A laboratory strain identified as “Hypsibius dujardini” is one of the best studied tardigrade strains: it is widely used as a model organism in a variety of research projects, ranging from developmental and evolutionary biology through physiology and anatomy to astrobiology. Hypsibius dujardini, originally described from the Île-de-France by Doyère in the first half of the 19th century, is now the nominal species for the superfamily Hypsibioidea. The species was traditionally considered cosmopolitan despite the fact that insufficient, old and sometimes contradictory descriptions and records prevented adequate delineations of similar Hypsibius species. As a consequence, H. dujardini appeared to occur globally, from Norway to Samoa. In this paper, we provide the first integrated taxonomic redescription of H. dujardini. In addition to classic imaging by light microscopy and a comprehensive morphometric dataset, we present scanning electron photomicrographs, and DNA sequences for three nuclear markers (18S rRNA, 28S rRNA, ITS-2) and one mitochondrial marker (COI) that are characterised by various mutation rates. The results of our study reveal that a commercially available strain that is maintained in many laboratories throughout the world, and assumed to represent H. dujardini sensu stricto, represents, in fact, a new species: H. exemplaris sp. nov. Redescribing the nominal taxon for Hypsibiidae, we also redefine the family and amend the definitions of the subfamily Hypsibiinae and the genus Hypsibius. Moreover, we transfer H. arcticus (Murray, 1907) and Hypsibius conifer Mihelčič, 1938 to the genus Ramazzottius since the species exhibit claws and eggs of the Ramazzottius type. Finally, we designate H. fuhrmanni as subjectively invalid because the extremely poor description precludes identifying neotype material.
Key words: barcoding, diagnostic key, development tracking, in vitro culture, M. berladnicorum, M. variefidum sp. nov. AbstractIn this paper we describe a new apochelan species, Milnesium variefidum sp. nov. from Scotland and provide novel morphological and molecular data for Milnesium berladnicorum Ciobanu et al., 2014. The new species differs from the most similar M. berladnicorum by the presence of developmental dimorphism in claw configuration, absent or weakly developed cuticular bars under claws I-III, a different arrangement of cuticular pseudoplates, and by differences in the sequences of three nuclear DNA fragments: 18S rRNA (p-distance: 0.6%), 28S rRNA (2.0%), ITS-2 (9.3%), and on mitochondrial gene COI (12.4%). Although ontogenetic claw configuration change was suspected to occur in some Milnesium species, we are the first to document it through the combined use of traditional, molecular and experimental methodologies. We discuss the implications of the observed phenomenon for the taxonomy of the genus and propose a new diagnostic key to all Milnesium species described up to the end of 2015. We also review other traits used for species differentiation in the genus and offer recommendations to improve the quality of future descriptions as well as suggest a need for integrative redescriptions of the known species. Finally, we propose to suppress M. dujiangensis and M. tardigradum trispinosum and suggest that M. alpigenum and M. quadrifidum are valid species that require thorough redescriptions.
Intraspecific variability of tardigrades is a poorly investigated field. The great majority of tardigrade species were described using single populations, sometimes even single individuals. Moreover, intraspecific variability between multiple, integratively verified populations has never been described for any tardigrade species. In this study, we analyzed morphological, morphometric, and genetic variability of nine European populations of Milnesium tardigradum, one of the first described tardigrade species that until recently was thought to be cosmopolitan. We found that the nine populations of M. tardigradum exhibited low to moderate diversity in the nuclear ITS‐2 marker (0.2%–4.0%, 1.1% on average) and low to high divergence in the mitochondrial COI fragment (0.2%–11.4%, 4.4% on average). Despite the considerable variability in COI, all M. tardigradum populations exhibited mostly uniform morphology and morphometry. Moreover, we analyzed ontogenetic variability in claw and lamellae configuration under scanning electron microscopy, which resulted in the description of novel taxonomically important traits for M. tardigradum. The phylogenetic analysis unambiguously showed that all GenBank sequences predating the redescription of M. tardigradum and labeled as “M. tardigradum” or “M. cf. tardigradum” represent other species. Thus, the neotype population from Germany and the additional eight European populations analyzed in this study are the only confirmed records of M. tardigradum to date. Finally, in order to clarify ontogenetic nomenclature, we propose distinct names for the first and second immature instars in the order Apochela (hatchling and juvenile, respectively) and concise definitions of time and direction of ontogenetic changes in claw configuration (i.e., early/late and positive/negative change).
Despite advances in molecular systematics, the taxonomy of tardigrades still depends largely on morphological and morphometric traits. The fact that the variability of any biological trait is determined by the interaction between genetics and environment prompts a very fundamental question: is it possible for tardigrades of the same genotype, but originating from various habitats that differ in environmental conditions, to have phenotypes so different that they would be erroneously classified as different species taxa by means of classical taxonomy? Here, we present the results of a broad and fully controlled laboratory experiment in which we investigated the phenotypic plasticity of a number of traits that are traditionally considered to be taxonomically important. In order to achieve this, we have cultured six tardigrade species belonging to four eutardigrade families (Milnesiidae, Hypsibiidae, Isohypsibiidae, and Macrobiotidae) under five experimental regimes, reflecting key environmental factors that are likely to vary in natural habitats (i.e. temperature and food availability). We then measured a number of key taxonomic traits and compared their dimensions between the treatments. Over two years of experimentation we have obtained more than 28 000 morphometric measurements for over 2300 individuals. Such an extensive data set allowed us to test some of the fundamental assumptions of classic tardigrade taxonomy. We found that in the five parachelan species analysed, the great majority of both absolute and relative traits differed significantly between the treatments, whereas there were no significant differences in the apochelan species. Overall, tardigrades grew largest under the low-temperature treatment, whereas the smallest specimens were observed under high-temperature and low-food regimes. However, the prevalent statistical significance resulted mainly from the considerable statistical power of our analyses, and not from effect sizes, which varied mostly between low and moderate. In other words, the differences, although consistent, were minor in terms of taxonomical significance, and probably would not be considered by classic taxonomists as sufficient to designate animals from different treatments as separate taxa.
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