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Tardigrades form an important component of meiofaunal communities across the globe. However, our knowledge on tardigrade ecology is very limited. Here, we report the results of 21 field samplings of the marine tardigradeHalobiotus crispaecollected over a period of 74 months at the locality of Vellerup Vig, Denmark, with the aim of providing novel insights into its ecology. Uniquely,H. crispaeis characterized by the presence of seasonal cyclic changes in the phenotype of the animal, i.e. cyclomorphosis. Our sampling data include (i) total number of animals extracted, (ii) dominant cyclomorphic stage found, and (iii) important environmental parameters such as temperature, salinity and pH. Our accumulated data constitute a tentative model for the annual fluctuations in animal density, which reveals an annual peak in abundance during the months of February and March. In contrast, tardigrade density appears to decrease in response to increasing temperatures during late spring/early summer. The thermal tolerance ofH. crispaewas therefore investigated experimentally for the pseudosimplex 2 stage, active stage and pseudosimplex 1 stage, revealing LTminvalues of 24.6°C, 14.2°C and 24.7°C, LT50values of 30.6°C, 29.6°C and 30.8°C, and LTmaxvalues of 35°C, 35.8°C and 35°C, respectively. In general, our thermal tolerance data reveal that (i) the entry into the pseudosimplex 1 stage represents an adaptive strategy that enables the survival of the Vellerup Vig population during periods of heat stress, and (ii) temperature is an important factor in limiting the geographic distribution ofH. crispae. [ABSTRACT FROM PUBLISHER]

Many species of tardigrades are known to tolerate extreme environmental stress, yet detailed knowledge of the mechanisms underlying the remarkable adaptations of tardigrades is still lacking, as are answers to many questions regarding their basic biology. Here, we present data on the inorganic ion composition and total osmotic concentration of five different species of tardigrades (Echiniscus testudo, Milnesium tardigradum, Richtersius coronifer, Macrobiotus cf. hufelandi and Halobiotus crispae) using high-performance liquid chromatography and nanoliter osmometry. Quantification of the ionic content indicates that Na+ and Cl- are the principal inorganic ions in tardigrade fluids, albeit other ions, i.e. K+, NH4+, Ca2+, Mg2+, F-, SO42- and PO43- were also detected. In limno-terrestrial tardigrades, the respective ions are concentrated by a large factor compared with that of the external medium (Na+, ×70-800; K+, ×20-90; Ca2+ and Mg2+, ×30-200; F-, ×160-1040, Cl-, ×20-50; PO43-, ×700-2800; SO42-, ×30-150). In contrast, in the marine species H. crispae, Na+, Cl- and SO42- are almost in ionic equilibrium with (brackish) salt water, while K+, Ca2+, Mg2+ and F- are only slightly concentrated (×2-10). An anion deficit of ~120 mEq l-1 in M. tardigradum and H. crispae indicates the presence of unidentified ionic components in these species. Body fluid osmolality ranges from 361±49 mOsm kg-1 in R. coronifer to 961±43 mOsm kg-1 in H. crispae. Concentrations of most inorganic ions are largely identical between active and dehydrated groups of R. coronifer, suggesting that this tardigrade does not lose large quantities of inorganic ions during dehydration. The large osmotic and ionic gradients maintained by both limno-terrestrial and marine species are indicative of a powerful ion-retentive mechanism in Tardigrada. Moreover, our data indicate that cryptobiotic tardigrades contain a large fraction of unidentified organic osmolytes, the identification of which is expected to provide increased insight into the phenomenon of cryptobiosis. [ABSTRACT FROM AUTHOR]

Aim To analyse the importance of climatic and geographical isolation in determining the patterns of speciation and distribution of species within the tardigrade genus Echiniscoides. Location Marine intertidal zone, globally. Methods DNA was extracted from 465 individual tardigrades from 48 localities world-wide. The tardigrades were divided into clusters using several distance-based criteria. The phylogeny of these clusters was estimated with Bayesian analyses. The relationships between genetic distance and substrate, climate, and geographical distance were tested with a new improved Mantel test which incorporates phylogenetic uncertainties by analysing the raw tree data instead of the averaged tree. Results Approximately 40 clusters, each probably corresponding to species, were recovered from the genetic analysis; the number of clusters fluctuated depending on the criterion used for cluster delimitation. Each cluster had a limited temperature range and all clusters were confined to single oceans under all realistic criteria for cluster delimitation. Apart from a tropical cluster, each cluster occurred only in one hemisphere. Occurrence on different substrata was not correlated with genetic distance between clusters. Both geographical distance and climate were correlated with genetic distance; however, the correlation between geographical and genetic distance disappeared when the non-independence of climatic and geographical distance were controlled for. Main conclusions The distribution of individual species of Echiniscoides is limited by climate and geographical distance. Distance does not appear to be a major factor influencing phylogeny in this genus, but ecological speciation along a temperature gradient appears to be important. [ABSTRACT FROM AUTHOR]

Intestinal schistosomiasis is a global disease of enormous public health importance. Assessing the local risk of transmission of the parasite causing the disease requires an appraisal of the distribution of its intermediate snail hosts, Biomphalaria spp.. In East Africa, the Lake Victorian Basin is a major freshwater ecosystem and highly endemic for intestinal schistosomiasis, although detailed distribution data for Biomphalaria have not been collected. We used on-the-ground malacological surveys and conjoint measurement of environmental determinants to develop models for analysis of variables associated with distribution and predictive snail mapping into data-deficient areas. Four expeditions were undertaken to collect snails along the Lake Victoria shoreline, visiting 223 sites overall. Environmental measurements were recorded at the time of inspection and water samples taken for quantification of anion and cation concentrations. The spatial distributions of Biomphalaria choanomphala and Biomphalaria sudanica were modelled in two Bayesian multivariate models: one non-spatial and one spatial with random effects. The results showed that chloride, nitrate, sulphate and the number of sympatric snail species were significant predictors of B. choanomphala whereas habitat, water depth, pH and sulphate were significant predictors for B. sudanica. The range of spatial autocorrelation was large (572.9 km for B. choanomphala and 175.3 km for B. sudanica). Interpolating snail abundance data by kriging revealed two 'hot-spots' of high abundance of Biomphalaria. These areas should be targeted in future expeditions to ground-truth the model's predictions. Our study is the first to use Bayesian methods for determination of biomedically important snail distributions and sets crucial limnological baseline data for assessment of future changes in snail biodiversity in the Lake Victoria Basin. [ABSTRACT FROM AUTHOR]

Genetic variability within and among Bulinus truncatus of the Albertine Rift freshwater bodies were assessed to investigate the degree of inbreeding and gene flow in the snail populations. The effect of ploidy on the genetic structuring of B. truncatus is also described. We characterized the genetic structure of seven B. truncatus populations from Lake Albert, Lake Kivu, and Katosho swamp in Tanzania using five polymorphic microsatellite loci. Genetic differentiation was quantified using pairwise FST values and Nei's standard genetic distances. Different alleles were observed across all loci and genetic diversity was low although it varied greatly across populations; observed heterozygosity was, however, higher than the expected heterozygosity in three of the populations studied. Significant heterozygote deficiencies were observed coupled with significant linkage disequilibria in five populations for all the five loci examined in this study. We found significant genetic differentiation among the seven freshwater bodies; private alleles were observed across all loci indicating restricted or absence of gene flow between populations. Limited snail dispersal and the reproductive biology of B. truncatus are the major forces shaping the genetic variation observed. Low genetic variation within B. truncatus populations exposes them to a high parasite infection risk as predicted in the Red Queen hypothesis. [ABSTRACT FROM AUTHOR]

Aim To analyse the phylogeographical history of intertidal tardigrades in the North Atlantic in order to improve our understanding of geographical differentiation in microscopic organisms, and to understand the potential importance of the Mid-Atlantic Islands as stepping stones between the American and European coasts of the Atlantic Ocean. Location Twenty-four localities from the Mid-Atlantic Islands (Greenland, Iceland and the Faroe Islands) and both sides of the North Atlantic Ocean. Methods A mitochondrial marker (cytochrome c oxidase subunit I) was sequenced from individual tardigrades belonging to the genus Echiniscoides. The existence of cryptic species was detected using generalized mixed Yule coalescence analysis; lineage ages were estimated with relaxed clock methods; and the degree of geographical differentiation was analysed with samova analyses, haplotype networks and Mantel tests. Results Echiniscoides hoepneri, previously known only from Greenland, was recovered throughout the Mid-Atlantic Islands. The Faroe Islands population was isolated from Greenland and Iceland, but overall genetic variation was low. The morphospecies Echiniscoides sigismundi had high genetic variation and consisted of at least two cryptic species. A northern and a southern species were both recovered on both sides of the Atlantic, but only the northern species was found on the Mid-Atlantic Islands. The northern species showed signs of long-term isolation between the Western and Eastern Atlantic, despite the potential of the Mid-Atlantic islands to act as stepping-stones. There was no sign of long-term isolation in the southern species. The Mid-Atlantic individuals of the northern species were of Eastern Atlantic origin, but Greenland and Iceland showed signs of long-term isolation. The genetic pattern found in the southern species is not clearly geographical, and can probably be best explained by secondary contact between former isolated populations. Main conclusions North Atlantic intertidal tardigrades from the genus Echiniscoides showed strong geographical differentiation, and the Mid-Atlantic Islands seemed unimportant as stepping stones across the Atlantic. The geographical variation of the northern species of E. sigismundi suggests postglacial recolonization from several refugia. [ABSTRACT FROM AUTHOR]

Changes in habitat stability may significantly shape evolutionary patterns and processes in ancient lakes. In the present study, we use a hierarchical combination of molecular phylogenetic and coalescent approaches to investigate the evolutionary history of the endemic species of the gastropod genus Bellamya in the African rift-lake Malawi. By integrating our findings with reported palaeontological and palaeolimnological data, we demonstrate that all but one evolutionary lineage of the Pliocene Bellamya fauna in Lake Malawi became extinct. Coalescent analyses indicate that the modern radiation underwent both a sudden demographic and a spatial expansion after a genetic bottleneck. We argue that a reflooding of the lake after severe Pleistocene low stands offers a straightforward explanation for this pattern and may have triggered speciation processes in the modern endemic Bellamya radiation in Lake Malawi. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, , 130-143. [ABSTRACT FROM AUTHOR]

The freshwater snails belonging to the genus Melanoides Olivier, 1804 are widespread across tropical regions of the world and endemic species have evolved in the African Lakes Malawi, Mweru and Tanganyika. The endemic Melanoides species of Lake Malawi have been investigated several times during the last century, due to their large conchological variation, but no unambiguous answer regarding the number of species has been given. The phylogenetic relationship between morphs or genetic clones of Melanoides in Lake Malawi was inferred by phylogenetic analyses of DNA sequence data from the mitochondrial genes 16S and COI. Additional sequences from GenBank were included to investigate the relationship to other morphs from different parts of the world. For the first time, a putative secondary structure was developed for a partial region of 16S in this genus to identify the variability of the secondary structure in stems and loops. The molecular analyses indicated that several genetic clones exist in Lake Malawi and that M. tuberculata is a paraphyletic taxon. It is not clear from the results whether invasions or dispersals account for the complex situation in Lake Malawi. The basal position of M. admirabilis, endemic to Lake Tanganyika, in the inferred phylogeny indicates that Africa might be the origin of the genus. The results further indicate that three major clades of Melanoides, consisting of several genetic clones, are present in Lake Malawi; one clade consisting of invasive M. tuberculata, another of native M. tuberculata and a third consisting of the M. polymorpha-complex. It appears as if the unique development of morphs within the Melanoides genus in Lake Malawi has evolved primarily by divergence of genetic clones instead of species differentiation. [ABSTRACT FROM AUTHOR]

Background: Tardigrada is a group of microscopic invertebrates distributed worldwide in permanent and temporal aquatic habitats. Famous for their extreme stress tolerance, tardigrades are also of interest due to their close relationship with Arthropoda and Cycloneuralia. Despite recent efforts in analyzing the musculature of a number of tardigrade species, data on the class Heterotardigrada remain scarce. Aiming to expand the current morphological framework, and to promote the use of muscular body plans in elucidating tardigrade phylogeny, the myoanatomy of two heterotardigrades, Actinarctus doryphorus and Echiniscoides sigismundi, was analyzed by cytochemistry, scanning electron and confocal laser scanning microscopy and 3D imaging. We discuss our findings with reference to other tardigrades and internal phylogenetic relationships of the phylum. Results: We focus our analyses on the somatic musculature, which in tardigrades includes muscle groups spanning dorsal, ventral, and lateral body regions, with the legs being musculated by fibers belonging to all three groups. A pronounced reduction of the trunk musculature is seen in the dorsoventrally compressed A. doryphorus, a species that generally has fewer cuticle attachment sites as compared to E. sigismundi and members of the class Eutardigrada. Interestingly, F-actin positive signals were found in the head appendages of A. doryphorus. Our analyses further indicate that cross-striation is a feature common to the somatic muscles of heterotardigrades and that E. sigismundi—as previously proposed for other echiniscoidean heterotardigrades—has relatively thick somatic muscle fibers. Conclusions: We provide new insights into the myoanatomical differences that characterize distinct evolutionary lineages within Tardigrada, highlighting characters that potentially can be informative in future phylogenetic analyses. We focus our current analyses on the ventral trunk musculature. Our observations suggest that seven paired ventromedian attachment sites anchoring a large number of muscles can be regarded as part of the ground pattern of Tardigrada and that fusion and reduction of cuticular attachment sites is a derived condition. Specifically, the pattern of these sites differs in particular details between tardigrade taxa. In the future, a deeper understanding of the tardigrade myoanatomical ground pattern will require more investigations in order to include all major tardigrade lineages. [ABSTRACT FROM AUTHOR]

Background: Tardigrades are renowned for their ability to enter cryptobiosis (latent life) and endure extreme stress, including desiccation and freezing. Increased focus is on revealing molecular mechanisms underlying this tolerance. Here, we provide the first transcriptomes from the heterotardigrade Echiniscoides cf. sigismundi and the eutardigrade Richtersius cf. coronifer, and compare these with data from other tardigrades and six eukaryote models. Investigating 107 genes/gene families, our study provides a thorough analysis of tardigrade gene content with focus on stress tolerance. Results: E. cf. sigismundi, a strong cryptobiont, apparently lacks expression of a number of stress related genes. Most conspicuous is the lack of transcripts from genes involved in classical Non-Homologous End Joining. Our analyses suggest that post-cryptobiotic survival in tardigrades could rely on high fidelity transcription-coupled DNA repair. Tardigrades seem to lack many peroxins, but they all have a comprehensive number of genes encoding proteins involved in antioxidant defense. The "tardigrade unique proteins" (CAHS, SAHS, MAHS, RvLEAM), seem to be missing in the heterotardigrade lineage, revealing that cryptobiosis in general cannot be attributed solely to these proteins. Our investigation further reveals a unique and highly expressed cold shock domain. We hypothesize that the cold shock protein acts as a RNA-chaperone involved in regulation of translation following freezing. Conclusions: Our results show common gene family contractions and expansions within stress related gene pathways in tardigrades, but also indicate that evolutionary lineages have a high degree of divergence. Different taxa and lineages may exhibit unique physiological adaptations towards stress conditions involving possible unknown functional homologues and/or novel physiological and biochemical mechanisms. To further substantiate the current results genome assemblies coupled with transcriptome data and experimental investigations are needed from tardigrades belonging to different evolutionary lineages. [ABSTRACT FROM AUTHOR]

It has recently been argued that the enigmatic tardigrades (water bears) will endure until the sun dies, surviving any astrophysical calamities in Earth’s oceans. Yet, our knowledge of stress tolerance among marine tardigrade species is very limited and most investigations revolve around species living in moist habitats on land. Here, we investigate desiccation tolerance in the cosmopolitan marine tidal tardigrade, Echiniscoides sigismundi, providing the first thorough analysis on recovery upon desiccation from seawater. We test the influence on survival of desiccation surface, time spent desiccated (up to 1 year) and initial water volume. We propose analysis methods for survival estimates, which can be used as a future platform for evaluating and analysing recovery rates in organisms subjected to extreme stress. Our data reveal that marine tidal tardigrades tolerate extremely rapid and extended periods of desiccation from seawater supporting the argument that these animals are among the toughest organisms on Earth. [ABSTRACT FROM AUTHOR]

Life unfolds within a framework of constraining abiotic factors, yet some organisms are adapted to handle large fluctuations in physical and chemical parameters. Tardigrades are microscopic ecdysozoans well known for their ability to endure hostile conditions, such as complete desiccation – a phenomenon called anhydrobiosis. During dehydration, anhydrobiotic animals undergo a series of anatomical changes. Whether this reorganization is an essential regulated event mediated by active controlled processes, or merely a passive result of the dehydration process, has not been clearly determined. Here, we investigate parameters pivotal to the formation of the so-called "tun", a state that in tardigrades and rotifers marks the entrance into anhydrobiosis. Estimation of body volume in the eutardigrade Richtersius coronifer reveals an 87 % reduction in volume from the hydrated active state to the dehydrated tun state, underlining the structural stress associated with entering anhydrobiosis. Survival experiments with pharmacological inhibitors of mitochondrial energy production and muscle contractions show that i) mitochondrial energy production is a prerequisite for surviving desiccation, ii) uncoupling the mitochondria abolishes tun formation, and iii) inhibiting the musculature impairs the ability to form viable tuns. We moreover provide a comparative analysis of the structural changes involved in tun formation, using a combination of cytochemistry, confocal laser scanning microscopy and 3D reconstructions as well as scanning electron microscopy. Our data reveal that the musculature mediates a structural reorganization vital for anhydrobiotic survival, and furthermore that maintaining structural integrity is essential for resumption of life following rehydration. [ABSTRACT FROM AUTHOR]

A large number of specimens (1563) of a new Florarctus species were collected in a single intertidal sample from the Ionian Sea, Italy. The postembryonic development of the new species was studied with emphasis on external morphology and functional anatomy. The new Florarctus species was recognized as having two distinct larval stages, during which the anus is formed and the gonads mature prior to the development of a gonopore. The larval stages are followed by one adult stage in females and two adult stages in males. Cuticular structures change in morphology during postembryonic development, so that they exhibit a juvenile and an adult appearance. The accelerated growth in gonadal development could be the result of paedomorphosis, which was observed in the tendency of adult males of the new Florarctus species to maintain juvenile character conditions. [ABSTRACT FROM AUTHOR]

The marine heterotardigrades belonging to Echiniscoididae inhabit a wide range of substrates and represent the dominant tardigrade group in intertidal zones worldwide. Notably, the evolution of this family is interesting as it holds key evidence as to how heterotardigrades colonized terrestrial and freshwater environments. We inferred the phylogeny of Echiniscoididae using 28S and COI sequences from GenBank and from new specimens collected at Roscoff, France. Morphological characterizations of several major clades, inferred from the molecular data, are currently lacking. However, one of the clades is erected as a new taxon, Isoechiniscoides gen. nov., for the former Echiniscoides higginsi-group, which comprises species with isonych claws, pillars in the epicuticle and exceptionally long sensory appendages. The type species for the new genus is described here as Isoechiniscoides sifae nov. gen., sp. nov. Isoechiniscoides comprises six clawed interstitial species, whereas Echiniscoides taxa possess a higher claw number and primarily inhabit algae, barnacles and lichens in tidal zones. The new genus, Isoechiniscoides, together with morphologically undetermined specimens from British Columbia (Canada), is sister-group to all other Echiniscoididae taxa. Consequently, we propose to establish Isoechiniscoidinae subfam. nov. to accommodate the new genus. Isoechiniscoidinae provides an evolutionary link between Echiniscoidea and Arthrotardigrada. The latter comprises exclusively marine and mostly interstitial species. [ABSTRACT FROM AUTHOR]

The so-called 'tun' state is best known from limno-terrestrial tardigrades and rotifers that rely on this compact body shape for anhydrobiotic survival. Little is known of tun formation in marine species and the evolutionary origin of the state is presently unknown. Here, we investigate desiccation tolerance and tun formation in the marine tidal echiniscoidean tardigrade, Echiniscoides sigismundi (M. Schultze, 1865). Groups of approximately 20 E. sigismundi sampled from Lynæs (Denmark) were dehydrated on filter paper from seawater as well as ultrapurified water and kept for 48 h at 5 °C, after which they were rehydrated in seawater. The activity and behaviour of the tardigrades was examined under a light microscope, whereas scanning electron microscopy was used for high-resolution three-dimensional imaging. When dehydrated from seawater, E. sigismundi enters a tun, however, when exposed to ultrapurified water, the tardigrade swells and becomes incapable of movement, and thus incapable of tun formation. Nonetheless, E. sigismundi tolerates being dehydrated from ultrapurified water, revealing an exceptional and unparalleled resilience towards losing structural integrity. Our results confirm previous investigations, which suggest that tun formation relies on a functional musculature. They further suggest that tun formation may have evolved as a response to elevated external pressure rather than desiccation per se. [ABSTRACT FROM AUTHOR]

Little is still known about the diversity and evolution of marine arthrotardigrades, as they are generally difficult to sample, resulting in a limited amount of molecular data for barcoding and phylogenetic studies. With the current study, we provide the first investigation into COI haplotype diversity in a marine tanarctid and at the same time readdress arthrotardigrade phylogeny. Specifically, we provide COI mtDNA, 18S and 28S rDNA sequences from a population of Actinarctus doryphorus (Tanarctidae) sampled off the coast of Roscoff, France and further provide new 18S sequences from two marine echiniscoidids. A. doryphorus COI sequences confirmed the presence of a single species and further revealed five haplotypes shared among nine sequenced individuals. Our 18S and 28S rDNA datasets were individually and combined analysed with Bayesian inference and Maximum Likelihood. Actinarctus doryphorus was placed together with Tanarctus sequences within a maximally supported Tanarctidae, confirming previous interpretations that the clade is distinct from Halechiniscidae. Although several studies in recent decades have concluded that the marine arthrotardigrades are paraphyletic, recent studies have argued that the clade may not be paraphyletic. Our phylogenetic analyses consistently inferred Arthrotardigrada as paraphyletic, as the clade includes the monophyletic Echiniscoidea. Accordingly, we propose that it is time to suppress the order Arthrotardigrada as it clearly does not reflect tardigrade phylogeny.

Tardigrades are microscopic aquatic animals renowned for their tolerance toward extreme environmental conditions. The current study is the first to investigate their tolerance toward heavy metals and we present a novel tardigrade toxicant tolerance assay based on activity assessments as a measure of survival. Specifically, we compare tolerance toward copper in four species representing different evolutionary lineages, habitats and adaptation strategies, i.e., a marine heterotardigrade, Echiniscoides sigismundi , a limno-terrestrial heterotardigrade, Echiniscus testudo , a limno-terrestrial eutardigrade, Ramazzottius oberhaeuseri , and a marine eutardigrade, Halobiotus crispae . The latter was sampled at a time of year, when the population is predominantly represented by aberrant P1 cysts, while the other species were in normal active states prior to exposure. Based on volume measurements and a general relation between body mass and copper tolerance, expected tardigrade EC50 values were estimated at 0.5-2 μg l -1 . Following 24 h of exposure, tolerance was high with no apparent link to lineage or habitat. EC50s (95% CI), 24 h after exposure, were estimated at 178 (168-186) and 310 (295-328) μg l -1 , respectively, for E. sigismundi and R. oberhaeuseri , whereas E. testudo and H. crispae were less affected. Highest tolerance was observed in H. crispae with a mean ± s.e.m . activity of 77 ± 2% ( n = 3) 24 h after removal from ~3 mg l -1 copper, suggesting that tardigrade cysts have increased tolerance toward toxicants. In order to identify putative tolerance related genes, an E. sigismundi transcriptome was searched for key enzymes involved in osmoregulation, antioxidant defense and copper metabolism. We found high expression of Na/K ATPase and carbonic anhydrase, known targets for copper. Our transcriptome, furthermore, revealed high expression of antioxidant enzymes, copper transporters, ATOX1, and a Cu-ATPase. In summary, our results indicate that tardigrades express well-known key osmoregulatory enzymes, supporting the hypothesis that copper inhibits sodium turnover as demonstrated for other aquatic organisms. Tardigrades, nevertheless, have high tolerance toward the toxicant, which is likely linked to high expression of antioxidant enzymes and an ability to enter dormant states. Tardigrades, furthermore, seem to have a well-developed battery of cuproproteins involved in copper homeostasis, providing basis for active copper sequestering and excretion.

Marine caves are known to support a rich macrofauna; however, few studies have focused on meiofauna. Marine cave meiofaunal tardigrades have been reported from Japan and the Mediterranean Sea and a preliminary list of species including a redescription of Actinarctus neretinus Grimaldi de Zio, D'Addabbo Gallo, Morone De Lucia, Vaccarella and Grimaldi, 1982 was reported from Fish Rock Cave and Jim's Cave on the coast of Australia. This study is the fourth in a series describing the unique meiofauna in two Australian submarine caves located off the coast of New South Wales, describing nine new species.        Only 67 tardigrades were collected from the two caves, yet these contained a high diversity of at least 16 different species which are quite different in the two caves. The fauna includes nine arthrotardigrade genera: Actinarctus, Batillipes, Dipodarctus, Halechiniscus, Raiarctus, Styraconyx, Tanarctus, Tholoarctus, and Wingstrandarctus. This fauna is different from that reported for the high energy beaches along the East Coast of Australia.        We describe nine new species comprising a single batillipedid and eight halechiniscids: Batillipes solitarius nov. sp., Dipodarctus australiensis nov. sp., Dipodarctus susannae nov. sp., Raiarctus jesperi nov. sp., Raiarctus katrinae nov. sp., Tanarctus hirsutospinosus nov. sp., Tholoarctus oleseni nov. sp., Wingstrandarctus stinae nov. sp. and Wingstrandarctus unsculptus nov. sp.