Factors influencing the distribution of aquatic plant communities in Irish canals

Terrestrial tardigrades are often found in the lichens and mosses growing on trees and rocks. The assertion that tardigrades in these habitats are very patchy in their distribution has rarely been backed by quantitative sampling. This study assesses spatial variability in tardigrade populations inhabiting small patches (0.1 cm² to over 5 cm²) of moss and lichen on trees and rocks at three sites in the United States of America. Tardigrades were collected from four replicate rocks in the Ouachita Mountains of Arkansas, with 30 lichen patches collected on two adjacent boulders and 20 moss patches on a second pair of boulders. In Fort Myers and in Citrus Springs, Florida, 30 lichen patches per tree were collected from two pairs of trees. The tardigrades in each sample were extracted, mounted, identified, and counted. The variation in tardigrade abundance among lichen or moss patches within rocks or trees was very high; the only consistent pattern was that very small patches usually lacked tardigrades. Tardigrade diversity and abundance also varied greatly within sites when lichens and mosses of the same species from different rocks and trees were compared (in the most extreme case one tree had numerous individuals of two tardigrade species present while the other had almost no tardigrades). The results of this quantitative sampling support the assertion that tardigrades are very patchy in distribution. Given the considerable time investment required for the quantitative processing of tardigrade samples, this high spatial variability in tardigrade diversity and abundance requires that researches testing ecological hypotheses about tardigrade abundance check variability before deciding how many samples to take.     

How to Crawl and Dehydrate on Moss

Tardigrades (Macrobiotus sp., Echiniscus testudo) were videotaped when moving on single plants of mosses with different water retaining capacities. On a fully hydrated Encalyptra contorta plant encased in a continuous water column, tardigrades could hardly crawl rapidly in a specific direction. This was probably due to their low specific weight and the poor ability of their claws to cling to the substrate. As tardigrades were not able to walk on the undersurface of the leaves under these conditions, they were rinsed out of a fully soaked moss cushion in the course of time. Fully hydrated Polytrichum formosum plants did not show a visible water film and locomotion of tardigrades could not be observed on this moss. Activity of animals appeared to be confined to the droplets in the leaf axils. During dehydration, the tardigrades stopped moving and formed a tun on the spot.     

Tardigrades of the Taimyr peninsula with descriptions of two new species

This is the first report on tardigrades from the Taimyr peninsula. Seventy-one species of tardigrades were recorded, new to Russian fauna and two of which are new to science. Diphascon boreale sp. nov. is closely related to D. brevipes (Marcus, 1936) but easily distinguished from it by claw structure and the absence of cuticular bars on the legs. Isohypsibius roberti sp. nov. belongs to the elegans group of the genus Isohypsibius ; it diners from other species in the group by having a long and thin buccal tube and large lunulae with small teeth on leg IV. The eggs of Ramazzottius montivagus are here described for the first time. The almost complete absence of tardigrades in and around Norilsk is noted. This is related to the heavy industrial pollution from a local nickel-copper plant.     

Experimentally induced anhydrobiosis in the tardigrade Richtersius coronifer: phenotypic factors affecting survival

The ability of some animal taxa (e.g., nematodes, rotifers, and tardigrades) to enter an ametabolic (cryptobiotic) state is well known. Nevertheless, the phenotypic factors affecting successful anhydrobiosis have rarely been investigated. We report a laboratory study on the effects of body size, reproductive condition, and energetic condition on anhydrobiotic survival in a population of the eutardigrade Richtersius coronifer. Body size and energetic condition interacted in affecting the probability of survival, while reproductive condition had no effect. Large tardigrades had a lower probability of survival than medium-sized tardigrades and showed a positive response in survival to energetic condition. This suggests that energy constrained the possibility for large tardigrades to enter and to leave anhydrobiosis. As a possible alternative explanation for low survival in the largest specimens we discuss the expression of senescence. In line with the view that processes related to anhydrobiosis are connected with energetic costs we documented a decrease in the size of storage cells over a period of anhydrobiosis, showing for the first time that energy is consumed in the process of anhydrobiosis in tardigrades.     

Dynamics of Long-term Anhydrobiotic Survival of Lichen-dwelling Tardigrades

It is not rare to find in references that anhydrobiotic tardigrades can survive for more than a century. However, a closer look at the empirical evidence provides very little support that tardigrades are capable of surviving dried for such a long time. In order to resolve this discrepancy, we carried out a study to evaluate the long-term survival of naturally dried tardigrades. A large fragment of dry lichen (Xanthoria parietina) was collected in the field two days after a rainy day in 1999. The dry lichen was stored inside a paper bag in the laboratory at room temperature and humidity and under atmospheric oxygen exposure. Replicates of the dry lichen were re-hydrated after various time periods of storage, with all tardigrades extracted and the survivors enumerated. Five species of tardigrades were found, but two of them only occasionally. Ramazzottius oberhaeuseri, Echiniscus testudo and Echiniscus trisetosus were sufficiently represented for statistical analysis. At the beginning of the experiment 91.1% of R. oberhaeuseri and 71.7% of Echiniscus spp. were alive. R. oberhaeuseri survived up to 1604 days, while Echiniscus spp. lived up to 1085 days. Recovery after four years of anhydrobiosis has to be considered a very good long-term survival, which is important from an ecological and evolutionary point of view.     

A survey of the terrestrial Tardigrada of the Vestfold Hills,Antarctica

A survey of the terrestrial tardigrades inhabiting growths of algae, lichens and mosses in the Vestfold Hills, Antarctica, was carried out at 11 and 35 sites during the austral summers of 1980 and 1982, respectively. In all, 24 species of plants were collected from which four genera and four species of Tardigrada were recovered. A key to the tardigrades of the area is presented. The distribution and associational patterns of the tardigrades are discussed in the context of other studies of antarctic Tardigrada.     

The geographic distribution of metazoan microfauna on East Antarctic nunataks

The abundance and distribution of nematodes, rotifers and tardigrades in samples from nunataks in continental Antarctica were investigated during four Antarctic expeditions in the austral summers of 1991/92, 1993/94, 1996/97 and 2001/02. Altogether 368 samples were collected from 14 nunataks and one oasis in East Antarctica. Nematodes were found in 35%, rotifers in 67% and tardigrades in 40% of all samples. Fifty-four microfaunal taxa were identified. Of these, 27 were nematodes, 8 tardigrades and 19 rotifers. The size and geographic location of the nunatak and oasis influenced the abundance and taxonomic composition of the microfauna. The highest abundance and diversity of nematodes were found on large nunataks close to the coast. Nematodes were not found on small inland nunataks. Very high population densities of tardigrades and rotifers were found on two small and isolated nunataks. No microfauna was found on the two southernmost nunataks (Okkenhaugrusta and Vardeklettane), or on the smallest one (Utsikta). The Sørensens Quotient of Similarity was generally low, especially between faunas on nunataks in different mountain ridges. The results indicate rather limited rates of dispersal and colonization between nunataks.     

Population density and species composition of moss-living tardigrades in a boreo-nemoral forest

This study investigates for the first time the tardigrade fauna in a variety of different mosses from a coniferous forest and an adjacent clear-cut area in southern Sweden. Tardigrades were found in a majority of the samples. Sixteen species were recorded, of which the cosmopolitan species Macrobiotus hufelandi was the far most common. Some mosses, particularly species with "wefts" growth form, contained more tardigrades than other mosses, indicating that growth form may have an impact on tardigrade abundance. Mosses of the same species collected from a forest and from a clear-cut, respectively, did not show a general trend in the overall abundance of tardigrades, but the forest tended to contain more species. Five species of tardigrades ( Murrayon dianae, Isohypsibius sattleri, Platicrista angustata, Diphascon belgicae and Diphascon pingue ) never previously reported from Sweden were recorded.     

Seasonal and Altitudinal Variation in the Distribution and Abundance of Tardigrada on Dugger Mountain, Alabama

A seasonal survey of the distribution of terrestrial tardigrades on Dugger Mountain, Alabama, was conducted during the time period from April 1997 through April 1998. Cryptogams from five trees (Quercus alba), three on north-facing slopes and two on south-facing slopes, were sampled seasonally at three stations (645 m, 410 m, 183 m) along an unnamed tributary of the South Fork of Terrapin Creek. Trees were chosen based on their location outside the riparian zone. Tardigrades were extracted from the samples, mounted individually in Hoyer's medium, and identified to species with phase microscopy. Seasonal and altitudinal variations in the distribution of the populations on the north- and south-facing slopes were determined. Present on Dugger Mountain were tardigrades belonging to 12 species (Macrobiotus cf. areolatus/tonollii, Macrobiotus cf. echinogenitus, Macrobiotus islandicus, Macrobiotus richtersi, Minibiotus intermedius, Milnesium tardigradum, Diphascon pingue, Hypsibius pallidus, Echiniscus cf. arctomys, Echiniscus virginicus, Pseudechiniscus ramazzottii, and Pseudechiniscus suillus). Due to the small numbers of individuals of each species, the total numbers of tardigrades of all species were pooled. There were no significant differences in the mean number of species or the mean number of all tardigrades per sample at each station (altitude). However, seasonal differences in both abundance and number of species were detected in pooled samples due to the high numbers collected in spring 1997.     

Tardigrades of the Australian Antarctic: Macquarie Island, sub-Antarctica

Sub-Antarctic Macquarie Island lies in the Southern Ocean between Tasmania and Antarctica and just above the Antarctic Convergence. Extensive flora and fauna samples were collected during the 1977—78 Australian Museum Expedition. Thirteen genera and 25 species of tardigrades are reported from among the 8725 specimens recovered from the 72 samples. Association between species of tardigrades and between species of tardigrades and plant species are discussed. Possible predator-prey relationships are suggested by patterns of association.     

Tardigrades of the Australian Antarctic Territories: the Windmill Islands, East Antarctica

Six species of tardigrades, Pseudechiniscus suillus, Macrobiotus sp., Hypsibius antarcticus, Ramajendas frigidus, Diphascon chilenense and Diphascon pingue were extracted from mosses and lichens from the ice-free regions of the Windmill Islands near Casey Base, East Antarctica. Significant positive associations were found between the three common species ( Pseudechiniscus suillus, Hypsibius antarcticus, Diphascon chilenense ) and bryophytes, whereas strong negative associations were found between these species and algae and lichens. There were additional interspecific associations between the common species of tardigrades as well as between tardigrades, nematodes and rotifers.     

First evidence of achiasmatic male meiosis in the water bears Richtersius coronifer and Macrobiotus richtersi (Eutardigrada, Macrobiotidae)

Chromosome behaviour during male meioses has been studied in two bisexual amphimictic populations of two tardigrade species, namely Richtersius coronifer and Macrobiotus richtersi (Eutardigrada, Macrobiotidae). Both bisexual populations exhibit a diploid chromosome number 2n=12 and no sex chromosomes were identified. DAPI staining and C-banding data indicate that all chromosomes of the bisexual population of R. coronifer are acrocentric. In both species, at male meiotic prophase, all six bivalent homologous chromosomes are aligned side by side along their length and show no evidence of chiasmata. However, in the oocytes of both species a chiasma is generally present in each bivalent at diplotene stage. Lack of recombination is previously unknown in tardigrades, but is a well known phenomenon in many other metazoans where it is always restricted to the heterogametic sex. In tardigrades there is no evidence of heterochromosomes, but it does not mean that in tardigrades, the heterogametic sex does not exist. The adaptive and evolutionary significance of achiasmatic meiosis is discussed.     

Tardigrade Fauna of the Region between Oka and Volga Rivers (Russia) and Patterns of their Distribution in Substrates and Habitats

The region between the Oka and Volga rivers has been largely neglected with respect to tardigrade biodiversity. In the present study a total of 21 species from 7 genera, 3 families, 2 orders and only 1 class of tardigrades were present, and some of their ecological preferences were noted. The tardigrades in the studied region were mainly cosmopolitan. Eurytopic, hygrophilic and xerophilic species were common while obligate freshwater species were not found. There is a marked similarity in the tardigrade fauna within similar habitats from different locations within the region.     

Partial mitochondrial gene arrangements support a close relationship between Tardigrada and Arthropoda

Regions (about 3.7-3.8 kb) of the mitochondrial genomes (rrnL-cox1) of two tardigrades, a heterotardigrade, Batillipes pennaki, and a eutardigrade, Pseudobiotus spinifer, were sequenced and characterized. The gene order in Batillipes was rrnL-V-rrnS-Q-I-M-nad2-W-C-Y-cox1, and in Pseudobiotus it was rrnL-V-rrnS-Q-M-nad2-W-C-Y-cox1. With the exception of the trnI gene, the two tardigrade regions have the same gene content and order. Their gene orders are strikingly similar to that of the chelicerate Limulus polyphemus (rrnL-V-rrnS-CR-I-Q-M-nad2-W-C-Y-cox1), which is considered to be ancestral for arthropods. Although the tardigrades do not have a distinct control region (CR) within this segment, the trnI gene in Pseudobiotus is located between rrnL-trnL1 and trnL2-nad1, and the trnI gene in Batillipes is located between trnQ and trnM. In addition, the 106-bp region between trnQ and trnM in Batillipes not only contains two plausible trnI genes with opposite orientations, but also exhibits some CR-like characteristics. The mitochondrial gene arrangements of 183 other protostomes were compared. 60 (52.2%) of the 115 arthropods examined have the M-nad2-W-C-Y-cox1 arrangement, and 88 (76.5%) the M-nad2-W arrangement, as found in the tardigrades. In contrast, no such arrangement was seen in the 70 non-arthropod protostomes studied. These are the first non-sequence molecular data that support the close relationship of tardigrades and arthropods.     

Aggregation effects on anhydrobiotic survival in the tardigrade Richtersius coronifer

For anhydrobiotic metazoans the rate of desiccation is an important factor influencing the probability of survival in a dry anhydrobiotic state. Formation of animal aggregations, in which the exposed body surface area of individual animals is reduced, represents one way to reduce the rate of evaporation. Such aggregations have earlier been documented in e.g., nematodes. We experimentally evaluate the effect of aggregation size (number of animals in a group of desiccating animals) on anhydrobiotic survival in the eutardigrade Richtersius coronifer. The experiment shows that aggregation provides a clear improvement on anhydrobiotic survival. The most likely explanation for this is that aggregated animals were exposed to a lower rate of desiccation. Although the empirical evidence of aggregation in tardigrades is scarce, our study suggests that aggregation could potentially be an important survival factor for tardigrades living in environments characterized by periods of rapid desiccation.     

Genome size variation in Tardigrada

The genome size of tardigrades has been documented to date in seven species of eutardigrades, and it is among the lowest in invertebrates. In this paper the evaluation of genome size is extended to eight other speeies of eutardigrades and to one species of heterolardigradcs, in order to obtain a more comprehensive picture of minimum DNA content evolution in this phylum. The genome size of the heterotardigrade Pseudechiniscus juanitae (0.6 pg), measured using le cells, is found to resemble that of some eutardigrades. Thus, it is not possible to discriminate between the two classes of tardigrades on this basis. Literature and new data on eutardigrades also suggest that families or subfamilies cannot be discriminated on the basis of genome size. Nevertheless, genome size and sperm cell shape are correlated. A lower DNA content always corresponds to a more specialized male gamete. In general, in tardigrades, the low genome size and its low variation could be related to the high specialization of the phylum. Preliminary data on nuclear AT contents suggest that the variations in those values are correlated with genome size variations.     

Interspecific Association and Substrate Specificity in Tardigrades from Florida, Southeastern United States

The distribution of terrestrial tardigrades in the Gulf Coast states of the United States is poorly known. Only one species has been reported in Florida. In this study moss, liverwort, lichen and fern samples (47 identified species) from trees and shrubs were collected from all 67 Florida counties. These samples contained 20 species of tardigrade. All possible pairs of tardigrade species and tardigrade and substrate were tested for interspecific association. Only three significant negative and one positive interspecific association between tardigrades were detected. Evidence for substrate specificity was weak. Although some tardigrade species were significantly associated with mosses or foliose lichens in general, no significant association between a tardigrade species and a substrate species was detected.