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Additional file 1: SM.1 List of primers and PCR programs.

Fractonotus gilvus (Biserov, 1986) n. comb. Isohypsibius gilvus Biserov, 1986: 984. REMARK Elliptical organs not always visible due to developed sculpturing in the cephalic portion of the body., Published as part of G��siorek, Piotr, Morek, Witold, Stec, Daniel, Blagden, Brian & Michalczyk, ��ukasz, 2019, Revisiting Calohypsibiidae and Microhypsibiidae: Fractonotus Pilato, 1998 and its phylogenetic position within Isohypsibiidae (Eutardigrada: Parachela), pp. 71-89 in Zoosystema 41 (6) on page 83, DOI: 10.5252/zoosystema2019v41a6, http://zenodo.org/record/3718524, {"references":["BISEROV V. I. 1986. - Terrestrial water bears from the North Caucasus. 2. Eutardigrada. Zoologicheskii Zhurnal 65 (7): 981 - 993."]}

Fractonotus verrucosus (Richters, 1900) n. comb. Macrobiotus ornatus var. verrucosus Richters, 1900: 41 (terra typica: Taunus, Germany). Macrobiotus scabrosus Murray, 1911: 10 (locus typicus: Clare Island, Ireland). Hypsibius verrucosus ��� Thulin 1911: 29 (Kiruna, Lapland, Sweden). ��� Marcus 1928: 180 (Vannsee, Berlin, Germany). ��� Cu��not 1932: 77 (the Vosges, France and Switzerland). ��� da Cunha 1947, 1948: 6, 2 (Serra d���Arga, Serra do Bu��aco, Serra da Estrela, Serra da Lous��, Portugal). Calohypsibius verrucosus ��� Thulin 1928: 239 (Sweden). Calohypsibius scabrosus Thulin, 1928: 239 (Sweden). Hypsibius scabrosus Cu��not, 1932: 77 (the Vosges, France and Switzerland). ��� da Cunha 1947, 1948: 6, 2 (Serra d���Arga, Serra do Bu��aco, Serra da Estrela, Serra da Lous��, Portugal). Hypsibius (Calohypsibius) verrucosus ��� Marcus 1936: 285 (Schwarzwald, Germany). ��� Franceschi 1951-1952: 12 (Val Camonica, Italy). ��� Mihelčič 1953: 247 (Tirol, Austria). ��� Fontoura 1981: 18 (Viseu, Arga, Amarante, Portugal). Hypsibius placophorus da Cunha, 1943: 1 (locus typicus: Cabril do Ceira, Portugal); 1947, 1948: 2, 2 (Serra d���Arga, Serra do Bu��aco, Serra da Estrela, Serra da Lous��, Portugal) n. syn. LOCALITIES. ��� Scotland. Creag Meagaidh (56��57���10������ N, 4��30���35������ W; 291 m a.s.l.; collection date: 1.X.2014), lichens from moorland rocks; Scotland, Hebrides, Isle of Lewis, Loch nan Muilne (58��21���08������ N, 6��35���14������ W; 27 m a.s.l.; collection date: 29.VII.2015), moss and lichen mix from stones on the lakeshores; Invermoriston, Loch Ness (57��12���39������ N, 4��35���59������ W; 20 m a.s.l.; 25.X.2015; Brian Blagden leg.), moss and lichen mix from stones on the lakeshores. MATERIAL EXAMINED. ��� 23 individuals, UJ (19 specimens, including one simplex, on slides GB.005.03-12, GB.008.01-3, GB.028.01- 2 and 4 specimens on two SEM stubs); 2 individuals, MNHN (slides GB.005.01-2); 3 individuals, NHMD (slides GB.008.04- 5); 2 individuals, UAM (slides GB.028.03-4); 1 individual, CU (slide GB.028.02). ETYMOLOGY (NOT PROVIDED IN THE ORIGINAL DESCRIPTION). ��� The name most likely refers to the rugged cuticular surface of the species (from Latin verruca = wart). DIFFERENTIAL DIAGNOSIS. ��� Fractonotus verrucosus n. comb. can be distinguished from F. caelatus and F. gilvus n. comb. (Fig. 2 A, B) by the presence of plaques (absent in the latter species). It also differs from F. gilvus n. comb. by shorter, stouter claws (anterior and posterior primary branches 4.1-6.4 ��m [N =10] and 4.3-7.4 ��m long [N =18], respectively, in Fractonotus verrucosus n. comb. vs 7.0- 13.0 ��m [N =21] and 10.5-16.5 ��m long [N =21], respectively, in F. gilvus n. comb.; compare Fig. 11 A-D). INTEGRATIVE DESCRIPTION Animals (see Table 3 for measurements) Body stubby, typically slightly rose in live animals, transparent in mounted specimens. Dorsum strongly sculptured from the first instar, although with substantial ontogenetic quantitative and qualitative variability in this trait (Fig. 1 A- F). Juveniles with ten transverse bands of numerous tu- bercles that increase in size towards the caudal end of the body, but fully formed plaques never present, legs covered with fine tubercles (Fig. 1 A). All ten bands not always easily identifiable under PCM in juveniles. In young adults, plaques present in bands 6-10, with the most prominent plaques in bands 8-10 (Fig. 1 B). In older adults, smooth spaces between the transverse bands becoming narrow and sometimes merge into larger areas (Fig. 1 C-F). Plaques larger and more numerous than in young adults, and typically developping in bands 4-10, but the most evident plaques present in the caudal part of the body (Fig.1 C-F). Tubercles more or less round or oval (Figs 3 A, B; 5 A, B), gradually increasing in size from juveniles to adults, and becoming scabrous with age (compare Figs 1 A-F; 5 A, B). Plaques, on the other hand, typically smooth and only sometimes slightly rough (Fig. 5 C, D, arrowheads); under stereomicroscope strongly opalescent. Plaques arranged symmetrically in respect to the longitudinal body axis, although deviations A from symmetry are not rare (Fig. 1 C, D). In adults, seven pairs of central plaques and four lateral plaque pairs. Central plaques triangular in shape, with their apices directed laterally and outwards. In rows where only central plaques are present, plaques slimmer and longer than in rows with lateral plaques. Central plaques present in bands aligned with legs I-III as well as in bands between those legs. First three pairs of lateral plaques in line with legs I-III and the last pair of double lateral plaques situated between legs III and IV (Fig. 1 E). Plaque configuration VII:4-2-4-2-4-2-6. Cephalic elliptical organs present but not easy to identify, given the rich cuticular sculpturing (Fig. 7 A). Eyes absent in live animals. Buccal apparatus of the Fractonotus - type (Fig. 7 B, C, E), i.e. with a long ventral AISM, and the dorsal AISM subdivided into the proximal, weakly developed thickening, and the distal, small blunt hook (Fig. 9 A). Mouth opening surrounded by six large and soft peribuccal lobes (visible only under SEM, Fig. 6 A). Oral cavity armature, visible only under SEM, consisting of a single row of minute conical teeth located on the ring fold (Fig. 8 A). Two distinct porous areas on the lateral sides of the buccal crown are visible in SEM only (Fig. 8 B). Stylet furcae of the modified Hypsibius shape, i.e. with very broad and trapezoid bases, thick arms and rounded apices (Figs 7 B, 8 D, 10 A). Buccal tube with slight lateral thickenings posterior to the stylets supports (Figs 7 B, C, E; 8 C). Round bulbus with large pharyngeal apophyses (almost as large as the placoids), and two granular macroplacoids (Figs 7 B, C, E; 8 E, F). In PCM, macroplacoids without constrictions, however slight central constrictions in both macroplacoids detectable under SEM (Fig. 8 E, F). Claws of the modified Isohypsibius - type (Figs 11 A-C; 12 A, B). Specifically, claw bases triangular, especially pronounced in claws IV (Figs 11 C, 12 B). Claw branches V-shaped, elongated and strongly curved. Apparent accessory points on the primary branches (Figs 11 A-C; 12 A, B). Weakly developed pseudolunulae present, particularly visible under the internal and anterior claws (Fig. 11 A, C). Claw septa and cuticular bars on legs absent. Eggs Roundish and smooth, deposited in exuviae (up to two eggs per exuvia recorded). MOLECULAR MARKERS The sequences for all DNA markers were of a good quality. The sequenced fragments were of the following lengths: 1.727 bp (18 S rRNA; MG 800855), 819 bp (28 S rRNA; MG 800856), and 499 bp (ITS-2; MG 800857). All markers, including the specimen without cuticular plaques, were represented by single haplotypes. The p-distances between 18S haplotypes of all available isohypsibioid species and Fractonotus verrucosus n. comb. ranged from 2.0% (I. prosostomus Thulin, 1928, EF 620404 from Denmark) to 7.1% (Hexapodibius micronyx Pilato, 1969, HQ 604915 from Italy), with an average distance of 5.2%. As our 28S rRNA primers obtain a different gene fragment to the one sequenced by previous authors, comparisons of this gene were not possible. Matrices with p-distances are provided in the Supplementary Material 2. REMARKS The vast part of the Richters Collection has been lost, thus the type material (if ever existed) is not available for examination. Moreover, no specimens from Germany were examined in this study, therefore the neotype series is not established. Hence, until the redescription from the terra typica in Germany is available, we propose to consider the description of the Scottish specimens only as the current perception of the species. PHYLOGENETIC POSITION OF FRACTONOTUS AMONG OTHER ISOHYPSIBIIDAE Both Bayesian Inference and Maximum Likelihood methods unreservedly located Fractonotus within Isohypsibioidea (Fig. 13), thus corroborating the phenotypic analysis (see above). The genus Isohypsibius s.s. (i.e. I. prosostomus and its closest relatives) appears paraphyletic with respect to Fractonotus. However, in general, all isohypsibioid lineages clearly remain in polytomy, with only the occasional sound Bayesian posterior probabilities characterising clades with morphologically similar taxa. Therefore, the exact relationships between different isohypsibioid clades remain unsolved., Published as part of G��siorek, Piotr, Morek, Witold, Stec, Daniel, Blagden, Brian & Michalczyk, ��ukasz, 2019, Revisiting Calohypsibiidae and Microhypsibiidae: Fractonotus Pilato, 1998 and its phylogenetic position within Isohypsibiidae (Eutardigrada: Parachela), pp. 71-89 in Zoosystema 41 (6) on pages 77-82, DOI: 10.5252/zoosystema2019v41a6, http://zenodo.org/record/3718524, {"references":["RICHTERS F. 1900. - Beitrage zur Kenntnis der Fauna der Umgebung von Frankfurt a. M. Bericht der Senckenbergischen Naturforschenden gesellschaft in Frankfurt am Main 21 - 44.","MURRAY J. 1911. - Clare Island Survey. 37: Arctiscoida. Proceedings of the Royal Irish Academy 31 (37): 1 - 16.","THULIN G. 1911. - Beitage zur Kenntnis der Tardigradenfauna Schwedens. Arkiv for Zoologi 7 (16): 1 - 60.","MARCUS E. 1928. - Spinnentiere oder Arachnoidea. IV: Bartierchen (Tardigrada). Tierwelt Deutschlands und der angrenzenden Meeresteile Jena 12: 1 - 230.","CUENOT L. 1932. - Tardigrades, in LECHEVALIER P. (ed.). Faune de France 24: 1 - 96.","CUNHA A. X. 1947 da. - Tardigrados da Fauna Portuguesa III. Memorias e estudos do Museu zoologico da Universidade de Coimbra 177: 1 - 9.","CUNHA A. X. DA 1948. - Tardigrados da Fauna Portuguesa IV. Memorias e estudos do Museu zoologico da Universidade de Coimbra 188: 1 - 8.","THULIN G. 1928. - Uber die phylogenie und das system der tardigraden. Hereditas 11: 207 - 266.","MARCUS E. 1936. - Tardigrada. Das Tierreich, de Gruyter & Co., Berlin and Leipzig 66: 1 - 340.","FRANCESCHI T. 1951 - 1952. - Contributo alla conoscenza dei Tardigradi d'Italia. Bollettino dei Musei e degli Istituti Biologici dell'Universita di Genova 24 (149): 5 - 15.","MIHELCIC F. 1953. - Contribucion al conocimiento de los tardigrados con especial consideracion de los tardigrados de Osttirol (II). Anales de Edafologia y Fisiologia Vegetal 12 (5): 431 - 479.","FONTOURA A. P. 1981. - Contribution pour l'etude des tardigrades terrestres du Portugal, avec la description d'une nouvelle espece du genre Macrobiotus. Publicac o es do Instituto de Zoologia ' Dr Augusto Nobre', Faculdade de Ciencias do Porto 160: 1 - 24.","CUNHA A. X. DA 1943. - Un Tardigrade nouveau de Portugal: Hypsibius placophorus sp. n. Memorias e estudos do Museu zoologico da Universidade de Coimbra 155: 1 - 5.","MICHALCZYK L. & KACZMAREK L. 2005. - The first record of the genus Calohypsibius Thulin, 1928 (Eutardigrada: Calohypsibiidae) from Chile (South America) with a description of a new species Calohypsibius maliki. New Zealand Journal of Zoology 32: 287 - 292. https: // doi. org / 10.1080 / 03014223.2005.9518420","NELSON D. R. & MCGLOTHLIN K. L. 1996. - A new species of Calohypsibius (Phylum Tardigrada, Eutardigrada) from Roan Mountain, Tennessee-North Carolina, U. S. A. Zoological Journal of the Linnean Society 116: 167 - 174. https: // doi. org / 10.1006 / zjls. 1996.0014","BISEROV V. I. 1986. - Terrestrial water bears from the North Caucasus. 2. Eutardigrada. Zoologicheskii Zhurnal 65 (7): 981 - 993.","KACZMAREK L., ZAWIERUCHA K., SMYKLA J. & MICHALCZYK L. 2012. - Tardigrada of the Revdalen (Spitsbergen) with the descriptions of two new species: Bryodelphax parvuspolaris (Heterotardigrada) and Isohypsibius coulsoni (Eutardigrada). Polar Biology 35: 1013 - 1026. https: // doi. org / 10.1007 / s 00300 - 011 - 1149 - 0","PILATO G. 1998. - Microhypsibiidae, new family of eutardigrades, and description of the new genus Fractonotus. Spixiana 21 (2): 129 - 134.","PILATO G. 1969 a. - Evoluzione e nuova sistemazione degli Eutardigrada. Bolletino di Zoologia 36: 327 - 345."]}

TAXONOMIC ACCOUNT OF THE CALOHYPSIBIIDAE SENSU STRICTO Superfamily HYPSIBIOIDEA Pilato, 1969 (emended by Bertolani et al. 2014a) Family CALOHYPSIBIIDAE Pilato, 1969 (emended by Bertolani et al. 2014a) EMENDED DIAGNOSIS. ��� Very small eutardigrades (typically below 150 ��m) with elliptical organs on the head. Dorsum covered with irregular, multangular protuberances, and sometimes also with spines (Figs 3 C, D; 4; 5 E, F). Claws miniaturised, but not reduced, of the Calohypsibius - type, i.e. asymmetrical with respect to the sequence of primary and secondary branches (2-1-2-1), but similar in their size, with bases as large as the sum of the primary and secondary branch widths, but devoid of sutures. Pseudolunulae absent. Accessory points symmetrical (Figs 11 E; 12 C, D). Six peribuccal papulae present (Fig. 6 B). AISM asymmetrical with respect to the frontal plane, with the dorsal apophysis subdivided in two portions of different shape (Fig. 9 B). Stylet furcae of the Hypsibius - type (Fig. 10 B). Pharyngeal apophyses smaller than the tiny granular macroplacoids. Smooth eggs laid in exuviae. COMPOSITION. ��� A monotypic family, comprising the genus Calohypsibius., Published as part of G��siorek, Piotr, Morek, Witold, Stec, Daniel, Blagden, Brian & Michalczyk, ��ukasz, 2019, Revisiting Calohypsibiidae and Microhypsibiidae: Fractonotus Pilato, 1998 and its phylogenetic position within Isohypsibiidae (Eutardigrada: Parachela), pp. 71-89 in Zoosystema 41 (6) on page 83, DOI: 10.5252/zoosystema2019v41a6, http://zenodo.org/record/3718524, {"references":["PILATO G. 1969 a. - Evoluzione e nuova sistemazione degli Eutardigrada. Bolletino di Zoologia 36: 327 - 345.","BERTOLANI R., GUIDETTI R., MARCHIORO T., ALTIERO T., REBEC- CHI L. & CESARI M. 2014 a. - Phylogeny of Eutardigrada: New molecular data and their morphological support lead to the identification of new evolutionary lineages. Molecular Phylogenetics and Evolution 76: 110 - 126. https: // doi. org / 10.1016 / j. ympev. 2014.03.006"]}

TAXONOMIC ACCOUNT OF THE MICROHYPSIBIIDAE SENSU STRICTO Superfamily HYPSIBIOIDEA Pilato, 1969 (emended by Bertolani et al. 2014a) Family MICROHYPSIBIIDAE Pilato, 1998 EMENDED DIAGNOSIS. ��� Very small eutardigrades (typically below 150 ��m in length) without elliptical organs on the head. Cuticle smooth. Claws minute and asymmetrical with respect to the sequence of primary and secondary branches (2-1-2-1), with thin bases continuous with the primary branches. External and internal, and anterior and posterior claws different in shape but similar in size. Pseudolunulae absent. Accessory points symmetrical (Fig. 11 G). Peribuccal papulae not visible under PCM. AISM asymmetrical with respect to the frontal plane, with the dorsal apophysis subdivided in two portions of different shapes (Fig. 9 A). Stylet furcae of the Hypsibius - type. Pharyngeal apophyses similar in size to macroplacoids. Smooth eggs laid in exuviae. COMPOSITION. ��� A monotypic family, comprising the genus Microhypsibius., Published as part of G��siorek, Piotr, Morek, Witold, Stec, Daniel, Blagden, Brian & Michalczyk, ��ukasz, 2019, Revisiting Calohypsibiidae and Microhypsibiidae: Fractonotus Pilato, 1998 and its phylogenetic position within Isohypsibiidae (Eutardigrada: Parachela), pp. 71-89 in Zoosystema 41 (6) on pages 83-84, DOI: 10.5252/zoosystema2019v41a6, http://zenodo.org/record/3718524, {"references":["PILATO G. 1969 a. - Evoluzione e nuova sistemazione degli Eutardigrada. Bolletino di Zoologia 36: 327 - 345.","BERTOLANI R., GUIDETTI R., MARCHIORO T., ALTIERO T., REBEC- CHI L. & CESARI M. 2014 a. - Phylogeny of Eutardigrada: New molecular data and their morphological support lead to the identification of new evolutionary lineages. Molecular Phylogenetics and Evolution 76: 110 - 126. https: // doi. org / 10.1016 / j. ympev. 2014.03.006","PILATO G. 1998. - Microhypsibiidae, new family of eutardigrades, and description of the new genus Fractonotus. Spixiana 21 (2): 129 - 134.","RICHTERS F. 1900. - Beitrage zur Kenntnis der Fauna der Umgebung von Frankfurt a. M. Bericht der Senckenbergischen Naturforschenden gesellschaft in Frankfurt am Main 21 - 44.","BISEROV V. I. 1986. - Terrestrial water bears from the North Caucasus. 2. Eutardigrada. Zoologicheskii Zhurnal 65 (7): 981 - 993.","MARCUS E. 1928. - Spinnentiere oder Arachnoidea. IV: Bartierchen (Tardigrada). Tierwelt Deutschlands und der angrenzenden Meeresteile Jena 12: 1 - 230.","THULIN G. 1928. - Uber die phylogenie und das system der tardigraden. Hereditas 11: 207 - 266.","KACZMAREK L., ZAWIERUCHA K., SMYKLA J. & MICHALCZYK L. 2012. - Tardigrada of the Revdalen (Spitsbergen) with the descriptions of two new species: Bryodelphax parvuspolaris (Heterotardigrada) and Isohypsibius coulsoni (Eutardigrada). Polar Biology 35: 1013 - 1026. https: // doi. org / 10.1007 / s 00300 - 011 - 1149 - 0"]}

Bryodelphax arenosus new species (Figs. 2���4, Tables 2, 3) Material examined. All individuals were mature females. Holotype (slide no. MY.813.01) and 14 paratypes (slides MY.813.01���11), six additional processed for DNA extraction and sequencing. Type locality: Bako Peninsula, lowland coastal rainforest close to the Tajor Waterfall (1��43���24���N, 110��27���50���E; 106 m asl), moss from a tree, coll. Piotr Gąsiorek & Artur Oczkowski, 5 th August 2017. Holotype and nine paratypes deposited in the Institute of Zoology and Biomedical Research, Jagiellonian University, Krak��w, Poland (slides MY.813.01���06), three paratypes deposited in the Natural History Museum of Denmark, Copenhagen (slides MY.813.07���09), and last two paratypes deposited in the Raffles Museum of Biodiversity Research, National University of Singapore (slides MY.813.10���11). Description of the new species. Adults (i.e., from the third instar onwards, with developed gonopore; measurements in Table 2). Body yellowish, pearly opalescent; eyes absent or not visible after preparation. Both primary and secondary clavae oval and small (Figs. 2, 3C, 4A). Cirri interni and externi with poorly developed cirrophores. Cirri A long (on average reaching ⅓ of the body length, whereas Bryodelphax typically has cirrus A /body length ratio around ��; Figs. 2, 3A). Dorsal plates with faint intra-cuticular pillars visible only under 100�� magnification with immersion oil (Figs. 3C, 4), better visible on the scapular and caudal plates (Figs. 3C, 4A, C), pillars on the posterior portion of the caudal plate are also the largest (Figs. 3C, 4C). Distinct dark ridges are present on lateral portions of the scapular, paired, and caudal plates, forming characteristic and stable ornamented pattern (Figs. 3B, C, 4). Pseudopores or pores (SEM observations are needed to verify the morphology of the cuticular depressions) minute and barely visible (visible in PCM as bright dots with blurred margins; Figs. 3C, 4), sometimes absent. Pseudopores/pores distributed unevenly, i.e., on the anterocentral portion of the scapular plate 8���21 pseudopores/pores per 100 ��m 2 were observed (x=17, N=10; Fig. 4A), on the central portion of the caudal plate: 11���18 pseudopores/pores per 100 ��m 2 (x=15, N=10; Fig. 4C), whereas sparse and bleak pseudopores/pores occur rarely on other, especially paired, plates. Scapular plate deeply faceted by dark ridges, with six or eight lighter rectangular areas in the central portion (Figs. 3A, C, 4A). Scapular sutures clearly separate lateral portions from the central part, extending from the base of cirrophore A to the posterior margin of the plate (Figs. 3B, C, 4A). Paired plates divided into two roughly equal anterior and posterior parts by a transverse stripe (Figs. 3, 4B). Caudal plate mildly faceted in the central part by the same dark ridges as the scapular plate (Fig. 3A), but the faceting is fainter and often poorly visible (Figs. 3C, 4C). Median plates 1 and 2 divided transversally into two unequal parts (Figs. 2, 3), and additionally the anterior portion of median 2 has a transverse suture (Fig. 3). Median plate 3 large, triangular in shape, and with a roundish posterior edge (Figs. 2, 3). Thin, non-sculptured supplementary lateral platelets present at the levels of median plates (three pairs of platelets on each body side: a pair between scapular plate and first pair of the segmental plates, a pair between paired plates, and a pair between second pair of segmental plates and caudal plate; Fig. 3B, C). Ventral cuticle smooth, without plates or intra-cuticular pillars. Papillae on legs I and IV absent or not detectable in PCM. Large, sclerotised pedal plates IV present, with irregular margin, but never forming a dentate collar (Fig. 3C, white arrowhead). External claws of all legs smooth, internal claws with minuscule bumps near their bases, which can be spurs (Fig. 3C, insert). However, they are not measurable in PCM since barely divergent from the claw branch. No juveniles, larvae (i.e., the second and first instars, respectively) or eggs were found. Remarks. All found specimens were mature females, thus it is likely that the new species is parthenogenetic. DNA sequences. New 18S and 28S rRNA sequences were obtained for two specimens; each fragment was represented by a single haplotype and is deposited in GenBank: MH 414953 (18S rRNA, 1112 bp long), XX000000 (28S rRNA, 688 bp long). p-distances between sequenced species of Bryodelphax ranged from 0.8% to 2.8% (18S rRNA), and from 7.5% to 19.3% (28S rRNA) (Table 3). Etymology. From Latin (h) arenosus = sandy, inhabiting sand. The new species was found in coastal lowland rainforest with peculiarly sandy soil (arenosols), atypical for this biome., Published as part of G��siorek, Piotr, 2018, New Bryodelphax species (Heterotardigrada: Echiniscidae) from Western Borneo (Sarawak), with new molecular data for the genus, pp. 371-381 in Raffles Bulletin of Zoology 66 on pages 374-377, DOI: 10.5281/zenodo.5358943

Bryodelphax instabilis sp. nov. (Figs 1���23, Tables 3���5) Locus typicus. Moss on carbonate bedrock from Homole Ravine in the Pieniny Mts. (49��24'17''N, 20��32'52''E; 595 m asl), Poland. Type material. Holotype (mature female, slide no. PL.189.19), coll. Maciej Barczyk, 25th May 2016, allotype (mature male, slide no. PL.189.01), 30 paratypes (21 mature females, 7 males, one juvenile, and one larva; slides PL.189.02���32) and additional 15 paratypes (both juveniles and adults) on SEM stubs. Found together with numerous Bryodelphax parvulus Thulin, 1928 (Fig. 24), Pseudechiniscus suillus (Ehrenberg, 1853), and single specimens of Testechiniscus spitsbergensis (Scourfield, 1897). All from the same moss sample. Holotype and most of paratypes deposited in the Institute of Zoology and Biomedical Research, Jagiellonian University, Krak��w, Poland, two paratypes (slides PL.189.10���11) deposited in the Department of Zoology, Comenius University in Bratislava, Slovakia, and two paratypes (slides PL.189.22���23) deposited in the Zoological Museum, University of Copenhagen, Denmark. Additional material examined. Two specimens in a moss sample from Chuda Alp in the Tatra Mts. (slide PL.258.06), coll. Piotr Gąsiorek, 24th September 2016, deposited together with type material. 128 specimens (44 males, 43 females, 3 juveniles, and 38 specimens of unidentified sex due to unsuitable orientation of a specimen on a slide; slides 390/7���10, 390/18, 390/20, 390/23, 390/25, 390/27���30, and 390/32; deposited in the Department of Zoology, Comenius University in Bratislava, except for 13 specimens deposited in the Institute of Zoology and Biomedical Research, Jagiellonian University, Krak��w ��� slide 390/18) and 9 specimens used for SEM analyses (not deposited) in a moss sample on a rock from Rajtop��ky hill, the Branisko Mts. (48��59'42''N, 20��51'59''E; ca. 1036 m asl ��� on the hill top area), Eastern Slovakia, coll. Peter Degma, 26th July 2003. Short description of holotype (Figs 1, 10; Table 3) Adult female; colour greyish; eyes not visible (after preparation); body length 137 ��m (685); cephalic cirri and papillae, and clava not enlarged; cirrus A 26.5 ��m (132.5); claws I���III of similar size, claw IV longer; dentate collar present, with three teeth; small papilla on leg IV. Dorsal plates with sparse pseudopores/pores; ventral plates not obvious. Female gonopore rosette-shaped. Short description of allotype (Figs 2, 12; Table 4) Adult male; colour greyish; eyes not visible (after preparation); body length 123 ��m (628); cephalic cirri and papillae, and clava enlarged; cirrus A 20.7 ��m (103.0); claws I���III of similar size, claw IV longer; dentate collar present, with three teeth; small papilla on leg IV not visible. Dorsal plates with sparse pseudopores/pores; suggestion of deeper faceting on scapular and caudal plates; ventral plates present. Male gonopore simple, circular. Details of the new species. Adults (i.e. from the third instar onwards; measurements and statistics for ♀♀ in Table 3, for ♂♂ in Table 4): Body greyish; eyes absent or not visible after preparation. Mouth opening surrounded by a ring of 10 papulae (visible in SEM only). Cephalic papillae greatly enlarged in males (compare Figs 1 and 2, 4), and clavae are also more prominent (Figs 13, 17���18). Internal cirri much shorter than external (Tables 3���4, Figs 1���4, 7���9). Cirri A thinner at the tip (visible only under SEM; Fig. 17). Dorsal plates covered with sparsely distributed pseudopores or pores of large and intermediate size (visible in PCM as bright dots with margins either blurred (pseudopores) or obvious (pores); Figs 1���2, 10, 12), true pores occur infrequently, being restricted to the posterior portion of the cephalic plate and the anterior portion of the scapular plate (Fig. 13) in the proximal body part, to the posterior parts of segmental plates (Fig. 14), and the anterior margin of the caudal plate in the distal body part. Pores rarely present on all plates (Figs 11, 15���16), but are often absent (Figs 6���7). Pseudopores/pores density similar on all dorsal plates, approximate mean density 10���13 pseudopores/pores per 100 ��m 2 in females and 8���10 pseudopores/ pores per 100 ��m 2 in males (Table 5). Intracuticular pillars visible as fine dark dots under PCM (Figs 10���12). Scapular plate typically distinctly faceted with a median longitudinal fold and three smaller transverse folds (Figs 2, 12). In females, sometimes only pseudopore/pore rows mark borders of faint facets (Figs 1, 10���11). Paired plates divided into two unequal anterior and posterior parts by a transverse stripe (Figs 10���12, 14, 16). Caudal plate faceted with two evident longitudinal folds (Figs 1���2, 6, 10���12), most often the central portion of this plate is subdivided into two parts by a transverse V-suture (Figs 2, 6, 12). Median plates 1 and 2 with transverse division into two unequal parts (compare Figs 10���12, 14, 16). Median plate 3 with faint transverse suture, triangular in shape, and with a roundish posterior edge (Figs 14, 16). Poorly developed supplementary lateral platelets present at the levels of median plates (three pairs of platelets on each body side: a pair between scapular plate and first pair of the segmental plates, a pair between paired plates, and a pair between second pair of segmental plates and caudal plate; Figs 1, 10���12), devoid of pores/pseudopores (Fig. 7). Ventral cuticle typically with weakly developed plates. Ventral intracuticular pillars either absent or present and well-visible (compare Figs 4���5) at 100x oil immersion, and always less evident than dorsal (compare Figs 5 and 11). Ventral plate configuration: VII/IX:(2)-(1)-2/4-2-2/4- 2-2-2-1. Two small subcephalic longitudinal plates just below the mouth opening and one subpharyngeal plate weakly outlined and usually only visible under SEM (compare Figs 5, 8���9). Typically, rows III���VIII/IX are discernible (Figs 4���5, 8���9, 21). Rarely, ventral plates invisible under PCM (e.g. holotype). Ventral granules absent or restricted to the plate surface (Fig. 5). Papilla on legs I absent, minute papilla on legs IV present (Fig. 1). Dentate collar on legs IV with 3���6 teeth (Figs 1���2, 20, arrowheads). External claws of all legs smooth, internal claws with tiny spurs pointing strongly downwards with very small gap between spur and claw base, making them almost invisible under PCM (however, always clear under SEM; Figs 19���20). Remarks on the sexual dimorphism: Sex differences in the new species are well-marked and embrace: longer, tubbier primary and secondary ♂ clavae (compare Figs 1���2 and Tables 3���4); stronger faceting of the ♂ scapular plate (compare Figs 10���11 and 12); significantly lower pseudopore/pore density on unpaired plates in ♂ (compare ranges presented in Table 5). Juveniles (i.e. the second instar, four-clawed without gonopore): In appearance like adults, but clearly smaller (75 ��m) and with indiscernible ventral plates (Fig. 22). It is impossible to state firmly whether the species exhibits ontogenetic shifts in ventral armature or the lack of ventral plates in the first two life stages is inconstant. This issue certainly requires more studies, since changes in ventral armature occurring during development were recently detected by Gąsiorek et al. (2017). Larvae (i.e. the first instar, two-clawed without gonopore; measurements in Table 4): Median plates with poorly delineated margins, lacking pseudopores/pores; supplementary lateral platelets and pedal (leg) plates undeveloped. Pores/pseudopores present only on the anterior and posterior margins of the cephalic, scapular, segmental, and caudal plates (Fig. 3). Claws with spurs formed as in adults. Legs IV without dentate collar. Ventral plates absent (Fig. 23). Eggs unknown. Etymology: The name instabilis underlines the variability in the number of ventral plates (see Remarks), which is unusual for Bryodelphax. Moreover, often only some of the ventral plates rows are visible under the light microscope, thus an animal���s venter seems to be devoid of plates in its proximal part. Remarks. Polish and Slovak populations have same ventral plates configuration but in three mature females from the Slovak population, row V has two additional, smaller ventral plates placed in a more marginal position, and two of the specimens also have such additional plates in row III (Figs 5 and 21). Differential diagnosis. Bryodelphax instabilis sp. nov. can easily be distinguished from the other members of the weglarskae group on the basis of the scapular plate faceting and being gonochorous. Nevertheless, the new species should be compared with four most similar taxa having last six ventral plates rows configuration same as the new species, i.e. 2-2-2-2-2-1 or 2-4-2-2-2-1 (due to the instability in ventral plate arrangement). It differs specifically from: Bryodelphax iohannis Bertolani et al., 1996, by a different ventral plate configuration (VII/IX:(2)-(1)-2/4-2-2/ 4-2-2-2- 1 in the new species vs X: 2-1-1-5-2-4-2-2-2- 1 in B. iohannis), by the presence of lateral platelets (absent in B. iohannis) and by longer teeth on the dentate collar (compare Figs 1���2 with Fig. 2A in Bertolani et al. 1996); Bryodelphax parvuspolaris Kaczmarek et al., 2012, by a different ventral plate configuration (VII/IX:(2)-(1)- 2/4-2-2/4-2-2-2- 1 in the new species vs VIII: 1-1-2-2-2-2-2- 1 in B. parvuspolaris), and by well-developed dentate collar (dentate collar with poorly developed teeth in B. parvuspolaris); Bryodelphax sinensis (Pilato, 1974), by a different ventral plate configuration (VII/IX:(2)-(1)-2/4-2-2/4-2-2-2- 1 in the new species vs VII:2-2-2-2-2-2- 1 in B. sinensis), and by the presence of well-developed dentate collar on legs IV in adults (dentate collar absent in B. sinensis); Bryodelphax weglarskae (Pilato, 1972), by a different ventral plate configuration (VII/IX:(2)-(1)-2/4-2-2/4-2- 2-2- 1 in the new species vs IX:2-1-5-2-4-2-2-2- 1 in B. weglarskae), and by non-bifurcated cephalic appendages (bifurcated in B. weglarskae). Moreover, Bryodelphax instabilis sp. nov. must be primarily compared with the first described gonochoristic representative of the genus, namely Bryodelphax tatrensis (Węglarska, 1959) because it resembles the latter species when the ventral armature is faint. Bryodelphax instabilis sp. nov. is distinguished from B. tatrensis on the basis of: (1) the average pseudopore/pore size is larger in the new species in comparison with the minute pores in B. tatrensis (the difference is especially obvious between females of both species, compare Figs 10���11 and 26), (2) more pronounced faceting of the scapular plate (clearly visible median suture and typically well-developed facets in the new species vs only faint median suture, which can be absent in B. tatrensis; compare Figs 11���12 and 26��� 27), and (3) the caudal plate consisting of four facets in males of the new species instead of three in B. tatrensis males (compare Figs 2, 12 and 25, 27). Additional discriminative criteria (e.g. morphometric) could be presented when measurements of a large population of B. tatrensis become available. Unfortunately, general rarity of B. tatrensis hinders the redescription of this taxon (Dastych 1988, personal observations). General remarks on Bryodelphax in Polish Pieniny Mts. and Tatra Mts. There are four sympatric Bryodelphax spp. that inhabit Poland and occur in the Pieniny Mts. (Dastych 1988); three (with the exception of B. weglarskae) are also present in the Tatra Mts. Our new discovery confirms an eucalciphil and subalpine preference for these Central European members of the genus. In Poland, Bryodelphax spp. are frequently found in close proximity. For example, in the Pieniny Mts. B. parvulus was found in the same moss cushions as B. instabilis sp. nov. or B. weglarskae; similarly, B. parvulus and B. tatrensis co-occur in mosses in the Tatra Mts., Published as part of G��siorek, Piotr & Degma, Peter, 2018, Three Echiniscidae species (Tardigrada: Heterotardigrada) new to the Polish fauna, with the description of a new gonochoristic Bryodelphax Thulin, 1928, pp. 77-96 in Zootaxa 4410 (1) on pages 79-85, DOI: 10.11646/zootaxa.4410.1.4, http://zenodo.org/record/1221257, {"references":["Thulin, G. (1928) Uber die Phylogenie und das System der Tardigraden. Hereditas, 11, 207 - 266. https: // doi. org / 10.1111 / j. 1601 - 5223.1928. tb 02488. x","Ehrenberg, C. G. (1853) Diagnoses novarum formarum. Verhandlungen der Koniglich Preussische Akademie der Wissenschaften zu Berlin, 8, 526 - 533.","Scourfield, D. J. (1897) Contributions to the non-marine fauna of Spitsbergen. Part I. Preliminary notes, and reports on the Rhizopoda, Tardigrada etc. Proceedings of the Zoological Society of London, 65 (3), 784 - 792.","Gasiorek, P., Stec, D., Morek, w., Marnissi, J. & Michalczyk, L. (2017) The tardigrade fauna of Tunisia, with an integrative description of Bryodelphax maculatus sp. nov. (Heterotardigrada: Echiniscidae). African Zoology, 52 (2), 77 - 89. https: // doi. org / 10.1080 / 15627020.2017.1297688","Bertolani, R., Guidi, A. & Rebecchi, L. (1996) Tardigradi della Sardegna e di alcune piccole isole circum-sarde. Biogeographia, 18, 229 - 247.","Kaczmarek, L., Zawierucha, K., Smykla, J. & Michalczyk, L. (2012) Tardigrada of the Revdalen (Spitsbergen) with the descriptions of two new species: Bryodelphax parvuspolaris (Heterotardigrada) and Isohypsibius coulsoni (Eutardigrada). Polar Biology, 35, 1013 - 1026. https: // doi. org / 10.1007 / s 00300 - 011 - 1149 - 0","Pilato, G. (1974) Tre nuove specie di Tardigradi muscicoli di Cina. Animalia, 1, 59 - 68.","Pilato, G. (1972) Prime osservazioni sui tardigradi delle Isole Egadi. Bollettino delle sedute dell'Accademia gioenia di scienze naturale in Catania, 11 (5), 111 - 124.","Dastych, H. (1988) The Tardigrada of Poland. Monografie Fauny Polski, 16, 1 - 255."]}

Adropion afroglacialis sp. nov. Zawierucha, Gąsiorek & Buda Material. Holotype (slide U/8) and paratypes including 35 individuals, one simplex and two exuviae (slides numbers: U/2���9, U/12���13, U/15���20) are deposited in the Department of Animal Taxonomy and Ecology at Adam Mickiewicz University, Poznań, Poland. Five paratypes (slides UG.007.01 [U/1] and UG.007.02[U/14], and three paratypes mounted on a SEM stub) are deposited in the Institute of Zoology and Biomedical Research, Jagiellonian University, Krak��w, Poland. Type locality. Mount Stanley, Rwenzori Mountains (0��22'31''N, 29��52'40''E; 4790 m asl), Uganda. Etymology. The name afroglacialis means ���occurring on an African glacier��� and refers to the ecosystem where the new species was found. Description. Body colourless or whitish in vivo as well as in slide mounts (Fig. 2A). Cuticle covered with faint irregular granulation (granules forming small dots), more visible on dorsal side and legs, sometimes poorly visible (Figs. 2B���E), for details see Remarks. Eyes absent in mounted specimens (Figs. 2A, 3A���B). Oral cavity armature not visible under PCM (Figs. 3A���B). Stylet furcae of the Hypsibius - type, unmodified (Figs. 3A���B). Drop-like thickening absent at the junction between the buccal and the pharyngeal tube (Figs. 2A, 3A���B). Annulation of the pharyngeal tube clearly visible in DIC (Fig. 3A) as a single narrow annuli (Fig. 3D). Pharynx with three double rows of macroplacoids, microplacoid and septulum (Figs. 3A���C). The apophyses clearly separated from the 1 st macroplacoids. All macroplacoids clearly separated. Macroplacoid length sequence 2Hypsibius type, with widened bases and with obvious accessory points on the primary branches (Figs. 4A���E). On the primary branch, at the border between accessory points and primary claw branch, a thick line is visible along entire branch length. Internal and anterior claws usually with two septa dividing the claw into the basal portion, the secondary branch, and the primary branch (Fig. 4C). Claws with poorly visible pseudolunulae (Figs. 4A���D, arrowheads). Bases of all claws smooth. Cuticular bars absent. ......continued on the next page Remarks and differential diagnosis. Granulation is poorly visible or not visible on some individuals, completely undistinguishable in SEM. Granulation is invisible in SEM, but is visible in both PCM and DIC, indicating that it may be comprised of depressions (fossae) covered with cuticle. On the holotype, granulation is visible only in some areas on the dorsal side. Thus, observation of cuticle should be done carefully. In five specimens, the first macroplacoid is shorter than the second; the difference ranges between 0.1���0.2 ��m. Currently, the genus Adropion consist of twenty species (Degma et al. 2009���2017), but from a molecular point of view the genus appears polyphyletic and potentially with cryptic diversity (Bertolani et al. 2014). It is first report of the genus Adropion from Uganda (McInnes et al. 2017). Apart from the peculiar granulation on cuticle, by having three macroplacoids, microplacoid and septulum in the pharynx, the new species is most similar to the following Adropion species: A. gordonense (Pilato, Claxton & Horning, 1991), A. greveni (Dastych, 1984), A. linzhiensis (Li, 2007), and A. onorei (Pilato, Binda, Napolitano & Moncada, 2002), but it specifically differs from: - A. gordonense by clearly smaller pharyngeal apophyses (compare: Fig. 1b in Pilato et al. 1991), and absence of thickened bars under internal claws I���III. - A. greveni by absence of long bars under internal claws I���III, the absence of spines under the claws, and much shorter macroplacoid row and all macroplacoids (macroplacoid row length 7.3���11.4 ��m in A. afroglacialis sp. nov. vs 18���24 ��m in A. greveni). - A. linzhiensis by having a less elongated body (compare: Figs. 6 ���7 in Li (2007)), a different macroplacoid length sequence (2vs. 1A. linzhiensis), relatively shorter macroplacoid row (36.5���50 % in A. afroglacialis sp. nov. vs 50���57.1 % in A. linzhiensis), and relatively shorter claws I���IV (external claw secondary branches 15.0���22.6 %, internal + anterior claw primary branches 11.4���25.7 %, and internal claw secondary branches 11.6���21.4 % in A. afroglacialis sp. nov. vs 25.0���37.5 %, 25.0���37.5 %, and 22.6���37.3 % in A. linzhiensis, respectively). - A. onorei by absence of bars under internal claws I���III, and absence of spines under posterior claws IV, the stylet supports inserted more anteriorly on the buccal tube (54.5���63.5 % in A. afroglacialis sp. nov. vs 64.2��� 65.3 % in A. onorei), much shorter macroplacoid row (7.3���11.4 ��m [36.5���50 %] in A. afroglacialis sp. nov. vs 14.9���17.7 ��m [78.8���84.3 %] in A. onorei)., Published as part of Zawierucha, Krzysztof, G��siorek, Piotr, Buda, Jakub, Uetake, Jun, Janko, Karel & Fontaneto, Diego, 2018, Tardigrada and Rotifera from moss microhabitats on a disappearing Ugandan glacier, with the description of a new species of water bear, pp. 311-328 in Zootaxa 4392 (2) on pages 314-319, DOI: 10.11646/zootaxa.4392.2.5, http://zenodo.org/record/1195435, {"references":["Bertolani, R., Guidetti, R., Marchioro, T., Altiero, T., Rebecchi, L. & Cesari, M. (2014) Phylogeny of Eutardigrada: New molecular data and their morphological support lead to the identification of new evolutionary lineages. Molecular Phylogenetics and Evolution, 76, 110 - 126. https: // dx. doi. org / 10.1016 / j. ympev. 2014.03.006","McInnes, S. J., Michalczyk L. & Kaczmarek, L. (2017) Annotated zoogeography of non - marine Tardigrada. Part IV: Africa. Zootaxa, 4284 (1), 1 - 7. https: // dx. doi. org / 10.11646 / zootaxa. 4284.1.1","Dastych, H. (1984) The Tardigrada from Antarctic with descriptions of several new species. Acta Zoologica Cracoviensia, 27, 377 - 436.","Li, X. C. (2007) A new species and a newly recorded species of tardigrade (Eutardigrada: Hypsibiidae) from China (Asia). Proceedings of the Biological Society of Washington, 120, 189 - 196. https: // dx. doi. org / 10.2988 / 0006 - 324 X (2007) 120 [189: ANSAAN] 2.0. CO; 2"]}

Class EUTARDIGRADA Richters, 1926 Order PARACHELA Schuster, Nelson, Grigarick and Christensen, 1980 Superfamily HYPSIBIOIDEA Pilato, 1969 (in Marley et al. 2011) Family HYPSIBIIDAE Pilato, 1969 Subfamily ITAQUASCONINAE Rudescu, 1964 Genus Mesocrista Pilato, 1987, Published as part of G��siorek, Piotr, Stec, Daniel, Morek, Witold, Zawierucha, Krzysztof, Kaczmarek, ��ukasz, Lachowska-Cierlik, Dorota & Michalczyk, ��ukasz, 2016, An integrative revision of Mesocrista Pilato, 1987 (Tardigrada: Eutardigrada: Hypsibiidae), pp. 2803-2828 in Journal of Natural History (J. Nat. Hist.) (J. Nat. Hist.) 50 on page 2809, DOI: 10.1080/00222933.2016.1234654, http://zenodo.org/record/5189370, {"references":["Richters F. 1926. Tardigrada. Handbuch Der Zoologie. III: 58 - 61.","Pilato G. 1969. Evoluzione e nuova sistemazione degli Eutardigrada. Boll Zool. 36: 327 - 345.","Marley NJ, McInnes SJ, Sands CJ. 2011. Phylum Tardigrada: A re-evaluation of the Parachela. Zootaxa. 2819: 51 - 64.","Rudescu L. 1964. Tardigrada. Fauna Republicii Populare Romine IV: 7 - 308."]}