The Central American Isthmus has had profound effects on the evolution and distribution of larger marine organisms, but the impacts on meiobenthic animals have received little attention. Tardigrades are microscopic metazoan lobopodians that are ubiquitous in benthic communities worldwide, but little is known about marine tardigrades in Central America. Only two marine tardigrades have been identified to species level in all of Central America, and these came from a single sample of barnacles from El Salvador collected in 1953. Additionally, multiple haplotypes of Echiniscoides were reported from Costa Rica and Panama, but species names were not assigned. Here we report an extensive survey of both intertidal and subtidal marine tardigrades from both the Caribbean and Pacific coasts of Costa Rica. We found 701 individuals in 65 samples. There was a total of 19 taxa, including seven species with a sufficient quality and quantity of specimens to describe here as new to science. The new species are Archechiniscus murilloi sp. nov., Batillipes homocercus sp. nov., Batillipes ichthyocercus sp. nov., Echiniscoides costaricensis sp. nov., Echiniscoides ritavargasae sp. nov., Styraconyx vargasi sp. nov. and Tanarctus breedyae sp. nov. Ten taxa were found only in the Caribbean, six were found only in the Pacific, and three were found on both coasts. We discuss the three species with trans-isthmus distributions and note two additional candidate geminate species pairs that warrant further investigation.

The main subject of my papers has been the phylum Tardigrada, with particular emphasis on the Antarctic fauna. While this is one of the ‘lesser known phyla’ the group forms an important element of Antarctic non-marine ecosystems. My work has centred on four interrelated themes: i) taxonomy, predominantly Tardigrada and with an Antarctic bias though including wider global species and high taxon groupings (I have also worked on other taxa such as Fungi-imperfecti and freshwater invertebrates); ii) biodiversity of tardigrades, particularly Antarctic habitats; iii) data-based studies based on the terrestrial Tardigrada and Antarctic freshwater crustaceans; and iv) biogeographic analysis of these databases. The thesis presented consists of papers published in major, peer-reviewed journals, along with book chapters published and in press. Several of my papers have been cited 10-30 times according to the ISI Web of Science citation system. Highlights of my work include being the first to publish a paper on the global terrestrial biogeography of a phylum and showing the extant limno-terrestrial tardigrade fauna reflect the early tectonic break-up of the Laurasia, West and East Gondwana super continents between 65 and 135 million years ago.

Background. The phylum Tardigrada consists of over 1,300 species that inhabit terrestrial, freshwater and marine environments throughout the world. In terrestrial habitats they live primarily in mosses, lichens, leaf litter and soil, whereas tardigrades in freshwater and marine environments are mainly found in sediments and on aquatic plants. More than 65 species have been previously reported in the state of Tennessee, USA. Methods. Tardigrades present in moss cushions (Grimmia sp.) collected from a xerothermic habitat on the East Tennessee State University campus, Johnson City, TN, USA, were extracted, mounted on slides, identified, and counted. Additional samples of fresh dried moss were used for integrative analyses, including morphological analysis with phase contrast (PCM) and scanning electron microscopy (SEM), as well as molecular analyses of COI, 18S rRNA, 28S rRNA, and ITS-2 of the Macrobiotus and Milnesium species. Results. Five species were found, including two species new to science: Viridiscus miraviridis sp. nov. and Macrobiotus basiatus sp. nov. Viridiscus miraviridis sp. nov. differs from other members of the genus mainly by having a different type of dorsal cuticle and some other, more subtle, morphometric characters. In addition to the two new species, Viridiscus perviridis and Viridiscus viridissimus were present, and males of Vir. viridissimus were found for the first time, the first record of males in the genus Viridiscus. Macrobiotus basiatus sp. nov. is most similar to Macrobiotus nelsonae, but it differs from Mac. nelsonae mainly by the stylet supports being situated in a more anterior position, shorter and narrower egg processes, and a smaller number of areoles around the egg processes. Moreover, the identification of Milnesium inceptum was confirmed as the first record for the USA by analysis of COI.

Although many estimates of species numbers have been attempted using various techniques, many smaller phyla remain poorly known without such estimates. For most of these it is unclear if they are species-poor or just poorly studied. The phylum Tardigrada is one of these phyla. Specialists have created a regularly updated checklist for the known tardigrade species, which as of 15 July 2013 listed 1190 taxa (species and subspecies). Of these, 1008 are limnoterrestrial and 182 are marine. These were the most up-to-date data at the time of our analysis. As species accumulation curves show little sign of levelling out, they do not provide a useful tool for estimating global tardigrade diversity from existing species numbers. A new technique has recently been developed that uses the more complete knowledge of higher taxonomic levels to estimate the asymptotic number of species. We applied this technique to limnoterrestrial and marine tardigrades. We estimate that the global total for limnoterrestrial tardigrades is 1145 (upper 95% CI = 2101), and the global total for marine tardigrades is 936 (upper 95% CI = 1803). This yields 87% completeness for our knowledge of limnoterrestrial tardigrades, and only 19% completeness for our knowledge of marine tardigrades. Thus, although many more marine species remain to be discovered, it appears that tardigrades are both poorly studied and relatively species poor.

This monograph describes the global records of marine water bears (Phylum Tardigrada). We provide a comprehensive list of marine tardigrades recorded from around the world, providing an up-to-date taxonomy and a complete bibliography accompanied by geographic co-ordinates, habitat, substrate and biogeographic comments. A link is provided to an on-line interactive map where all occurrences for each species are shown. In total we list 197 taxa and their 2240 records from 39 oceans and seas and 18 Major Fishing Areas (FAO). It is hoped this work will serve as a reference point and background for further zoogeographic and taxonomic studies on marine tardigrades.

A new species, Ramazzottius belubellus, is described from a single lichen sample collected in the Great Smoky Mountains National Park in western North Carolina, U.S.A. The new species is easily distinguishable from all other members of the genus Ramazzottius by the presence of long dorsal, sharp triangular spines not arranged in transverse bands as opposed to small tubercles arranged in bands, or a thin reticulum, or a smooth cuticle present in all other described species of the genus. The new species could be confused with R. baumanni but differs from it by the presence of sharp triangular spines on the dorsal side of the body instead of flat, hemispherical tubercles.

One of the largest inventories of tardigrades ever conducted occurred from 2000 2010 in the Great Smoky Mountains National Park, USA. Over 16,000 specimens were catalogued, 85 species were identified, 11 species new to science were described, and 16 other possible new species await further study. More than 20 papers have resulted from the GSMNP tardigrade inventory, making the Smokies the most thoroughly studied area in North America for tardigrades. Several species lists have been published over this 20-year period, but many taxonomic revisions and new identifications have led to significant changes to the list. Biogeographical studies citing species records from earlier studies could yield serious errors. Here we update the species list from the Smokies to accommodate the many recent changes in tardigrade taxonomy, we re-Analyze some species in light of delineations of cryptic species groups that have occurred recently via integrative taxonomy, and we provide a table of all synonyms that have been used in previous publications. We also make available, for the first time, the Smokies tardigrade database, complete with all locations, elevations, and substrates.

This is the first survey of subtidal marine tardigrades from the Bahamas, and we compare our results with earlier studies of Bahamian intertidal habitats. In 2011 and 2014 we collected 60 subtidal sand samples from Bimini, the Berry Islands, New Providence, Eleuthera, and the Exumas. We found 11 species only one of which, Dipodarctus subterraneus (Renaud- Debyser, 1959), had been found in the previous intertidal Bahamian collections. Thus, 10 species are new records for the Bahamas, and four of these are new to science. We describe two of the new species (Archechiniscus bahamensis sp. nov. and Anisonyches eleutherensis sp. nov.). Additional material is needed for a conclusive identification of the undescribed species of Batillipes and Florarctus. Based on examination of the holotype of Anisonyches diakidius Pollock, 1975, we redescribe this species and give an emended diagnosis of the genus and the species. We also provide new keys to the spe-cies in the genera Archechiniscus and Anisonyches. A Chao2 species richness estimate indicated that our sampling effort was extensive and probably recovered most common species. Correlations of abundance and diversity indices with regard to depth, distance to nearest shore, and sediment grain size were not significant.

Homeotherms and many poikilotherms display a positive relationship between body size and latitude, but this has rarely been investigated in microscopic animals. We analysed all published records of marine Tardigrada to address whether microscopic marine invertebrates have similar ecogeographical patterns to macroscopic animals. The data were analysed using spatially explicit generalized least squares models and linear models. We looked for latitudinal patterns in body size and species richness, testing for sampling bias and phylogenetic constraints. No latitudinal pattern was detected for species richness, and sampling bias was the strongest correlate of species richness. A hump-shaped increase in median body size with latitude was found, and the effect remained significant for the Northern Hemisphere but not for the Southern. The most significant effect supporting the latitudinal gradient was on minimum body size, with smaller species disappearing at higher latitudes. Our results suggest that biogeographical signals were observed for body size, albeit difficult to detect in poorly studied groups because of swamping from biased sampling effort and from low sample size. We did not find a significant correlation with the latitudinal pattern of body size and ecologically relevant net primary productivity.