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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 nomeninquirendum, 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. [ABSTRACT FROM AUTHOR]

The phylum Tardigrada comprises ~ 1400 described species that inhabit a wide range of ecosystems throughout the globe. Tardigrades are generally considered taxonomically challenging due to a strongly limited number of taxonomically informative morphological traits and microscopic size. Of all tardigrade groups, the taxonomy of Milnesium Doyère, 1840 is particularly difficult because in comparison with most other eutardigrades, the genus lacks the taxonomically informative egg shell ornamentation and/or placoids in the muscle pharynx. Therefore, any new morphological traits that could be used in species delineation and identification are priceless. In this contribution, we review and evaluate taxonomic value of the dorsal cuticle morphology. Specifically, by means of experimental taxonomy, we demonstrate the first extreme case of ontogenetic variability in dorsal epicuticle sculpturing using a new species from Portugal, Milnesium decorumsp. nov. Furthermore, we verify the existence of dorsal gibbosities in Milnesium reticulatum Pilato, Binda, Lisi, 2002, the only species of the genus allegedly exhibiting these structures. Finally, we provide a diagnostic key to the Milnesium granulatum morphogroup. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

The Malay Archipelago comprises thousands of islands that house a variety of biomes, from tropical rainforests of Borneo, Sumatra and Celebes to the Lesser Sundas deciduous forests. In this paper, we present an extensive genetic and morphological dataset for the genus Kristenseniscus, demonstrating the presence of the walteri complex that contains several pseudocryptic species. One of them, Kristenseniscus exanthema sp. nov., is described from Ambon and Seram (the Moluccas). First genetic data are released for two potentially pantropical Echiniscus species: E. africanus Murray, 1907 and E. pusae Marcus, 1928. Furthermore, Echiniscus minutus sp. nov. (Sulawesi Tengah), exhibiting trunk spines and contrasting dorsal sculpturing that comprise both pores and epicuticular granules, is found to constitute a separate evolutionary lineage within the Echiniscus spinulosus morphogroup (epicuticular granules are typically absent in the spinulosus group). Based on the phylogenetic reconstructions involving ca. 80 spp. of echiniscids representing the Echiniscus evolutionary line, we hypothesise that the tropical and subtropical regions served as the main radiation zone for this heterotardigrade clade. Finally, Pseudechiniscus (Meridioniscus) celebesiensis sp. nov. (Sulawesi Tengah) is a characteristic species with an apomorphic lack of claw spurs and intricate dorsal cuticular sculpturing. Our findings suggest that the tropical areas of the globe harbour a phylogenetically important but yet mostly uncovered portion of Earth’s tardigrade diversity. [ABSTRACT FROM AUTHOR]

Inhibition of return (IOR) is the effect of slower responses to validly than invalidly cued targets. The discovery of IOR raised controversy as to whether it has two "flavors", i.e., attentional/perceptual and motoric, or whether it is a homogeneous visual-motor phenomenon that should be understood in terms of the preparation of different effectors (mainly eye movement). Since manipulation of fixation offset (0 and 200 ms gap) is believed to affect the latency of saccades, we measured its influence on saccadic and manual IOR with a simple keypress response when eye movements were forbidden. In the two experiments which we carried out, the fixation offset decreased IOR in both the saccadic and the manual conditions. The results suggest the limitations of the attentional hypothesis, which assumes that manual IOR is independent of the motoric component; they are also in line with the tenets of the oculomotor hypothesis of IOR. [ABSTRACT FROM AUTHOR]

Climate change is affecting biodiversity, but proximate drivers remain poorly understood. Here, we examine how experimental heatwaves impact on reproduction in an insect system. Male sensitivity to heat is recognised in endotherms, but ectotherms have received limited attention, despite comprising most of biodiversity and being more influenced by temperature variation. Using a flour beetle model system, we find that heatwave conditions (5 to 7 °C above optimum for 5 days) damaged male, but not female, reproduction. Heatwaves reduce male fertility and sperm competitiveness, and successive heatwaves almost sterilise males. Heatwaves reduce sperm production, viability, and migration through the female. Inseminated sperm in female storage are also damaged by heatwaves. Finally, we discover transgenerational impacts, with reduced reproductive potential and lifespan of offspring when fathered by males, or sperm, that had experienced heatwaves. This male reproductive damage under heatwave conditions provides one potential driver behind biodiversity declines and contractions through global warming. [ABSTRACT FROM AUTHOR]

Tardigrada are a group of microscopic metazoans that inhabit a variety of ecosystems throughout the world, including polar regions, where they are a constant element of microfauna with densities exceeding hundreds of individuals per gram of dry plant material. However, despite a long history of research and their ubiquity in tundra ecosystems, the majority of tardigrade species have limited and outdated diagnoses. One such example is Pilatobius recamieri, a common tardigrade that is widely distributed in the Arctic. The aim of this study is to redescribe this species using new material from the type locality and tools of integrative taxonomy, viz. by combining classical imaging and morphometry by light microscopy and scanning electron microscopy imaging with DNA sequencing of four markers with various mutation rates: three nuclear (18S rRNA, 28S rRNA, and ITS-2) and one mitochondrial (COI). The sequences of the three latter markers are also the first to be presented for the genus Pilatobius. This study therefore provides the first necessary step towards the verification of the geographic range of P. recamieri, which is currently assumed to be very broad. A detailed comparison of P. recamieri with Pilatobius secchii (Bertolani and Rebecchi, 1996) from Italy revealed no morphological or morphometric differences between the two species, thus we designate P. secchii as a nomen inquirendum until molecular data for the taxon become available. Finally, we propose to replace the term 'lunula' in the superfamilies Hypsibioidea and Isohypsibioidea with the more appropriate 'pseudolunula' to differentiate it from the true lunula in other parachelans. [ABSTRACT FROM AUTHOR]

The acute toxicity of ammonia on Thulinius ruffoi (Bertolani, 1981), a eutardigrade isolated from a small waste water treatment plant (WWTP) in Poland, was estimated. Our results show that no active individuals survived a 24 h exposure to solutions equal to or higher than 125 mg/L of total ammonia nitrogen (NH-N + NH-N), which, under the conditions in our experiment, was equivalent to 1.17 mg/L of un-ionised ammonia (NH). The LC concentration of total ammonia nitrogen was equal to 52 mg/L (or 0.65 mg/L un-ionised ammonia). Given that the norms for the concentration of ammonia in treated waters leaving WWTPs are usually several times lower than the LC for T. ruffoi, this species does not seem to be a good bioindicator candidate for WWTPs. In this paper we also note that various ecotoxicological studies use different methodological approaches and we suggest that a more uniform methodology may aid interspecific comparisons of LC values. [ABSTRACT FROM AUTHOR]

Reproduction through sex carries substantial costs, mainly because only half of sexual adults produce offspring. It has been theorized that these costs could be countered if sex allows sexual selection to clear the universal fitness constraint of mutation load. Under sexual selection, competition between (usually) males and mate choice by (usually) females create important intraspecific filters for reproductive success, so that only a subset of males gains paternity. If reproductive success under sexual selection is dependent on individual condition, which is contingent to mutation load, then sexually selected filtering through 'genic capture' could offset the costs of sex because it provides genetic benefits to populations. Here we test this theory experimentally by comparing whether populations with histories of strong versus weak sexual selection purge mutation load and resist extinction differently. After evolving replicate populations of the flour beetle Tribolium castaneum for 6 to 7 years under conditions that differed solely in the strengths of sexual selection, we revealed mutation load using inbreeding. Lineages from populations that had previously experienced strong sexual selection were resilient to extinction and maintained fitness under inbreeding, with some families continuing to survive after 20 generations of sib × sib mating. By contrast, lineages derived from populations that experienced weak or non-existent sexual selection showed rapid fitness declines under inbreeding, and all were extinct after generation 10. Multiple mutations across the genome with individually small effects can be difficult to clear, yet sum to a significant fitness load; our findings reveal that sexual selection reduces this load, improving population viability in the face of genetic stress. [ABSTRACT FROM AUTHOR]

(1) During the Arctic summer, little auks ( Alle alle) deposit considerable amounts of guano on land. Ecosystems subsidised in nutrients are known to hold greater biodiversity and to produce grander biomass of plants and animals compared with areas where seabirds do not nest. (2) The aim of this study was to look into the relationship between guano fertilisation and body size of invertebrates inhabiting tundra. (3) The specimens of Macrobiotus islandicus islandicus Richters, 1904, a tardigrade dwelling in mosses and lichens of the Arctic, from six different populations from Spitsbergen (Hornsund fjord) were measured. Tardigrades were collected from areas different in terms of seabird guano effects on the tundra ecosystem. An overall body size index for tardigrades was calculated using a principal component analysis. (4) Here, we show that the body size of M. i. islandicus is larger in vicinities of the little auk colonies than in areas devoid of bird nesting sites. (5) Given that fitness of many invertebrates is positively correlated with their condition, our study underlines the ecological importance of a side effect of seabirds biology-the transfer of nutrients from the sea to the land. [ABSTRACT FROM AUTHOR]

Polyandrous mating is extremely common, yet for many species the evolutionary significance is not fully resolved. In order to understand the evolution of mating systems, it is crucial that we investigate the adaptive consequences across many facets of reproduction. We performed experimental evolution with the naturally polygamous flour beetle Tribolium castaneum subjected to either polyandry or enforced monogamy, creating contrasting selection regimes associated with the presence or absence of sexual selection. After 36 generations, we investigated male and female adaptations by mating beetles with an unselected tester strain to exclude potential effects of male-female coevolution. Reproductive success of focal monogamous and polyandrous beetles from each sex was assessed in separate single male and multiple male experiments emulating the different selection backgrounds. Males and females from the polyandrous regime had more offspring in the experiments with multiple males present than monogamous counterparts. However, in single male experiments, neither females nor males differed between selection regimes. Subsequent mating trials with multiple males suggested that adaptations to polyandry in both sexes provide benefits when choice and competition were allowed to take place. Polyandrous females delayed the first copulation when given a choice of males and polyandrous males were quicker to achieve copulation when facing competition. In conclusion, we show that the expected benefits of evolutionary adaptation to polyandry in T. castaneum depended on the availability of multiple mates. This context-dependent effect, which concerned both sexes, highlights the importance of realistic competition and choice experiments. [ABSTRACT FROM AUTHOR]

Despite a century long history of research, tardigrade fauna of the Svalbard Archipelago remains poorly known. In order to deepen our knowledge of tardigrade biodiversity in the Arctic, we collected forty-one moss and lichen samples from the Revdalen and on the south-east slopes of the Rotjesfjellet (Spitsbergen, Svalbard Archipelago) in June 2010. In these samples, twenty-five tardigrade species were found, including two new for science: Bryodelphax parvuspolaris sp. nov. and Isohypsibius coulsoni sp. nov. B. parvuspolaris sp. nov. belongs to the weglarskae group but differs from all other species of the group by a unique configuration of ventral plates. I. coulsoni sp. nov. differs from the most similar species I. ceciliae Pilato and Binda, 1987 mainly by the absence of ventral sculpture. Two additional species, Milnesium asiaticum Tumanov, 2006 and Diphascon ( Adropion) prorsirostre Thulin, 1928, are recorded from the Svalbard Archipelago for the first time. [ABSTRACT FROM AUTHOR]

rare marine tardigrade Halobiotus crispae Kristensen, 1982 was found abundant in intertidal flats at Josephbukta, Bellsund fiord, West Spitsbergen. This is the first record of this species from the Svalbard Archipelago. So far, the species has been found only in seven other localities in the northern hemisphere (between 56° and 69°N). The present record greatly extends the known geographical range to the north and also indicates a much wider distribution of the species than formerly reported. Moreover, a potential indicative character of this species and of the study area in monitoring climatic changes is discussed. [ABSTRACT FROM AUTHOR]

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]