taxonID	type	description	language	source
9541850EC509B326846EF931E414F897.taxon	description	Family: Doryphoribiidae Gąsiorek et al., 2019	en	Gąsiorek, Piotr (2025): Grevenius granulifer (Thulin, 1928) revised: a fresh look at one of the most intensively studied water bears (Eutardigrada: Isohypsibioidea). Organisms Diversity & Evolution 25 (1): 29-41, DOI: 10.1007/s13127-024-00658-7, URL: https://doi.org/10.1007/s13127-024-00658-7
9541850EC50BB3218787F90FE3C4FA9D.taxon	description	(Figs. 1, 2, 3, 4, 5 and 6, Table 1 with measurements of adults). Adults. Body large and stocky, clearly bent dorsoventrally (Fig. 1 A). Cuticular sculpturing well-developed dorsally throughout dorsum, extending to lateral sides and legs IV; legs I – III with weak wrinkling, mainly in the centromedian portion (Fig. 1 A). Ventral cuticle smooth, not sculptured, only with some wrinkling (Fig. 1 B). Muscle attachment points (MAPs) visible dorsolaterally on trunk (Fig. 2 A, E). Cephalic region with a different type of sculpturing comprising deep wrinkling (Fig. 2 B), smoothly passing at the level of legs I into the reticulum covering the rest of dorsum. Reticulum with thickened edges of polygons (Fig. 2 A, C, G); corners of polygons in the form of larger tubercles / granules, which sometimes may be more evident than reticulum itself (Fig. 2 D – F). Bucco-pharyngeal apparatus of the Isohypsibius type (Fig. 3). OCA visible in light microscope as black dots (Fig. 3, insert); in fact, it comprises two bands: the first containing three to five rows of small conical teeth and the second composed of one row of tear-shaped teeth (Fig. 4 A – D). AISMs of the Isohypsibius type (Figs. 3 E and 4 E). Buccal tube narrow and terminated with pharyngeal apophyses. Pharynx with three macroplacoids, of which the first and the second are roughly of the same length, and the third is longer (Fig. 3). Macroplacoids are connected by a thin cuticular list, and usually malformed, with rugged margins (Fig. 4 F – H). Especially the terminal portion of the third macroplacoid can be separated from the remaining macroplacoid by a deep groove, causing an impression that a microplacoid is present (Fig. 4 H). Peribuccal lobes absent, a continuous peribuccal lamina present (Fig. 4 A, B). Claws of the Pseudobiotus type, with secondary and primary branches similar in height (br 0.81 – 1.00, 0.93 on average; this agrees with previous data published in Gąsiorek et al. (2019): br 0.83 – 0.99, 0.91 on average). Elongated (chaliceshaped) basal tracts and prominent humps on primary branches of internal and anterior claws present (Fig. 5). Accessory points closely adjacent to primary branches. Pseudolunulae present (Fig. 5 C, E), but often weakly developed and not observable in light microscope. Short and faint cuticular bars present under internal claws I – III (Fig. 5 B – D), yet usually difficult to identify. Bars are subcuticular since not visible in SEM (Fig. 5 A). Hatchlings. 111 – 177 μm long, 144 μm on average (N = 20). Body elongated, not arched. The cuticular sculpturing in the form of delicate, rugose wrinkling (Fig. 1 C), which is uniform throughout the dorsum. At first glance, the cuticle may even seem to be smooth. The difference in sculpturing between adults and juveniles has already been noted (Ramazzotti & Maucci, 1983). Exuviae. Containing 23 – 50 oval to roundish eggs; 34 on average (N = 15). Eggs are densely packed in exuvia (Fig. 6 A) and sometimes fall out through the anterior opening made by an individual laying eggs and shedding cuticle. Chorion smooth. Remarks. Many largest adults, which probably can reach even up to 600 μm (not measured due to unsuitable position on slides), seemed to be emaciated after egg deposition and Grevenius granulifer (Thulin, 1928) revised: a fresh look at one of the most intensively studied … 35 likely die soon after the last ecdysis. This is supported by analogous behaviour in another freshwater tardigrade, Pseudobiotus megalonyx (pers. observ.). However, it is also possible that animals were exhausted by massive infection of a parasitic fungus from the genus Ballocephala (Fig. 6 B, C), as unicellular assimilative hyphae developed in many recently deceased specimens. Ballocephala is a kickxellomycotin previously reported for eutardigrades dwelling in leaf litter and sheep dung (Drechsler, 1951; Richardson, 1970).	en	Gąsiorek, Piotr (2025): Grevenius granulifer (Thulin, 1928) revised: a fresh look at one of the most intensively studied water bears (Eutardigrada: Isohypsibioidea). Organisms Diversity & Evolution 25 (1): 29-41, DOI: 10.1007/s13127-024-00658-7, URL: https://doi.org/10.1007/s13127-024-00658-7
9541850EC500B32E87A9F97DE413FF39.taxon	description	Since the year of its description (Thulin, 1928), the species was reliably recorded from many Palaearctic localities, with a detailed drawing / microphotographic documentation (Cuénot, 1932, Marcus, 1936, Rudescu, 1964, Greven & Blom, 1977, Betolani, 1982, Ramazzotti & Maucci, 1983, Maucci, 1986, Dastych, 1988). The Nearctic record by Schuster et al. (1978) was published with a SEM picture of the cuticle, which indicated that the identification was correct. Also, the record from Korea (Moon et al., 1989) seems to truly represent G. granulifer. Therefore, assuming that a biogeographic structuring is present among limnic tardigrades as in their cryptogam-dwelling relatives (Gąsiorek, 2023), the species should be tentatively regarded as Holarctic until new evidence shows otherwise. Non-Holarctic records (McInnes, 1994) are thus treated as uncertain and not considered herein. The first comprehensive DNA barcodes for G. granulifer were published in Møbjerg et al. (2007), and when cross-checked with new data, they positively verified the presence of this species in Denmark (see also above). Recently, Massa et al. (2024) introduced a division of Grevenius into four morphogroups to ease species identification. Grevenius granulifer was included within the asper morphogroup (three macroplacoids in the pharynx and no microplacoid), which is clearly separate from the annulatus morphogroup (two macroplacoids in the pharynx and no microplacoid). The latter should be probably separated from Grevenius s. s. as an independent genus. However, the species recorded by Massa et al. (2024) from the British Columbia does not belong to Grevenius, but to Isohypsibius s. s., as it was found in moss (Grevenius is primarily aquatic) and represents a classical Isohypsibius prosostomus morphotype (Dastych, 1988; Thulin, 1928) with sculpturing in the form of reticulum. A precise diagnosis of I. prosostomus, and the more precise diagnosis of Isohypsibius s. s., is still lacking due to the unavailability of modern redescription.	en	Gąsiorek, Piotr (2025): Grevenius granulifer (Thulin, 1928) revised: a fresh look at one of the most intensively studied water bears (Eutardigrada: Isohypsibioidea). Organisms Diversity & Evolution 25 (1): 29-41, DOI: 10.1007/s13127-024-00658-7, URL: https://doi.org/10.1007/s13127-024-00658-7
