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ORI GI NAL PAPER Tardigrada of the Revdalen (Spitsbergen) with the descriptions of two new species: Bryodelphax parvuspolaris (Heterotardigrada) and Isohypsibius coulsoni (Eutardigrada) Łukasz Kaczmarek ã Krzysztof Zawierucha ã Jerzy Smykla ã Łukasz Michalczyk Received: 22 June 2011 / Revised: 17 December 2011 / Accepted: 19 December 2011 / Published online: 19 January 2012 Ó The Author(s) 2012. This article is published with open access at Spri
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  ORIGINAL PAPER Tardigrada of the Revdalen (Spitsbergen) with the descriptionsof two new species:  Bryodelphax parvuspolaris  (Heterotardigrada)and  Isohypsibius coulsoni  (Eutardigrada) Łukasz Kaczmarek  ã Krzysztof Zawierucha  ã Jerzy Smykla  ã Łukasz Michalczyk Received: 22 June 2011/Revised: 17 December 2011/Accepted: 19 December 2011/Published online: 19 January 2012   The Author(s) 2012. This article is published with open access at Springerlink.com Abstract  Despite a century long history of research,tardigrade fauna of the Svalbard Archipelago remainspoorly known. In order to deepen our knowledge of tar-digrade biodiversity in the Arctic, we collected forty-onemoss and lichen samples from the Revdalen and on thesouth-east slopes of the Rotjesfjellet (Spitsbergen, SvalbardArchipelago) in June 2010. In these samples, twenty-fivetardigrade species were found, including two new for sci-ence:  Bryodelphax parvuspolaris  sp. nov. and  Isohypsibiuscoulsoni  sp. nov.  B. parvuspolaris  sp. nov. belongs to the weglarskae  group but differs from all other species of thegroup by a unique configuration of ventral plates.  I. coul-soni  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 asi-aticum  Tumanov, 2006 and  Diphascon  (  Adropion )  prorsi-rostre  Thulin, 1928, are recorded from the SvalbardArchipelago for the first time. Keywords  Arctic    Faunistics    New records   Tardigrada    Taxonomy    Biodiversity Introduction Although the first studies on water bears (Tardigrada) of the Svalbard Archipelago took place as early as in thenineteenth century (Scourfield 1897) and were continuedby a number of researchers throughout the twentieth cen-tury (e.g. Richters 1903, 1904, 1911; Murray 1907; Marcus 1928; We˛glarska 1965; Dastych 1985; Pugh and McInnes 1998; Łagisz 1999), our knowledge about the tardigrade fauna of this region is still relatively poor. The mostcomprehensive work on Spitsbergen tardigrades (Dastych1985) provided a complete species list, new records andalso investigated the ecology of these microscopic inver-tebrates. A decade later, Pugh and McInnes (1998) ana-lysed the srcin of the Arctic Tardigrada. Other paperswere published sporadically, and the majority were limitedto reports and descriptions of new species from the Sval-bard Archipelago (e.g. Binda et al. 1980; Pilato et al. 1982; Dastych 1983; De Smet et al. 1987, 1988; Pilato and Binda 1987; Van Rompu and De Smet 1988, 1991, 1994 ; DeSmet and Van Rompu 1994; Maucci 1996; Tumanov 2007; Smykla et al. 2011). In the region of Hornsund, studieswere previously conducted by We˛glarska (1965), Dastych(1985), Maucci (1996), Janiec (1996) and Łagisz (1999). Up to now, only 84 species were recorded from the Ł. Kaczmarek ( & )    K. ZawieruchaDepartment of Animal Taxonomy and Ecology,A. Mickiewicz University, Umultowska 89,61-614 Poznan, Polande-mail: kaczmar@amu.edu.plK. Zawieruchae-mail: krzysiu_zaw@wp.plJ. SmyklaDepartment of Biodiversity, Institute of Nature Conservation,Polish Academy of Sciences, Mickiewicza 33,31-120 Krako´w, Polande-mail: smykla@iop.krakow.pl Present Address: J. SmyklaDepartment of Biology and Marine Biology, University of NorthCarolina Wilmington, 601 S. College Rd., Wilmington,NC 28403, USAŁ. Michalczyk School of Biological Sciences, University of East Anglia,Norwich Research Park, Norwich NR4 7TJ, UK e-mail: LM@tardigrada.net  1 3 Polar Biol (2012) 35:1013–1026DOI 10.1007/s00300-011-1149-0  Svalbard Archipelago, but none specifically from theRevdalen (Coulson 2011).Here, we provide a list of tardigrades species from theRevdalen, including two new records from the SvalbardArchipelago and descriptions of two species new for sci-ence,  Bryodelphax parvuspolaris  sp. nov. and  Isohypsibiuscoulsoni  sp. nov. The genus  Bryodelphax  consists of only17 species, but its distribution is global—it has beenrecorded from the polar regions to tropical rain forests(Kaczmarek and Michalczyk  2004; Kaczmarek et al. 2005; Kristensen et al. 2010; Degma et al. 2011). In contrast, the genus  Isohypsibius  is one of the largest in the phylumTardigrada, with more than 130 species and subspeciesdescribed from all over the world (McInnes 1994; Degmaet al. 2011). Materials and methods Moss and lichen samples for this study were collected fromthe Revdalen and the Rotjesfjellet, which are located on thenorth coast of Hornsund (Spitsbergen, Svalbard Archipel-ago; Fig. 1). The total of forty-one moss and lichen sampleswere collected on the 26th June 2010 from the Revdalen andon the 29th June 2010 from the south-east slopes of theRotjesfjellet (see Table 1). Twenty-nine (over 70%) of thesamples provided a total of 461 specimens and 83 eggs.All specimens and eggs were mounted on microscopicslides in Hoyer’s medium and then examined and photo-graphed with a Phase Contrast Microscope (PCM). Specieswere identified using the key to the World Tardigrada(Ramazzotti and Maucci 1983) and srcinal descriptionsfrom the literature.All measurements are given in micrometres ( l m).Structures were measured only if their orientations weresuitable. Body length was measured from the anterior tothe posterior end of the body, excluding the hind legs.Measurements of the species used in differential diagnosesare given or calculated according to the srcinal descrip-tions (i.e. Pilato 1972, 1974; Bertolani et al. 1995; Kristensen et al. 2010). Claws of   Isohypsibius coulsoni  sp.nov. were measured according to Beasley et al. (2008).In eutardigrades, the  pt   ratio is the ratio of the length of a given structure to the length of the buccal tube, expressedas a percentage (Pilato 1981). Similarly, to provide relativemeasurements in echiniscids, the  sc  ratio of the length of agiven structure to the length of the scapular plate is used(e.g. Fontoura and Morais 2011). Both values are always Table 1  The list of localities in the Hornsund area, from which samples containing tardigrades were collectedLocality no. Locality name and coordinates Plant Substrate m aslI Northern part of the Revdalen, near the Revvatnet and the Revelva (77  01 0 29 00 N; 15  22 0 39 00 E) Moss Rock 51II–IV Northern part of the Revdalen, near the Revvatnet and the Revelva (77  01 0 41 00 N; 15  22 0 21 00 E) Moss Soil 67V Northern part of the Revdalen, near the Revvatnet and the Revelva (77  01 0 41 00 N; 15  22 0 21 00 E) Moss Rock 67VI–VIII Northern part of the Revdalen, near the Revvatnet and the Revelva (77  01 0 39 00 N; 15  22 0 47 00 E) Moss Rock 76IX Northern part of the Revdalen, near the Revvatnet and the Revelva (77  01 0 34 00 N; 15  23 0 12 00 E) Moss Rock 76X Northern part of the Revdalen, near the Revvatnet and the Revelva (77  01 0 26 00 N; 15  23 0 30 00 E) Moss Soil 68XI Northern part of the Revdalen, near the Revvatnet (southern edge) and the Revelva(77  01 0 09 00 N; 15  24 0 34 00 E)Moss Rock 50XII–XIII Northern part of the Revdalen, near the Revvatnet (southern edge) and the Revelva(77  01 0 09 00 N; 15  24 0 34 00 E)Moss Soil 50XIV Northern part of the Revdalen, near the Revvatnet (southern edge) and the Revelva(77  01 0 09 00 N; 15  24 0 34 00 E)Moss,lichenSoil 50XV The Revdalen, south-east of the Revvatnet and the Revelva (77  00 0 35 00 N; 15  28 0 20 00 E) Moss Soil 36XVI The Revdalen, south-east of the Revvatnet and the Revelva (77  00 0 22 00 N; 15  29 0 02 00 E) Moss Soil 29XVII–XVIII The Rotjesfjellet, south-east slope (77  00 0 16 00 N; 15  24 0 02 00 E) Moss Soil 50XIX–XX The Rotjesfjellet, south-east slope (77  00 0 19 00 N; 15  23 0 55 00 E) Moss Soil 100XXI–XXII The Rotjesfjellet, south-east slope (77  00 0 26 00 N; 15  23 0 42 00 E) Moss Soil 201XXIII The Rotjesfjellet, south-east slope (77  00 0 29 00 N; 15  23 0 35 00 E) Moss Soil 250XXIV The Rotjesfjellet, south-east slope (77  00 0 31 00 N; 15  23 0 21 00 E) Moss Soil 301XXV The Rotjesfjellet, south-east slope (77  00 0 31 00 N; 15  23 0 21 00 E) Moss,lichenSoil 301XXVI–XXVIIThe Rotjesfjellet, south-east slope (77  00 0 35 00 N; 15  22 0 58 00 E) Moss Rock 399XXVIII–XXIXThe Rotjesfjellet, the top (77  00 0 40 00 N; 15  22 0 20 00 E) Moss Rock 4371014 Polar Biol (2012) 35:1013–1026  1 3  provided in italics, in order to differentiate them from othermeasurements and ratios.The configuration of ventral plates in the genus  Bryodel- phax  is described using an analogous system to that used forthe description of cuticular gibbosities in some eutardigrades(Michalczyk and Kaczmarek  2010), i.e. a Roman numberwith a colon at the beginning of the sequence indicates thetotal number of rows of ventral plates and following Arabicnumbersseparated bydashesrefer tothenumbers ofplatesineach row, starting from the most anterior row.Alltheinvestigatedmaterialispreservedinthecollectionof the first author, at the Department of Animal Taxonomyand Ecology, A. Mickiewicz University, Poznan´, Poland. Results Taxonomic accounts of the new species foundin the study  Bryodelphax parvuspolaris sp. nov. (Figs. 2 – 8; Table 2)  Material examined   Holotype (female) (slide 14.2/20) and 8paratypes (females) (slides: 14.2/4, 14.2/8, 14.2/9, 14.2/12,14.2/13, 14.2/14, 14.2/16, 14.2/19).  Description (measurements in Table 2)  Body (Figs. 2–4) transparent to slightly rose, eyes absent or not visible afterthe preparation. Apart from the head appendages (cirrus internus  and  externus  and drop-shaped cephalic papillae(secondary clava)), only lateral cirrus  A  (with clava nearthe base (primary clava) present).Dorsal plates covered with fine, but distinct dark dotsthat appear as granulation under PCM, but are in factcuticular pillars within cuticle (Michalczyk and Kaczmarek 2006, 2007). ‘‘Granulation’’ distinctly larger on the scap- ular and the terminal plate. In addition to the ‘‘granula-tion’’, slightly larger and irregularly distributed pores arevisible mainly on the margins of all dorsal plates (Fig. 5).Scapular plate facetted with a median longitudinal fold anda few smaller transverse folds. Paired plates divided intotwo unequal anterior and posterior parts by a transversestripe without ‘‘granulation’’. Median plates 1 and 2 divi-ded, and median plate 3 undivided. Twelve supplementaryplates poorly visible near median plates 1–3. The terminalplate facetted with two longitudinal folds.Ventral plates arranged in eight rows: 1 plate in row I(between legs I), 1 plate in row II (between legs I and II), 2plates in row III (between legs I and II), 2 plates in row IV(between legs II), 2 plates in row V (between legs II andIII), 2 plates in row VI (between legs III), 2 plates in row Fig. 1  The study area:  a  Svalbard Archipelago  b  Hornsund, West Spitsbergen,  c  Revdalen and Rotjesfjellet (maps from Norsk Polarinstitute)Polar Biol (2012) 35:1013–1026 1015  1 3  VII (in line with the gonophore) and 1 plate in row VIII(below the gonophore); i.e. the ventral plate configurationVIII:1-1-2-2-2-2-2-1 (Figs 4, 8). All ventral plates with fine and indistinct ‘‘granulation’’ (Fig. 6).Spine on legs I and papilla on legs IV absent or notvisible under PCM. Collar on legs IV with poorly devel-oped and irregular teeth. External claws of all legs smooth,internal claws with very small spurs directed downwards(Fig. 7).Eggs unknown.  Remarks  In some specimens, ventral plates are indis-tinct; thus, an examination of at least several specimens toensure correct identification is strongly recommended. Etymology  The name ‘  parvuspolaris ’, meaning ‘a smalldweller from the polar regions’, was chosen by the par-ticipants of the XXXIII Polar Expedition of the PolishAcademy of Sciences, who provided us with logisticalsupport and helped collecting samples from Spitsbergen. Type locality  Hornsund, northern part of the Revdalen,near the Revvatnet and the Revelva, mosses from soil,67 m asl, 77  01 0 41 00 N, 15  22 0 21 00 E, 26.06.2010, coll.Łukasz Kaczmarek and Jerzy Smykla. Type depositories  Holotype (slide 14.2/20) and para-types (slides: 14.2/4, 14.2/8, 14.2/9, 14.2/12, 14.2/13, 14.2/ 14, 14.2/16, 14.2/19) are deposited at the Department of Animal Taxonomy and Ecology, Institute of Environmen-tal Biology, A. Mickiewicz University, Umultowska 89,61-614 Poznan´.  Differential Diagnosis Bryodelphax parvuspolaris  sp.nov. has ventral plates and thus belongs to the  weglarskae group (Kristensen et al. 2010); we therefore only comparedother species of this group (see Fig. 8), using the ventralplates and other characters. The new species differs from: ã  B. aaseae  Kristensen et al., 2010 by: a different ventralplate configuration (VIII:1-1-2-2-2-2-2-1 in the newspecies and X:2-1-4-4-2-4-2-1-2-1 in  B. aaseae ) andthe presence of dentate collars on hind legs. ã  B. iohannis  Bertolani et al., 1995 by: a different ventralplate configuration (VIII:1-1-2-2-2-2-2-1 in the newspecies and X:2-1-1-5-2-4-2-2-2-1 in  B. iohannis ), aslightly smaller body size (87.4–125.0 in the newspecies and 113.9–179.5 in  B. iohannis ), and thepresence of dentate collars on hind legs. ã  B. sinensis  Pilato, 1974 by: a different ventral plateconfiguration (VIII:1-1-2-2-2-2-2-1 in the new speciesand VII:2-2-2-2-2-2-1 in  B. sinensis ), slightly longerlateral appendages  A  (up to 37.4 in the new species andup to 27.0 in  B. sinensis ), and the presence of dentatecollars hind legs. ã  B. weglarskae  Pilato, 1972 by: a different ventral plateconfiguration (VIII:1-1-2-2-2-2-2-1 in the new speciesand IX:2-1-5-2-4-2-2-2-1 in  B. weglarskae ) and by notbifurcated appendages.  Isohypsibius coulsoni sp. nov. (Figs. 9 – 13; Table 3)  Material examined   Holotype (slide 34.2/29) and 80 para-types (slides: 31.4/1, 31.4/3, 34.2/6, 34.2/7, 34.2/8, 34.2/9,34.2/10,34.2/11,34.2/12,34.2/13,34.2/14,34.2/15,34.2/16, Table 2  Measurements and  sc  values of selected morphological structures of nine specimens (including the holotype) from the type populationof   Bryodelphax parvuspolaris  sp. nov.Character N Range Mean SD Holotype l m  sc  l m  sc  l m  sc  l m  sc Body length 9 87–125  523 – 556   110  542  11  17   122  546  Scapular plate length 3 16.7–22.3 – 19.6 – 2.8 – 22.3 –Head appendages lengthsCirrus  internus  7 4.4–7.3  22.4 – 26.3  5.5  24.3  1.1  2.7   5.0  22.4 Cephalic papilla 6 2.4–4.0  15.2 – 17.9  2.9  16.5  0.6  2.0  4.0  17.9 Cirrus  externus  7 9.0–16.1  52.5 – 57.6   11.8  55.0  2.2  3.6   11.7  52.5 Clava 8 1.4–4.2  14.1 – 18.8  2.7  16.5  0.9  3.3  4.2  18.8 Cirrus  A  8 22.9–37.4  144.4 – 164.6   29.9  154.5  5.0  14.3  32.2  144.4 Cirrus  A  /Body length ratio 8 21%–32% – 27% – 5% – 26% –Cirrus  int   /  ext   length ratio 7 43%–53% – 47% – 3% – 43% –Claw 4 lengthsBranch 9 4.5–6.3  26.9 – 31.8  5.6  28.6   0.6  2.8  6.0  26.9 Spur 9 0.7–1.3  3.5 – 4.9  0.9  4.4  0.2  0.8  1.1  4.9 Spur/branch length ratio 9 11%–22% – 16% – 3% – 18% –  N   number of specimens or structures measured,  Range  the smallest and the largest structure found among all specimens measured,  SD  standarddeviation,  sc  ratio of the length of a given structure to the length of the scapular plate, expressed as a percentage1016 Polar Biol (2012) 35:1013–1026  1 3

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