taxonID	type	description	language	source
039F879BFF94FF854AD8F981FE1DF8B4.taxon	description	Body cylindrical, with weakly delineated and blunt cephalic region (Fig. 3 A). Eyes present as a group of pigment granules, which dissolve in Hoyer’s medium. Entire dorsal cuticle wrinkled (Figs 4 A, 5 A, B) and with marked muscle attachment points in the middle of the trunk; posteriorly, the wrinkling changes into distinct sculpturing in the form of large, plaque-like protuberances in the caudal body portion (Figs 4 B – D, 5 A, B, E – H). The protuberances are opalescent under stereomicroscope and in PCM. Most often, the sculpturing is arranged in two belts: more anterior with smaller protuberances (rarely shaped in a rectangular, plate-like manner; Fig. 4 C), and posteriormost with large, multangular protuberances (Figs 4 B – D, 5 B). In SEM, the dorsoposterior sculpturing can be characterized as locally thickened wrinkling (Fig. 5 E – H). Moreover, flattened structures are identifiable in SEM (not visible in PCM) and interpreted as rudimentary elliptical sensory organs (Fig. 5 C, D). A pair of cribriform muscle attachment sites (Kihm et al. 2023) flanking the mouth opening present (Fig. 6 F). Six peribuccal lobes present around the mouth ring (Fig. 6 E), lamellae or papulae absent. Bucco-pharyngeal apparatus of the Ramazzottius type (Figs 3 B – D, 6 A – D). OCA absent (Fig. 6 E). Apophyses for the insertion of stylet muscles (AISMs) asymmetrical with respect to the frontal plane: dorsal apophysis is shorter, higher, and stumpy (Fig. 3 D), with a prominent caudal apex (Figs 6 A, B, D, 7 A, B). Ventral apophysis slenderer (Fig. 3 D), with more developed caudal processes (Fig. 6 C). Lateral porous areas present in the buccal crown (Figs 6 E, 7 B). Furcae unmodified, of the Hypsibius type, with well-developed condyles (Figs 3 D, 6 G, H). Buccal tube narrow, with a clear dorsal longitudinal wall thickening (Fig. 6 A, B, D), posteriorly bent and terminating with pharyngeal apophyses (Figs 3 D, 6 A – D). Pharynx circular (Fig. 3 C, D), with only two granular macroplacoids (Figs 3 B – D, 6 A – D, 7 C – F). Macroplacoids are of similar size or the first is only slightly longer than the second. Both macroplacoids with a central or subcentral constriction (Fig. 7 C – F), but only the first constriction visible in PCM (Fig. 3 B – D). Claws Ramazzottius - like (sensu Tumanov 2020), with elongated primary branches of external / posterior claws and thinned connective portions in the form of light-refracting units (Fig. 8 C – L). SEM microphotographs revealed that these units are particularly slender and flexible (Fig. 8 E, G, J – L). Posterior claws are higher than external claws I – III. Accessory points on all primary branches evident and clearly divergent (Fig. 8 C – L). Pseudolunulae at claw bases present (Fig. 8 D, H, I) but often poorly visible. Typically, pseudolunulae are most prominent under claws IV. Pulvini absent. Almost none of the bar types defined for hypsibiids (Gąsiorek et al. 2024) are present (as described by Cuénot 1929, 1932), but in some specimens, transverse basal bars on the legs I – III (Fig. 8 D) are developed asymmetrically and irregularly. These structures must lie under the cuticle as they are unidentifiable in SEM, and probably serve as muscle attachment points for retractor leg muscles. Eggs laid in exuviae (up to five eggs per exuvia were found), but not smooth. Chorion with densely granulated surface (Fig. 3 E), but without any processes. The granulation is developed already within the ovary during chorion formation in the female’s body. No specimens with spermatozoa were found. Comparison with related species Hypsibius sp. nov., the new species represented only by one population from the vicinity of Kolding (Jutland), is similar to H. scabropygus in most characters, but differently developed caudal sculpturing, which is less sclerotized, without protuberances, and resembles folding of the cerebral cortex (Fig. 4 F, G), allows for a quickdelimitationunderalightmicroscope. Itsclawlight-refracting units seem to be weakly developed (Fig. 8 A) when compared with those of H. scabropygus (Fig. 8 C – L). Due to the scarcity of material, we refrain from formal description of this species. Importantly, paratypes of H. camelopardalis also have faint light-refracting units (Fig. 8 B). The population of H. cf. camelopardalis from Italy shows that the sculpturing can be formed as typical multangular plaques (Fig. 4 E), weakly elevated above the epicuticle. Among the better-documented literature records, H. cf. scabropygus from the Baltic States (Zawierucha et al. 2014) exhibits posterior bars and certainly represents another new species. Also H. ragonesei Binda and Pilato, 1985 and H. stiliferus Abe, 2004 show posterior bars; all other bar types are absent in the H. scabropygus complex of spp. Given the morphological oddities of H. scabropygus and several other spp. (Table 2), a new hypsibiin genus is erected.	en	Gąsiorek, Piotr, SØrensen, Martin V., Lillemark, Marie Rathcke, TØttrup, Frederik LeerhØi Anders P. (2024): Too rough to be a Hypsibius: multipopulation survey accentuates the phylogenetic position and apomorphies of Hypsibius scabropygus (Eutardigrada: Hypsibiidae). Zoological Journal of the Linnean Society (Zool. J. Linn. Soc.) 202 (4): 1-18, DOI: 10.1093/zoolinnean/zlae160, URL: https://doi.org/10.1093/zoolinnean/zlae160
039F879BFF96FF844850F8F2FE76F893.taxon	description	urn: lsid: zoobank. org: act: 4516 C 76 C-F 0 EB- 443 F-A 809 - C 29 EB 43 F 38 FB.	en	Gąsiorek, Piotr, SØrensen, Martin V., Lillemark, Marie Rathcke, TØttrup, Frederik LeerhØi Anders P. (2024): Too rough to be a Hypsibius: multipopulation survey accentuates the phylogenetic position and apomorphies of Hypsibius scabropygus (Eutardigrada: Hypsibiidae). Zoological Journal of the Linnean Society (Zool. J. Linn. Soc.) 202 (4): 1-18, DOI: 10.1093/zoolinnean/zlae160, URL: https://doi.org/10.1093/zoolinnean/zlae160
039F879BFF96FF844850F8F2FE76F893.taxon	type_taxon	Type species: Hypsibius scabropygus Cuénot, 1929. Diagnosis: Hypsibiids with Ramazzottius - like claws and rigid buccal tube bent posteriorly. Only posterior bars may be present. AISMs asymmetrical, dorsal AISM shorter and higher than ventral, with prominent apex and less evident caudal processes. Pharynx with two granular macroplacoids. Rudimentary elliptical sensory organs present on head. Cuticular sculpturing well developed as protuberances at least in the dorsoposterior body portion, or protuberances cover an entire dorsum. Eggs laid within the exuviae, with granulated chorion.	en	Gąsiorek, Piotr, SØrensen, Martin V., Lillemark, Marie Rathcke, TØttrup, Frederik LeerhØi Anders P. (2024): Too rough to be a Hypsibius: multipopulation survey accentuates the phylogenetic position and apomorphies of Hypsibius scabropygus (Eutardigrada: Hypsibiidae). Zoological Journal of the Linnean Society (Zool. J. Linn. Soc.) 202 (4): 1-18, DOI: 10.1093/zoolinnean/zlae160, URL: https://doi.org/10.1093/zoolinnean/zlae160
039F879BFF96FF844850F8F2FE76F893.taxon	etymology	Etymology: From Greek para - = near, close + Hypsibius, referring to the close relationship between these genera.	en	Gąsiorek, Piotr, SØrensen, Martin V., Lillemark, Marie Rathcke, TØttrup, Frederik LeerhØi Anders P. (2024): Too rough to be a Hypsibius: multipopulation survey accentuates the phylogenetic position and apomorphies of Hypsibius scabropygus (Eutardigrada: Hypsibiidae). Zoological Journal of the Linnean Society (Zool. J. Linn. Soc.) 202 (4): 1-18, DOI: 10.1093/zoolinnean/zlae160, URL: https://doi.org/10.1093/zoolinnean/zlae160
039F879BFF96FF844850F8F2FE76F893.taxon	description	Composition: P. biscuitiformis (Bartoš, 1960) comb. nov.; P. calcaratus (Bartoš, 1935) comb. nov.; P. camelopardalis (Ramazzotti and Maucci, 1983) comb. nov.; P. macrocalcaratus (Beasley, 1988) comb. nov.; P. ragonesei (Binda and Pilato, 1985) comb. nov.; P. roanensis (Nelson and McGlothlin, 1993) comb. nov.; P. scabropygus (Cuénot, 1929) comb. nov.; P. stiliferus (Abe, 2004) comb. nov.; and P. runae (Bartoš, 1941) comb. nov. (uncertain, type specimens are unusable due to the crystallization of mounting medium, thus the transfer is done based on simplistic original description).	en	Gąsiorek, Piotr, SØrensen, Martin V., Lillemark, Marie Rathcke, TØttrup, Frederik LeerhØi Anders P. (2024): Too rough to be a Hypsibius: multipopulation survey accentuates the phylogenetic position and apomorphies of Hypsibius scabropygus (Eutardigrada: Hypsibiidae). Zoological Journal of the Linnean Society (Zool. J. Linn. Soc.) 202 (4): 1-18, DOI: 10.1093/zoolinnean/zlae160, URL: https://doi.org/10.1093/zoolinnean/zlae160
039F879BFF96FF844850F8F2FE76F893.taxon	distribution	Geographic distribution: Essentially Holarctic. Among all the species included in the genus, only H. scabropygus was recorded outside of the Holarctic region — from the Palaeotropics (McInnes 1994). The conspecificity of European and African populations has yet to be demonstrated.	en	Gąsiorek, Piotr, SØrensen, Martin V., Lillemark, Marie Rathcke, TØttrup, Frederik LeerhØi Anders P. (2024): Too rough to be a Hypsibius: multipopulation survey accentuates the phylogenetic position and apomorphies of Hypsibius scabropygus (Eutardigrada: Hypsibiidae). Zoological Journal of the Linnean Society (Zool. J. Linn. Soc.) 202 (4): 1-18, DOI: 10.1093/zoolinnean/zlae160, URL: https://doi.org/10.1093/zoolinnean/zlae160
039F879BFF96FF844850F8F2FE76F893.taxon	diagnosis	Differential diagnosis: The new genus is compared with all two-macroplacoid hypsibiid genera having a rigid buccal tube, and with Acutuncus Pilato and Binda, 1997 (in Pilato and Binda 1997 b) due to the similar mode of egg deposition in the Acutuncus mariae group (Vecchi et al. 2023). Table 3 shows the differences between all relevant taxa.	en	Gąsiorek, Piotr, SØrensen, Martin V., Lillemark, Marie Rathcke, TØttrup, Frederik LeerhØi Anders P. (2024): Too rough to be a Hypsibius: multipopulation survey accentuates the phylogenetic position and apomorphies of Hypsibius scabropygus (Eutardigrada: Hypsibiidae). Zoological Journal of the Linnean Society (Zool. J. Linn. Soc.) 202 (4): 1-18, DOI: 10.1093/zoolinnean/zlae160, URL: https://doi.org/10.1093/zoolinnean/zlae160
