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The Armoured Marine Tardigrades (Arthrotardigrada, Tardigrada) Abstract

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The Armoured Marine Tardigrades (Arthrotardigrada, Tardigrada) Abstract SCIENTIA The armoured marine tardigrades DANICA (Arthrotardigrada, Tardigrada) • SERIES By Jesper Guldb erg Hansen, Reinhardt Møbjerg Kristensen & Aslak Jørgensen B • BIOLOGICA • VOL. Det Kongelige Danske Videnskabernes Selskab The Royal Danish Academy of Sciences and Letters The armoured marine tardigrades (Arthrotardigrada, Tardigrada) Abstract During the last three decades several new species of the marine stygarctid tardi- grades have accumulated in the tardigrade collection at the Natural History Mu­ seum of Denmark, University of Copenhagen. The major aims of the present investigation were to describe some of the new species, to further increase the knowledge of the morphological diversity and to conduct phylogenetic analyses of all currently described species within the marine family Stygarctidae. The ma­ jor objectives of the phylogenetic analyses were to investigate the internal phylo­ genetic relationships of Stygarctidae, their relationship to the most closely relat­ ed previously recognized families (Neostygarctidae and Renaudarctidae), and the character evolution within the Stygarctidae. The outgroups were chosen to represent the extant members of presumed ancestral lineages of arthrotardi- grades (Coronarctus, Coronarctidae and Neoarctus, Neoarctidae). In order to evaluate the significance of the morphological diversity within the Stygarctidae, all species currently described have been personally examined by the authors and species new to science were described in each of the genera: Faroestygarctus nov. gen., Mesostygarctus, Parastygarctus, Pseudostygarctus and Renaudarc- tus. The character matrix consists of 31 species and 81 morphological characters. The characters have been scored from six main characters systems, i.e. the ar­ rangement of head lobes, the cuticular segmental plates (head, body and caudal plates), the seminal receptacles, the legs and claws, the sense organs and the buc­ cal apparatus. All 81 characters were parsimony informative and 51 are multistate characters. The most notable results from the phylogenetic analyses are: 1) Neostygarctus is the sister-group to the previously known genera in Stygarctinae and is included in this subfamily; 2) Faroestygarctus nov. gen. is the sister-group to the other genera in Stygarctinae; 3) Mesostygarctus is a valid taxon and is the sister-group to Pseu- dostygarctus-, 4) Stygarctusspinifer is the sister-group to the other Stygarctus species and is not a member of Parastygarctus as recently suggested. In our opinion Megastygarctides is very different from the members of Stygarc­ tinae and a position higher in the taxonomic hierarchy might be justified. Not surprisingly the Renaudarctidae is the sister-group to the Stygarctidae clade (Megastygarctidinae + Stygarctinae). Key words: marine armoured tardigrades, new species, cladistics, phylogeny, evolution. 'Jesper Guldberg Hansen, 'Reinhardt Møbjerg Kristensen, sAslak Jørgensen 'Department of Invertebrate Zoology, Zoological Museum, The Natural History Museum of Denmark, Univer­ sity of Copenhagen, Copenhagen, Denmark; “Laboratory of Molecular Systematics, The Natural History Mu­ seum of Denmark, University of Copenhagen, Copenhagen, Denmark e-mail: [email protected] The armoured marine tardigrades (Arthrotardigrada, Tardigrada) By Jesper Guldberg Hansen, Reinhardt Møbjerg Kristensen &Aslak Jørgensen Scientia Danica. Series B, Biologica • vol. 2 DET KONGELIGE DANSKE VIDENSKABERNES SELSKAB The armoured marine tardigrades (Arthrotardigrada, Tardigrada) © Det Kongelige Danske Videnskabernes Selskab 2012 Printed in Denmark by Specialtrykkeriet Viborg a-s issN 1904-5484 • isbn 978-87-7304-368-4 MIX Papir fra £5 ansvarlige kilder FSC FSC*C010558 Submitted to the Academy December 2011 Published December 2012 Table of Contents Introduction 7 Reinstatement of the genus Meso stygarctus Renaud- Mornant, 1979 31 Material and Methods 9 Mesostygarctusspiralis nov. sp. 32 Taxon sampling 9 Genus Parastygarctus Renaud-Debyser, 1965 37 Sampling sites 9 Parastygarctus robustus nov. sp. 37 Material examined 12 Parastygarctus svennevigi nov. sp. 41 Light (LM) and scanning electron microscopy (SEM) 12 Genus Pseudostygarctus McKirdy, Schmidt & McGinty- ßayiy, 1976 47 Phylogenetic Analysis 12 Pseudostygarctusgalloae nov. sp. 47 Terminology 14 Pseudostygarctus triungulatus McKirdy, Schmidt & Morphology 14 McGinty-Bayly, 1976 56 Postembryonic development and larval Comparative morphology of the seminal characters 14 receptacles 60 Heterochrony 15 Phylogenetic analysis 66 Abbreviations 17 Character states 66 Taxonomic descriptions 18 Clades, clade support (all analyses) and character Family Renaudarctidae Kristensen & Higgins, tracing 66 1984 18 Character tracing of different character systems 76 Genus RenaudarctusKristensen & Higgins, 1984 18 Discussion 80 Renaudarctusfossorius nov. sp. 18 Conclusions 82 Family Stygarctidae Schulz, 1951 25 Acknowledgements 82 Faroestygarctus nov. gen. 25 References 83 Faroestygarctus dezioae nov. gen. et nov. sp. 25 Appendix 1: List ofcharacters and character states 86 5 Introduction The phylum Tardigrada is constituted by microscopic erected a new family, Neostygarctidae, for this mono­ metazoans related to either panarthropods (Arthro- typic genus. The phylogenetic position of Neostygarctus poda and Onychophora) or nematoidans (Nematoda is of special interest as it resembles the members of and Nematomorpha) within the superclade Ecdyso- Stygarctinae in general body outline. However, zoa (Dunn et al. 2008). Recently, tardigrades have Neostygarctus exhibits cuticular extensions on the body received increased attention due to their adaptation and has unique claws, but these features might simply to survival in extreme environments (reviewed in Mø­ be autapomorphic features. Of special interest is the bjerg et al. 2011). Most of the described species of tar­ phylogenetic position of Mesostygarctus intermedius Re­ digrades are semi-terrestrial, however, the greatest naud-Mornant, 1979 and Stygarctus spinifer Hiruta, 1985 morphological diversity is present within the marine within Stygarctidae; it has recently been proposed arthrotardigrades. that these taxa be assigned to Pseudostygarctus and Para- Since the description of the first stygarctid Stygarc­ stygarctus, respectively (Gallo D’Addabbo et al. 2001). tus bradypus Schulz, 1951, a further 19 species have been Two new species belonging to the genus Megastygarcti- described in the family Stygarctidae. Representatives des have recently been described (Hansen & Kristen­ of this family are often a characteristic component in sen 2006). This study revealed several taxonomic im­ tidal and subtidal beach meiofaunas around the portant characters of the genus Megastygarctides not world. Following the monumental work by McKirdy shared by any other members of the Stygarctidae. et al. (1976) and several works by Renaud-Debyser The morphological difference between the two (1965) and Renaud-Mornant (1970, 1979, 1981a and subfamilies of Stygarctidae (Stygarctinae and Mega- 1981b) most recent studies have been published by stygarctidinae) is pronounced, as is the variation Italian investigators, who have significantly increased within the different character systems. The plate and our knowledge of the diversity, ecology and systema­ the sense organ character systems constitute approxi­ tics of these arthrotardigrades (De Zio Grimaldi et al. mately half the characters used in the data matrix for 1998; Gallo D’Addabbo et al. 2000 and 2001; Grimal­ phylogenetic analyses. Furthermore, we introduce the di de Zio et al. 1987). seminal receptacles as a new character system, which The phylogeny has been investigated by Bello & shows a surprisingly high diversity within the Styg­ Grimaldi de Zio (1998), who inferred the intergeneric arctidae. The morphological diversity of the seminal relationships of the family. Kristensen & Higgins (1984) receptacles has not been thoroughly studied in other and Bello & Grimaldi de Zio (1998) regard Renaudarc- arthrotardigrade families. tidae as being the sister-group to Stygarctidae. Dewel & A Stygarctidae-like animal has been pictured as Dewel (1997), Renaud-Mornant (1982) and Simonetta the ancestral tardigrade in recent paleontological stu­ & la Cave (1991) have argued for a position of the Styg­ dies on panarthropod evolution (Dewel & Dewel arctidae close to the root of the tardigrade phylogenetic 1997; Budd 2001). The genus Stygarctus was used as tree. Stygarctus was recently included in a phylogenetic one of the terminals in the strict consensus tree of study by Jørgensen et al. (2010) and its position was Dewel and Dewel (Figure 10.5, 1997). In their phylo­ inferred to be near the base of the arthrotardigrades, genetic analysis Stygarctus branches off between the however, the 28S rRNA sequence was somewhat aber­ extinct lobopodian Aysheaia and the extinct lateral­ rant with regard to the other arthrotardigrades. flapped Kerygmachela. These new paleontological ana­ Neostygarctus Grimaldi de Zio et al., 1982 was placed lyses were not very different from the conclusions of in Stygarctidae until Grimaldi de Zio et al. (1987) older reports on the phylogeny of lobopods and tardi- 7 INTRODUCTION SCI.DAN.B. 2 grades (Renaud-Mornant 1987; Simonetta & la Cave cted in the present study was thus expected to incre­ 1991). However, Renaud-Mornant (1987) argued that ase the phylogenetic resolution at higher levels, parti­ the genus Parastygarctus might resemble the stem group cularly as a new genus (and subfamily) is described for all
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