the first time.

A molecular approach to arthrotardigrade phylogeny (Heterotardigrada, Tardigrada)

The marine order Arthrotardigrada (class Heterotardigrada, phylum Tardigrada) is known for its conspicuously high morphological diversity and has been traditionally recognized as the most ancestral group within the phylum. Despite its potential importance in understanding the evolution of the phylum, the phylogenetic relationships of Arthrotardigrada have not been clarified. This study conducted molecular phylogenetic analyses of the order encompassing all families except Neoarctidae using nuclear 18S and 28S rRNA fragments. Data from two rare families, Coronarctidae and Renaudarctidae, were included for the first time. The analyses confirmed the monophyly of Heterotardigrada and inferred Coronarctidae as the sister group to all other heterotardigrade taxa. Furthermore, the results support a monophyletic Renaudarctidae + Stygarctidae clade, which has been previously suggested on morphology. Our data indicated that two subfamilies currently placed in Halechiniscidae are only distantly related to this family. We propose that these taxa are each elevated to family level (Styraconyxidae (new rank) and Tanarctidae (new rank)). The morphology of tardigrades is discussed in the context of the inferred phylogeny.