Search

The family Macrobiotidae is one of the most speciose and diverse groups among tardigrades. Although there have been attempts to reconstruct the phylogeny of this family, the evolutionary relationships within Macrobiotidae are only superficially determined as available genetic data cover only a small fraction of this vast group. Here, we present the first extensive molecular phylogeny of the family based on four molecular markers (18S rRNA, 28Sr RNA, ITS-2 and COI) associated with detailed morphological data for the majority of taxa. The phylogenetic analysis includes nearly two hundred sequences representing more than sixty species, including sixteen taxa that have never been sequenced and/or analysed phylogenetically before. Our results recovered a new monophyletic group, comprising Macrobiotus spectabilis Thulin, 1928 and Macrobiotus grandis Richters, 1911, for which we erect a new genus, Sisubiotusgen. nov., to accommodate its evolutionary distinctiveness. The largest, so far, dataset for the family Macrobiotidae showed that the genus Xerobiotus is nested within the clade representing the genus Macrobiotus deeper than it was earlier assumed, therefore we propose to suppress Xerobiotus and transfer its species to Macrobiotus. Moreover, mapping key morphological traits onto macrobiotid phylogeny exposed complex evolution of phenotypes within the Macrobiotus hufelandi group, i.e. Macrobiotus s.s. Finally, our findings enabled a detailed revision and discussion on species compositions of the most ubiquitous tardigrade genera, species groups and species complexes, which resulted in changes of taxonomic statuses of a number of macrobiotid species. All this contributes to the reconstruction of the morphological evolution within Macrobiotidae., (Copyright © 2020 The Author(s). Published by Elsevier Inc. All rights reserved.)

Tardigrade research in Japan dates back over 100 years, and to date, 167 species of this ecdysozoan phylum have been reported from the country. Of these species, the Macrobiotus hufelandi complex has been represented only by the nominal taxon of this group, Macrobiotus hufelandi. In this article, a new species of the hufelandi group from Japan, Macrobiotus shonaicus sp. nov., is described using integrative taxonomy. In addition to the detailed morphological and morphometric data, obtained using phase contrast light microscopy (PCM) and scanning electron microscopy (SEM), we provide DNA sequences of four molecular markers (both nuclear and mitochondrial). The new species belongs to the persimilis subgroup and is most similar to M. anemone from USA, M. naskreckii from Mozambique, and M. patagonicus from Argentina, but it can be easily distinguished from these species by the presence of thin flexible filaments on terminal discs of the egg process. By the latter character, the new species is most similar to M. paulinae and M. polypiformis, but it can be easily distinguished from them by having a solid egg surface between egg processes (i.e., without pores or reticulum). A phylogenetic analysis of available DNA sequences of the COI marker for the hufelandi group revealed that the new species clusters with the two other species that exhibit filaments on egg process discs (M. paulinae and M. polypiformis) and with two species that have entire egg processes modified into filaments (M. kristenseni and M. scoticus). All five species form a clade distinct from all other sequenced species of the hufelandi group with typical mushroom- or inverted goblet-shaped egg processes, which may suggest that the ancestor of the five species with atypical egg processes had a mutation allowing derivations from the mushroom or inverted chalice-like shape of egg processes.

A new freshwater tardigrade species, Pseudobiotus spinifer sp. nov., is described from the sand bottom of Nakdong River, South Korea. The new species is most similar to Pseudobiotus vladimiri from Biwa Lake, Japan, but differs from it by having small accessory points ending in the center of the primary branches of all claws, relatively longer claws and macroplacoids, and well developed cuticular spines/spicules over most of the dorsal and lateral surfaces of the body. Pseudobiotus vladimiri is redescribed on the basis of the holotype. A revised key to the species of the genus Pseudobiotus is also given.

Tardigrade taxonomy is most often hindered by prevalent outdated species descriptions, lack of integrative redescriptions, scarce genetic information, and fragmentary sampling. Here, we diagnose the problems obscuring phylogenetic inference and the taxonomy of the cosmopolitan family Ramazzottiidae. We carried out the most extensive phylogenetic analysis of this family to date, with a considerable influx of new genetic data from poorly sampled regions of the world. We found two new distinct evolutionary lineages defined by distinct morphological traits (dorsal cuticular sculpturing and/or body shape), characterized by restricted geographic distributions, and we delineated them as new species complexes: the Neotropical Ramazzottius baumanni complex and the Afro-Oriental Ramazzottius szeptyckii complex. Nevertheless, we could not confidently ascertain the taxonomic status of both complexes due to: (i) the current state of the ramazzottiid taxonomy, with outdated and/or imprecise species descriptions with heterogeneous terminology and, in many cases, without accounting for intraspecific variation; (ii) the missing genetic information for key taxa; and (iii) the possible lack of monophyly of Cryoconicus and Ramazzottius as suggested by our results. In addition to diagnosing the problems of ramazzottiid phylogeny and systematics, we also propose possible solutions that could accelerate the progress in our understanding of the evolution of this group. [ABSTRACT FROM AUTHOR]

Numerous evolutionary lineages representing an armoured, limno-terrestrial family Echiniscidae are restricted to high mountainous or polar localities, exhibiting clear cold stenothermic preferences. One such group is the genus Claxtonia , with its type species Claxtonia wendti , originally described from Svalbard and later on reported globally. In this paper, we re-describe this taxon by establishing the neotype from Spitsbergen. We sequenced five genetic markers (18S rRNA, 28S rRNA, ITS-1, ITS-2, and COI) for multiple populations of Claxtonia , including C. mauccii , C. molluscorum , C. wendti , and several potentially new species, collected in the Alps, Andes, Carpathians, Scotland, Iceland, Southern Appalachians, Spitsbergen, Scandinavian Peninsula, Borneo, and the West Indies. The relationships between these species are elucidated thanks to the phylogenetic reconstructions based on nuclear loci. Various Claxtonia species frequently co-occur in mosses and lichens, which hampers delineating their probable geographic ranges. All historical records of C. wendti are questioned in the light of our revision. Delineation of new species within the genus is nipped in the bud due to numerous problems with original species descriptions, unknown intraspecific (including ontogenetic) variability, and the lack of DNA barcodes. Finally, it is hypothesized that Claxtonia potentially comprises two lineages: a cold-stenothermic one and a thermophilic subtropical–tropical one. [ABSTRACT FROM AUTHOR]

The genus Barbaria , recently established to accommodate the former Echiniscus bigranulatus group, is a tardigrade group emblematic for the South American tardigrade fauna. This unappendaged echiniscid lineage is widely recognized for the so-called 'double' sculpturing composed of endocuticular pillars and pseudopores or pores in the dorsal cuticle. The phylogenetic relationships in the genus have so far been completely unknown, but the discovery of two new species (B. paucigranulata sp. nov. and B. weglarskae sp. nov.), together with new genetic data for further six species (B. bigranulata , B. charrua comb. nov. , B. danieli , B. jenningsi , B. madonnae and B. ollantaytamboensis), create an opportunity not only to uncover phyletic relationships, but also to reconstruct morphological evolution in the genus. To achieve this, we sequenced five genetic markers (18S rRNA, 28S rRNA, ITS1, ITS2, COI) for multiple populations of eight species of Barbaria (two-thirds of all known species) collected in Alabama (USA), Argentina and the Antarctic, and we analysed them in tandem with detailed morphological data. Our phylogentic analysis and the reconstruction of evolution of morphological traits suggests that the ancestor of the genus inhabited the Neotropics, and it was morphologically most similar to B. bigranulata. We also analyse literature records of Barbaria and conclude that the genus is most likely limited to the Neotropics, Antarctica and southern parts of the Nearctic. The findings are discussed in the context of the phylogeny of the Echiniscus evolutionary line. [ABSTRACT FROM AUTHOR]

The document titled "15th International Symposium on Tardigrada – Section I: Biodiversity, Taxonomy, Biogeography, Phylogeny & Evolution" discusses the topic of tardigrades, also known as water bears. The document includes an illustration of major tardigrade lineages climbing a phylogenetic tree, representing our current understanding of tardigrade phylogeny. The authors, Łukasz Michalczyk and Kamil Janelt, declare no conflict of interest. The document is authored by researchers from the Department of Invertebrate Evolution at the Institute of Zoology and Biomedical Research, Faculty of Biology, Jagiellonian University in Kraków, Poland. [Extracted from the article]

Tardigrades constitute a phylum of cosmopolitan, microscopic invertebrates, capable of withstanding harsh environmental conditions thanks to cryptobiosis. These tiny animals are taxonomically challenging due to the limited number of meaningful morphological traits and morphological stasis. One of the most extreme examples is the order Apochela, in which the morphology is even more simplified and conserved than in the majority of other tardigrades. Despite this, apochelans are currently popular among researchers, as almost half of the 45 known species have been described only in the last five years. During the examination of samples from Kazakhstan we encountered specimens belonging to the genus Milnesium Doyère, 1840, certainly representing a species new to science. Milnesium pentapapillatum sp. nov. is described in the integrative taxonomy framework, which includes morphometrics, imaging both in light and scanning electron microscopy, ontogenetic variability testing, and DNA sequencing of four standard markers, which allowed for pinpointing its phylogenetic position. Although this species, with a [3-3]-[3-3] claw configuration and smooth cuticle, represents the most common morphotype found in Milnesium , it can be easily differentiated from all other congeners by the presence of five (instead of six) peribuccal papillae. Given that to date all apochelans were characterised by six peribuccal papillae, the discovery of M. pentapapillatum sp. nov. entails a change in the diagnosis of the order Apochela, family Milnesiidae, and the genus Milnesium. Moreover, we propose to suppress the recently erected class Apotardigrada and restore the suppressed order Parachela, since grouping the Apochela and Parachela under Eutardigrada reflects tardigrade phylogeny. [ABSTRACT FROM AUTHOR]

The family Hypsibiidae comprises four subfamilies, among which Itaquasconinae are the most abundant in genera. Although being morphologically diverse and cosmopolitan, itaquasconins are scarcely reported and their taxonomy is entangled. Here, we present a phylogeny based on three DNA markers and morphological clues obtained from a detailed scanning electron microscopy analysis of bucco‐pharyngeal apparatuses of 25 species representing seven genera. Our study revealed that (a) the polyphyletic Adropion consists of at least three evolutionary lineages, with Guidettion gen. nov. being the sister group of all remaining itaquasconins; (b) stylet supports were independently lost in Astatumen and Insulobius gen. nov.; (c) pharyngeal tube annulation can be divided into two general types, simple and complex, and is stable at the genus level, but being homoplasious (similarly to the apophyses for the insertion of stylet muscles and the shape of the claw bases), it is not useful in inferring phyletic relationships within the subfamily; and (d) Astatumen and Platicrista contain numerous similar species, challenging the current taxonomic classification; moreover, (e) three new taxa are proposed to accommodate the newly recognised lineages: Guidettion gen. nov. (the former Adropion prorsirostre group), Insulobius gen. nov. (for a new Malayan species with a mixture of Astatumen and Itaquascon traits), and Raribius gen. nov. (former Itaquascon spp. with an elongated and narrowed pharyngeal tube terminating with a spherical pharynx). The simple type of annulation is synapomorphic to Hypsibiidae, with single reversals to the uniform buccal tube in Parascon (Itaquasconinae) and in Hypsibiinae. The pharyngeal tube evolved convergently several times, and is of different character in classes Heterotardigrada and Eutardigrada. [ABSTRACT FROM AUTHOR]

The largest apochelan genus, Milnesium , is increasingly gaining attention. However, our knowledge on the phylogeny of Milnesiidae is strongly limited. In fact, so far, apochelan species usually served only as outgroups in phylogenetic analyses of other tardigrade groups. Thus, the goal of this study was to reconstruct the first multilocus molecular phylogeny of Milnesium and test several hypotheses on the evolution and dispersion of species in the genus by mapping taxonomically important morphological traits, reproductive mode and geographic distribution onto the phylogenetic tree. The phylogeny was inferred from four DNA markers (nuclear 18S rRNA, 28S rRNA, ITS2 and mitochondrial COI) and for 34 populations representing 25 species collected across the globe. Phylogenetic analysis revealed distinct molecular lineages within the genus, but rather than by morphological characters, species clustered by zoogeographic zones. In fact, no obvious patterns of phenotypic traits or reproductive mode were observed, which may indicate high levels of convergent evolution in morphology throughout the genus. The lack of congruence between morphology and phylogeny did not allow splitting this largest apochelan genus into separate taxonomic units such as genera or subgenera. On the other hand, the correlation between phylogeny and geography may suggest limited dispersal of at least some Milnesium species. [ABSTRACT FROM AUTHOR]

Incomplete descriptions of nominal taxa are one of the most significant obstacles in modern taxonomy, including the taxonomy of Tardigrada. Another major problem in tardigrade systematics is the lack of tests for the reliability of genetic markers in species delineation. Here, we employ an integrative taxonomy approach to redescribe the nominal taxon for the P. areolatus complex, Paramacrobiotus areolatus. Moreover, we obtained multilocus DNA sequences for another 16 populations representing 9–12 Paramacrobiotus species collected from Europe, North America, Africa and Australia, enabling us to reconstruct the most extensive phylogeny of the genus to date. The identification of a pair of potentially cryptic dioecious P. areolatus complex species with divergent genetic distances in ITS2 (1.4%) and COI (13.8%) provided an opportunity to test the biological species concept for the first time in the history of tardigrade taxonomy. Intra- and interpopulation crosses did not differ in reproductive success in terms of F1 offspring. However, because of the low F1 family sizes, we were unfortunately unable to test F1 hybrid fertility. Although our results are only partially conclusive, they offer a baseline not only for further taxonomic and phylogenetic research on the areolatus complex, but also for studies on species delineation in tardigrades in general. [ABSTRACT FROM AUTHOR]

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). Intraspecific, interpopulation variability of a tardigrade Milnesium tardigradum was investigated by means of integrative taxonomy, including morphology (light and scanning electron microscopy), morphometry, and barcoding of a nuclear (ITS‐2) and a mitochondrial marker (COI). The nine analysed European populations revealed low to moderate variability in morphology and in ITS‐2 (≤4.0%). In contrast, COI exhibited low to surprisingly high divergence (≤11.4%), questioning the possibility of setting a universal barcoding threshold for tardigrade species delineation. Additionally, novel, taxonomically important traits for M. tardigradum are described. [ABSTRACT FROM AUTHOR]

Species constituting the family Echiniscidae are highly derived, armoured and inhabit terrestrial habitats, in contrast to other heterotardigrades that are predominantly marine. The genus Echiniscus C.A.S. Schultze, 1840, nominal for the family Echiniscidae, is currently the most speciose tardigrade genus. However, the great morphological variability, in comparison with other heterotardigrade genera, suggests the polyphyletic character of the genus. Here, we analyse new specimens of Echiniscus pseudelegans Séméria, 1994 collected in Japan and conclude that the species as well as two other related taxa, E. elegans Richters, 1907 and E. latifasciatus Dudichev and Biserov, 2000, represent a new genus, Stellariscus , gen. nov. The new genus is characterised by a mixture of peculiar morphological apomorphies: black eyes, star-like dorsal plate sculpturing, no trunk appendages (only cephalic cirri present), two types of ventral plates, and striking sexual dimorphism in both qualitative and quantitative traits. Morphological phylogeny of the family Echiniscidae suggests a close affinity between Stellariscus , Hypechiniscus Thulin, 1928 and Pseudechiniscus Thulin, 1911. The polyphyletic status of both Echiniscus and Testechiniscus Kristensen, 1987 is also inferred. The taxonomic significance of ventral armature in echiniscid phylogeny and taxonomy is discussed. Many tardigrades can be easily found in garden mosses, but the members of the family Echiniscidae are quite rare, thus poorly known. This paper presents morphological clues for an artificial character of the largest echiniscid genus Echiniscus , by erecting new genus restricted to Asia. Therefore, by demonstrating the value of cuticular pattern in the systematics, further changes can be made to elucidate echiniscid classification. [ABSTRACT FROM AUTHOR]

Mesocristais a Holarctic tardigrade genus currently classified within the subfamily Itaquasconinae (Eutardigrada: Parachela: Hypsibiidae). The position of the genus has been so far inferred solely on the basis of light microscopy observations. Here, we present the first-ever scanning electron microscopy images of a dissected bucco-pharyngeal apparatus and DNA sequences forMesocristafrom Spitsbergen, mainland Norway and Poland. The new data allowed an integrative redescription of the nominalM. spitzbergensisfrom thelocus typicusand uncovered a new species,M. revelata, from the European localities. The two species differ phenotypically by the oral cavity armature and claw morphology as well as by a number of morphometric traits, and they are also distinct genetically in all sequenced DNA markers, three nuclear (18S rRNA, 28S rRNA, ITS-2) and one mitochondrial (COI). Both molecular and morphometric data presented in this paper suggest that previous records ofM. spitzbergensisshould be treated with caution, since most probably some of them signify differentMesocristaspecies. Our phylogenetic analysis confirmed thatMesocristais indeed a member of the Itaquasconinae. We also found that the most closely related, among genera for which there are molecular data available, isAdropion, from whichMesocristadiffers by the shape of the apophyses for insertion of stylet muscles and the width of the bucco-pharyngeal tube. Finally, we advocate thatDiphascon marcusi, a rare species hypothesised to represent eitherMesocristaorAdropion, should be transferred to the latter genus and that the species should be redescribed to confirm its validity and systematic position. http://zoobank.org/urn:lsid:zoobank.org:pub:6515D58D-8469-4854-8687-177232112BDA [ABSTRACT FROM PUBLISHER]

The family Richtersiidae, although established recently with the use of phylogenetic methods, was considered potentially paraphyletic at the time of its erection. Until now, the family comprised four genera, Richtersius, Diaforobiotus, Adorybiotus and a newly erected genus Crenubiotus. However, the genetic characterisation for the latter two genera was very limited or absent. To address concerns about the phylogenetic affinity of these two genera, we present a multilocus phylogeny of the families Richtersiidae and Murrayidae based on four molecular markers (18S rRNA, 28S rRNA, ITS-2 and COI). Our results show a distinct evolutionary lineage composed of Adorybiotus and Crenubiotus, which is sister to Murrayidae. In order to accommodate the phylogenetic and morphological distinctiveness of this lineage, we erect a new family, Adorybiotidae fam. nov. The new taxon differs morphologically from other families in the superfamily Macrobiotoidea by a unique combination of traits: (1) the presence of tubercles/cushions with aggregations of microgranules on their surfaces present on all legs and on the dorso-caudal cuticle, (2) a system of internal septa in claws, and (3) buccal apparatus morphology. Moreover, in order to stabilise the taxonomy and nomenclature in the genus Crenubiotus, we redescribe its type species, Crenubiotus crenulatus, by means of integrative taxonomy and designate a new neotype based on a population from the original terra typica. [ABSTRACT FROM AUTHOR]

Isohypsibioidea are most likely the most basally branching evolutionary lineage of eutardigrades. Despite being second largest eutardigrade order, phylogenetic relationships and systematics within this group remain largely unresolved. Broad taxon sampling, especially within one of the most speciose tardigrade genera, Isohypsibius Thulin, 1928, and application of both comparative morphological methods (light contrast and scanning electron microscopy imaging of external morphology and buccal apparatuses) and phylogenetic framework (18S + 28S rRNA sequences) resulted in the most comprehensive study devoted to this order so far. Two new families are erected from the currently recognised family Isohypsibiidae: Doryphoribiidae fam. nov., comprising all aquatic isohypsibioids and some terrestrial isohypsibioid taxa equipped with the ventral lamina; and Halobiotidae fam. nov., secondarily marine eutardigrades with unique adaptations to sea environment. We also split Isohypsibius into four genera to accommodate phylogenetic, morphological and ecological variation within the genus: terrestrial Isohypsibius s.s. (Isohypsibiidae), with smooth or sculptured cuticle but without gibbosities; terrestrial Dianea gen. nov. (Isohypsibiidae), with small and pointy gibbosities; terrestrial Ursulinius gen. nov. (Isohypsibiidae), with large and rounded gibbosities; and aquatic Grevenius gen. nov. (Doryphoribiidae fam. nov.), typically with rough cuticle and claws with branches of very similar heigths. Claw morphology is reviewed and, for the first time, shown to encompass a number of morphotypes that correlate with clades recovered in the molecular analysis. The anatomy of pharynx and cuticle are also shown to be of high value in distinguishing supraspecific taxa in Isohypsibioidea. Taxonomy of all isohypsibioid families and genera is discussed, with special emphasis on the newly erected entities. Finally, a dychotomous diagnostic key to all currently recognised isohypsibioid families and genera is provided. [ABSTRACT FROM AUTHOR]