Green armoured tardigrades (Echiniscidae: Viridiscus), including a new species from the Southern Nearctic, exemplify problems with tardigrade variability research.

Ranges of tardigrade intraspecific and interspecific variability are not precisely defined, both in terms of morphology and genetics, rendering descriptions of new taxa a cumbersome task. This contribution enhances the morphological and molecular dataset available for the heterotardigrade genus Viridiscus by supplying new information on Southern Nearctic populations of V. perviridis, V. viridianus, and a new species from Tennessee. We demonstrate that, putting aside already well-documented cases of significant variability in chaetotaxy, the dorsal plate sculpturing and other useful diagnostic characters, such as morphology of clavae and pedal platelets, may also be more phenotypically plastic characters at the species level than previously assumed. As a result of our integrative analyses, V. viridianus is redescribed, V. celatus sp. nov. described, and V. clavispinosus designated as nomen inquirendum, and its junior synonymy with regard to V. viridianus suggested. Morphs of three Viridiscus species (V. perviridis, V. viridianus, and V. viridissimus) are depicted, and the implications for general echiniscid taxonomy are drawn. We emphasise that taxonomic conclusions reached solely through morphological or molecular analyses lead to a distorted view on tardigrade α-diversity.

The importance of being integrative: a remarkable case of synonymy in the genus Viridiscus (Heterotardigrada: Echiniscidae).

There are two predominant sources of taxonomically useful morphological variability in the diverse tardigrade family Echiniscidae: the internal structure and surface sculpture of the cuticular plates covering the dorsum (sculpturing) and the arrangement and morphology of the trunk appendages (chaetotaxy). However, since the appendages often exhibit intraspecific variation (they can be reduced or can develop asymmetrically), sculpturing has been considered more stable at the species level and descriptions of new echiniscid species based solely on morphology are still being published. Here, we present a case study in which a detailed analysis of the morphology and multiple genetic markers of several species of the genus Viridiscus shows that cuticular sculpture may also exhibit considerable intraspecific variation and lead to false taxonomic conclusions. In a population collected from the eastern Nearctic, in the type locality of the recently described species V. miraviridis, individuals with transitional morphotypes between those reported for V. viridissimus and V. miraviridis were found. Importantly, all morphotypes within the viridissimus-miraviridis spectrum were grouped in a single monospecific clade according to rapidly evolving markers (ITS-1, ITS-2 and COI). Given the morphological and genetic evidence, we establish V. miraviridis as a junior synonym of V. viridissimus. This study explicitly demonstrates that a lack of DNA data associated with morphological descriptions of new taxa jeopardizes the efforts to unclutter tardigrade systematics. Additionally, V. perviridis and V. viridissimus are reported from Lâm Dong Province in southern Vietnam, which considerably broadens their known geographic ranges.

New Indomalayan Nebularmis species (Heterotardigrada: Echiniscidae) provoke a discussion on its intrageneric diversity.

Recent years have brought undeniable progress in tardigrade taxonomy, and speciose complexes were detected in a number of phylogenetic lineages. The family Echiniscidae is one such lineage; it is one of the most diverse groups of limno-terrestrial tardigrades and can be characterized as having achieved great evolutionary success. In this contribution, using populations representing several species that originated from the Indomalayan region, we reconstructed phylogenetic affinities within Nebularmis, a recently erected genus within the Echiniscus lineage. Nebularmis auratus sp. nov. and Nebularmis burmensis sp. nov. are described from the Eastern Yoma Mountains and the Shan Hills (Myanmar), Nebularmis bhutanensis sp. nov. is described from the Eastern Himalayas (Bhutan), and Nebularmis indicus sp. nov. is described from the foothills of the Western Ghats (Goa, India). Moreover, males are reported in populations of the last two species. All known members of the genus can be phenotypically differentiated based on minute details of their dorsal sculpture and claws. Moreover, a very wide tropical distribution is demonstrated for Nebularmis cirinoi, recorded for the first time from islands of the Malay Archipelago. Furthemore, novel morphological, genetic, and geographic data allowed for the clarification of the generic diagnosis. Currently available data favor a scenario under which Nebularmis evolved in Southeast Asia and later dispersed to other regions of the globe.

Phylogeography and morphological evolution of Pseudechiniscus (Heterotardigrada: Echiniscidae).

Tardigrades constitute a micrometazoan phylum usually considered as taxonomically challenging and therefore difficult for biogeographic analyses. The genus Pseudechiniscus, the second most speciose member of the family Echiniscidae, is commonly regarded as a particularly difficult taxon for studying due to its rarity and homogenous sculpturing of the dorsal plates. Recently, wide geographic ranges for some representatives of this genus and a new hypothesis on the subgeneric classification have been suggested. In order to test these hypotheses, we sequenced 65 Pseudechiniscus populations extracted from samples collected in 19 countries distributed on 5 continents, representing the Neotropical, Afrotropical, Holarctic, and Oriental realms. The deep subdivision of the genus into the cosmopolitan suillus-facettalis clade and the mostly tropical-Gondwanan novaezeelandiae clade is demonstrated. Meridioniscus subgen. nov. is erected to accommodate the species belonging to the novaezeelandiae lineage characterised by dactyloid cephalic papillae that are typical for the great majority of echiniscids (in contrast to pseudohemispherical papillae in the suillus-facettalis clade, corresponding to the subgenus Pseudechiniscus). Moreover, the evolution of morphological traits (striae between dorsal pillars, projections on the pseudosegmental plate IV', ventral sculpturing pattern) crucial in the Pseudechiniscus taxonomy is reconstructed. Furthermore, broad distributions are emphasised as characteristic of some taxa. Finally, the Malay Archipelago and Indochina are argued to be the place of origin and extensive radiation of Pseudechiniscus.

High Mountain Echiniscid (Heterotardigrada) Fauna of Taiwan.

Taiwan lies at the transitional zone between the East Palaearctic and Oriental regions, which translates into both Palaearctic and Indomalayan taxa being present on the island. Furthermore, large habitat heterogeneity and high mountains contributed to the rise of conditions favouring allopatric speciation and the emergence of endemic species. The tardigrade fauna of Taiwan is poorly studied, and the aim of this contribution is to provide new data on the members of the family Echiniscidae, the largest limno-terrestrial group of the class Heterotardigrada, found at high elevations in central Taiwan. We report 11 species grouped in 5 genera: Claxtonia (1 species), Echiniscus (3 species), Hypechiniscus (1 species), Nebularmis (2 species), and Pseudechiniscus (4 species). All are new to Taiwan, including 5 species that are new to science, 4 or which are described herein by means of integrative taxonomy: Hypechiniscus crassus sp. nov. (the exarmatus morphogroup), Pseudechiniscus (Meridioniscus) dreyeri sp. nov., Pseudechiniscus (Pseudechiniscus) formosus sp. nov., and Pseudechiniscus (Pseudechiniscus) totoro sp. nov. The new findings also help to clarify the description of Echiniscus clevelandi Beasley, 1999, and supplement the phylogenies of the Echiniscus virginicus complex and of the genera Hypechiniscus, Nebularmis and Pseudechiniscus.

Continued Exploration of Tanzanian Rainforests Reveals a New Echiniscid Species (Heterotardigrada).

The Afrotropical tardigrade fauna is insufficiently studied, and consequently its diversity in this region is severely underestimated. Ongoing sampling in the Udzungwa Mountains, Morogoro Region of Tanzania has revealed a new representative of the genus Echiniscus C.A.S. Schultze, 1840 (Echiniscidae). Echiniscus tantulus sp. nov. belongs to the spinulosus group, but it stands out from other members of this speciose Echiniscus clade by having a heteromorphic sculpture of the dorsal plates and an uncommonly stable body appendage configuration A-C-C d -D d -E. The new species is characteristic by being equipped with long dorsal spines and very short lateral spicules, which so far have been found only in one other species of the group, Echiniscus spinulosus (Doyère, 1840). An updated checklist of Tanzanian Echiniscidae is provided, incorporating recent advances in their classification.

Revised Cornechiniscus (Heterotardigrada) and new phylogenetic analyses negate echiniscid subfamilies and tribes.

Echiniscidae are undoubtedly the most thoroughly studied lineage of the class Heterotardigrada. Recently, new subfamilies and tribes grouping echiniscid genera based on traditionally recognized morphological clues have been proposed. Here, by integrative analyses of morphology and DNA sequences of numerous populations of a rare genus Cornechiniscus, we show that this hypothesized classification is artificial. Specifically, we demonstrate that Echiniscinae are diphyletic, as Bryodelphax forms a distinct phyletic lineage within Echiniscidae, and Pseudechiniscinae are polyphyletic, with Mopsechiniscus being indirectly related to Pseudechiniscus, which is closer to the Echiniscus-like genera than to other genera with pseudosegmental plates. Consequently, the subfamilies and tribes are considered as unsupported from the phylogenetic and morphological point of view. The genus Cornechiniscus is revised, and the phenotypic diagnoses of several species are updated thanks to new rich material from Africa, Asia and Europe. Cornechiniscus imperfectus sp. nov. is described from mountains of Kyrgyzstan, being the second appendaged species within the genus and the third known to exhibit dioecy. A taxonomic key to the genus is provided. Systematic positions of Acanthechiniscus and Multipseudechiniscus are also discussed. Acanthechiniscus goedeni is confirmed to be a member of the genus Acanthechiniscus.

Untangling the Echiniscus Gordian knot: paraphyly of the "arctomys group" (Heterotardigrada: Echiniscidae).

The genus Echiniscus C.A.S. Schultze, 1840, one of the earliest established and speciose tardigrade genera, has been hypothesized to be polyphyletic. Moreover, the presence or absence of trunk appendages, the type of cuticular sculpturing and body colour have been argued to hold taxonomic significance at the genus level in Echiniscus-line taxa. Here, by combining morphological and genetic analyses, we demonstrate that the so-called "arctomys group", i.e. Echiniscus spp. lacking trunk appendages, comprises numerous separate evolutionary lineages within the family Echiniscidae. As a result, we erect five new echiniscid genera: Barbariagen. nov., presumably of Neotropical (Gondwanan) origin, previously classified as the Echiniscus bigranulatus group; the pantropical and subtropical Kristenseniscusgen. nov. (the tessellatus group), characterized by a peculiar subdivision of dorsal plates; Claxtoniagen. nov. (the wendti group), with large and evident endocuticular pillars in the form of polygons; Nebularmisgen. nov. (the reticulatus group), with an elusive dorsal sculpturing; and Viridiscusgen. nov. (the viridis group), with body colour ranging from light green through brownish to even almost black. Additionally, we briefly address appendaged Echiniscus s.s. and divide the genus into several groups based on dorsal plate sculpturing and suggest that these could also represent separate supraspecific entities.

Revision of Testechiniscus Kristensen, 1987 (Heterotardigrada: Echiniscidae) refutes the polar-temperate distribution of the genus.

The family Echiniscidae comprises limno-terrestrial heterotardigrades with a strongly sclerotised dorsum, typically covered with plates. Among other members of the Echiniscus evolutionary line, the genus Testechiniscus Kristensen, 1987 stands out with well-developed ventral armature and polygonal sculpturing of the dorsal plates. It has alleged bipolar distribution (with satellite alpine records in the Holarctic). Thanks to fresh material from terra typica (Svalbard), we integratively redescribe (i.e. using light and electron microscopy imaging, morphometry, and molecular methods) the nominal species for the genus, Testechiniscus spitsbergensis (Scourfield, 1897). A comparison of the neotype series with a number of Holarctic records revealed morphological variability suggesting that the species may encompass several taxa, which, most likely, will be possible to delineate primarily with molecular tools. Moreover, based on material from Simien Mountains (Northern Ethiopia), we describe a new sibling subspecies, Testechiniscus spitsbergensis tropicalis ssp. nov. Extensive morphometric datasets are provided for the genus members for the first time. A new generic definition is proposed, embracing the two subspecies of T. spitsbergensis and T. laterculus (Schuster et al., 1980), but excluding two circum-Antarctic species, T. macronyx (Richters, 1907) and T. meridionalis (Murray, 1906). The later species are likely to be erected in the future as separate genera, and their autapomorphies are described here. In the light of our findings, the genus          Testechiniscus should be recognised as a native element of the Northern Hemisphere, with mainly circum-Arctic distribution and additional, insular alpine records from the Nearctic, Palearctic and Eastern Afrotropic.

An integrative redescription of Hypsibius dujardini (Doyère, 1840), the nominal taxon for Hypsibioidea (Tardigrada: Eutardigrada).

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 Mihelcic, 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.

Three Echiniscidae species (Tardigrada: Heterotardigrada) new to the Polish fauna, with the description of a new gonochoristic Bryodelphax Thulin, 1928.

During a faunistic survey in the Pieniny and Tatra Mountains three species of Echiniscidae new to Poland, including one new to science, were found. Bryodelphax instabilis sp. nov. is characterised by an instable number of ventral plates, deep faceting of the scapular plate, having dorsal plates covered either with pseudopores or pores, and striking sexual dimorphism. Two first records, namely Echiniscus militaris and E. spiniger, are rare taxa of upland-mountain character, which have already been reported from a few European countries since the original description at the beginning of the XXth century. The interspecific appendage length variability and development of pedal plates in the spinulosus group, to which E. spiniger belongs, and their taxonomic importance is discussed. Notes relating to sexual dimorphism within the newly recorded Pseudechiniscus facettalis, are also presented. Echiniscus testudo is reported from the Polish part of the Tatra Mountains for the first time. Succeeding findings confirm the high tardigrade α-diversity in the Polish mountain ranges. An amended key for Polish Heterotardigrada is provided.

Tardigrada and Rotifera from moss microhabitats on a disappearing Ugandan glacier, with the description of a new species of water bear.

Glaciers and ice sheets are a peculiar biome with characteristic abiotic and biotic components. Mountain glaciers are predicted to decrease their volume and even to melt away within a few decades. Despite the threat of a disappearing biome, the diversity and the role of microscopic animals as consumers at higher trophic levels in the glacial biome still remain largely unknown. In this study, we report data on tardigrades and rotifers found in glacial mosses on Mount Stanley, Uganda, and describe a new tardigrade species. Adropion afroglacialis sp. nov. differs from the most similar species by having granulation on the cuticle, absence of cuticular bars under the claws, and a different macroplacoid length sequence. We also provide a morphological diagnosis for another unknown tardigrade species of the genus Hypsibius. The rotifers belonged to the families Philodinidae and Habrotrochidae. In addition, we discuss the diversity of microinvertebrates and potential role of tardigrades and rotifers on mountain glaciers as top consumers. As for any organism living apparently exclusively in glacial habitats on tropical glaciers, their extinction in the near future is inevitable, possibly before we can even discover their existence.

Feeding strategies and competition between terrestrial Haemadipsa leeches (Euhirudinea: Arhynchobdellida) in Danum Valley rainforest (Borneo, Sabah).

Haemadipsid leeches are among the most successful terrestrial invertebrates in Bornean rainforests. They are very common ectoparasites of vertebrates, and their abundance has facilitated the conduction of numerous projects in the fields of ecology, zoogeography and taxonomy. We undertook research on two species inhabiting lowland dipterocarp forest, Haemadipsa picta Moore, 1929 and Haemadipsa subagilis (Moore, 1929), in order to address the following questions: (a) is there a difference in leech abundance between trails and off-trails?; (b) is ambush location dependent on specimen size or is species-specific?; (c) is intra- and interspecific competition limited by differences in foraging behaviours or vertical niche partitioning? Our results clearly show that H. picta is more abundant on trails than on off-trails and is vertically dispersed within the understory; the size of a specimen is strongly correlated with plant height. Haemadipsa subagilis was found not to exhibit such patterns. We suggest a possible lowering of interspecific competition between these species as a result of: (i) size-dependent dispersion of H. picta (together with reduction of intraspecific competition); and (ii) habitat specialisation of H. subagilis. Moreover, we provide new observations on their foraging behaviour.

Determinants and taxonomic consequences of extreme egg shell variability in Ramazzottius subanomalus (Biserov, 1985) (Tardigrada).

Nearly a half of known eutardigrade species lay ornamented eggs. The ornamentation is thought to provide attachment of the egg to the substrate and protection for the developing embryo, but from the taxonomic point of view chorion morphology may also provide key characters for species differentiation and identification, especially between closely related taxa. Nonetheless, despite the evolutionary and taxonomic importance of the egg shell, the determinants of its morphology are very poorly, if at all, understood. Here, we combine morphological, molecular and experimental approaches in an attempt to separate the genetic and environmental factors that shape egg chorion morphology in Ramazzottius subanomalus (Biserov, 1985). Our integrative study, based on a population of R. subanomalus isolated from a single moss sample, revealed (1) remarkable variation in egg shell morphology, but (2) relatively little variation in animal morphometric traits, and (3) genetic differentiation, expressed as two ITS-2 haplotypes, but no parallel polymorphism in COI. Although animals did not differ morphometrically between the haplotypes, eggs laid by haplotype 1 and 2 females exhibited highly statistically significant differences in all measured traits. The study demonstrates, for the first time, a correlation between phenotypic and genetic variability within a tardigrade species. The revealed congruence between genetic and morphological traits might be viewed as an example of incipient speciation that illustrates early evolutionary steps leading to species complexes that differ primarily in terms of egg shell morphology. Moreover, our data confirm the value of the ITS-2 fragment in distinguishing very closely related tardigrade lineages.