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The Deep Sea Elements of the Faroe Bank Tardigrade Fauna with a Description of Two New Species

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The Deep Sea Elements of the Faroe Bank Tardigrade Fauna with a Description of Two New Species G. Pilato and L. Rebecchi (Guest Editors) Proceedings of the Tenth International Symposium on Tardigrada J. Limnol., 66(Suppl. 1): 12-20, 2007 The deep sea elements of the Faroe Bank tardigrade fauna with a description of two new species Jesper GULDBERG HANSEN Department of Invertebrate Zoology, Zoological Museum, University of Copenhagen, Universitetsparken 15, DK-2100 Copenhagen, Denmark e-mail: [email protected] ABSTRACT Two new marine Tardigrada species are described from the calcareous sediments at the steep slope of the Faroe Bank in the North Atlantic Ocean. Parmursa torquata sp. nov. can be distinguished mainly by small cylindrical secondary clavae, and the presence of caudal and cephalic vesicles, and a large ventral plate. Coronarctus verrucatus sp. nov. is characterised by its unique cuticular sculpture, with numerous small wart-like excrescences, regularly distributed all over the body. These new records from the relatively shallow water of the Faroe Bank (200-260 m) further widen the range of Parmursa and Coronarctus distribution and diversity, especially regarding the genus Parmursa, which to date has remained monospecific. Key words: Tardigrada, Arthrotardigrada, Faroe Bank, Parmursa torquata sp. nov, Coronarctus verrucatus sp. nov. ethanol and acetone prior to critical point drying. The 1. INTRODUCTION dehydrated specimens were then mounted on aluminium stubs, coated with gold and observed in a JEOL JSM- Although deep-sea tardigrades have been known 840 scanning electron microscope. The type-material is since the mid-1960's, the published information is deposited in the collections of the Zoological Museum, scattered and data about their worldwide distribution are Copenhagen (ZMUC), Denmark. In addition to the still very scarce. This study adds yet another piece to the examination of two paratypes (AR 316 and AR 317) of puzzle of deep-sea tardigrades. Since the erection of Parmursa fimbriata Renaud-Mornant, 1984, this study Parmursa Renaud-Mornant, 1984 and the addition of also includes the examination of a single specimen of four new species to the genus Coronarctus by Renaud- Parmursa sp. collected at 1813 m depth from Lihir Mornant in 1987, no further species have been ascribed Island in the New Ireland Basin, by the German FS to either of these bathyal and abyssal genera. Among the Sonne (Herzig 1998). This species cannot be attributed collections gathered during the BIOFAR programme in to either P. fimbriata or Parmursa torquata sp. nov.; the North Atlantic Sea (Kristensen 1992, 1999; Hansen however a detailed description awaits the finding of et al. 2001; Hansen 2005), new species of both further specimens of this particular species. Parmursa and Coronarctus were found. Abbreviations: an: anus, ap: apophysis, bt: buccal tube, 2. METHODS ca: caudal ala, cB-E: cirrus B-E, cg: claw gland, ec: external cephalic cirrus, go: gonopore, gu: midgut, ic: The new species from the Faroe Bank was internal cephalic cirrus, lc: lateral cirrus, ma: muscle encountered during expeditions in 1990, 1992 and 1998. attachment, mc: median cirrus, mo: mouth cone, mu: BIOFAR Station 627 was sampled by the German R/V muscle, pb: pharyngeal bulb, pc: primary clava, pl: Valdivia (1990), BIOFAR Station 785 was sampled by placoid, pv: ventral plate, rs: seminal receptacle, sc: the Faroese Coast Guard vessel Olavur Halgi (1992) and secondary clava, se1-4: sense organ of leg 1-4, sp4: BIOFAR Station 2011 was sampled by R/V Magnus spine-like projection of leg 4, st: stylet, te: testis, sv: Heinason of the Faroese Fishery Investigations (1998). seminal vesicle, v.ca: caudal vesicle, v.ce: cephalic Details on the sampling-localities are provided in vesicle. Nørrevang et al. (1994). To release the tardigrades from the sediment, a number of sub-samples at each station 3. RESULTS were fresh-water shocked, decanted through a 32 µm mesh net and fixed in 4% buffered formaldehyde for 3.1. Family Halechiniscidae Thulin, 1928 later sorting. Specimens were mounted on micro slides Diagnosis: Arthrotardigrades without sclerotised dorsal in glycerol and studied using phase-contrast and segmental plates. The complete set of cephalic Nomarski-technique (DIC). A single specimen of each appendages in halechiniscids is 11, but the secondary species was prepared for scanning electron microscopy clavae may be indistinguishable (9) or a pair of tertiary (SEM). They were dehydrated in a graded series of clavae may be present (13). Each adult leg with four Deep Sea Tardigrades of the Faroe Bank 13 Fig. 1. Parmursa torquata sp. nov.: adult holotypic male, ventral view. Scale bar = 25 µm. Drawing by Stine B. Elle. digits with claws. Claws simple or with accessory Cephalic cirri with a funnel-shaped scapus. Erect hooks, spurs or thin bristles. Peduncles can be present secondary clavae. Stylet supports absent. on all digits or only on external digits. Two cuticular Type species: Parmursa fimbriata Renaud-Mornant, seminal receptacles always present in adult females. 1984. 3.2. Subfamily Euclavarctinae Renaud-Mornant, 1983 3.4. Parmursa torquata sp. nov. (Figs 1-4) Halechiniscidae with unplated body without Material: Holotype [slide TAR 690, ZMUC], and 4 projections. Conical head bearing two sets of unequally paratypes, [slide TAR 691-694, ZMUC] (all males). shaped clavae. Cirri A and primary clavae are inserted separately. The claws are simple or with dorsal spur on Type locality: BIOFAR station 2011 (61˚12.0' N; 08˚31 internal digits. W), Faroe Bank, North Atlantic Sea. Depth range: 200 m. The sediment is represented by fine sand, shell Type genus: Euclavarctus Renaud-Mornant, 1975. gravel, cobbles. 3.3. Genus Parmursa Renaud-Mornant, 1984 Etymology: From torquis (Latin) twisted collar. Diagnosis (emended): Euclavarctinae with cuticular 3.4.1. Diagnosis aliform expansions supported externally by dorsal cuticular ribs. Trapezoid head, completely proportionate Parmursa with small cylindrical secondary clavae. to the arrangement of the cephalic sense organs. Primary clavae with a sausage-shaped distal part. A pair 14 J. Guldberg Hansen Fig. 2. Parmursa torquata sp. nov.: adult paratypic male, dorsal view. Scale bar = 25 µm. Drawing by Stine B. Elle. of cephalic vesicles and a caudal vesicle are present. that the ala actually is continuous, extending from the Lateral alae extending on both sides of the body, from primary clava to leg IV. Dorsal cuticular ribs support the primary clavae to the fourth pair of legs and a caudal the alae externally (Fig. 4c). ala extending between the fourth pair of legs. Erect The trapezoid head (Figs 3b, d and 4b) is dorsal cephalic ala, extending between the primary differentiated from the body by the large pedestal of clavae. Simple external claws, internal claws with a primary clavae and lateral cirri. All the cephalic cirri long dorsal accessory spine. consist of a cirrophore, funnel-shaped scapus and a Holotype: The holotype (Figs 1 and 3) is 142 µm long. flagellar portion. The internal cirri (17 µm) arise at the The body is ovoid, being broadest (83 µm) at the level anterior margin of the head. The external cirri (20 µm) between the second and third pair of legs. The dorsal are inserted ventrally and the median cirrus (20 µm) cuticle has a plated appearance with four transverse mid-dorsally, anterior to the cephalic ala. The primary inter-segmental depressions: one anterior to the first pair clava (20 µm) has a swollen proximal part and a of legs, one between the first and second pair of legs, sausage-shaped distal part. A Van der Lands’s body is one between the second and third pair of legs and one visible inside its base. Primary clava and lateral cirrus between the third and fourth pair of legs (Figs 2 and 4a). (38 µm) arise separately, on the same pedestal. The An erect dorsal cephalic ala (Figs 2 and 4a, b), two secondary clavae are small (9 µm) and cylindrical, lateral alae (Figs 2 and 4a, c), and a caudal ala (Figs 2 without an evident cirrophore. They arise ventrally, and 4d) are present. Folds give the impression that each bending upwards and extend dorsally. The leg I sense lateral ala is divided into three separate alae, however it organ is an unsegmented spine with a slightly swollen is evident using scanning electron microscopy (SEM) base and a terminal tube. The sense organs of leg II and Deep Sea Tardigrades of the Faroe Bank 15 Fig. 3. DIC-micrographs of Parmursa torquata sp. nov., holotypic male. a. Overview. b. Close up on the head, dorsal view. c. Close up on the gonopore. d. Close up on the head, ventral view. Scale bars: a = 25 µm, b, c, d = 10 µm. III are unsegmented tapering spines. The fourth leg have a long accessory spine. All the claws can be sense organ is an elongate papilla with a basal Van der retracted into claw sheaths. Lands’s body and a terminal tube. The cirrus E (50 µm) The mouth cone consists of four parts: a large basal has a prominent cirrophore, strongly annulated scapus cone, and three telescopic segments. The buccal tube is and a long tapering flagellum (Fig. 4d). 39 µm long and thin. The stylets are 42 µm long and A pair of cephalic vesicles and a caudal vesicle are very thin, each with branches of furcae so diverging as present (Figs 3a and b). The cephalic vesicles open to give a rectilinear appearance (Fig. 3d). Stylet sup- dorsally, and the caudal vesicle opens ventrally. Their ports are lacking. The almost straight placoids are short, function is presently unknown, but their contents look thick, and sharply pointed. The pharyngeal apophyses like bacteria, indicating a function similar to the vesicles are very small and droplet-shaped. From the pharyngeal seen in florarctids (Kristensen 1984). bulb, a short oesophagus leads to the diverticulated mid- The leg consists of coxa, femur, tibia and tarsus.
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