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Echiniscus quadrispinosus quadrispinosus Richters, 1902 Figs 1–7, Tables 1–4, Supp. file 2 Material examined Neotype GERMANY • Taunus Mountains Range, near Königstein im Taunus; 50°11′49″ N, 08°27′15″ E; 485 m a.s.l.; 2 Sep. 2019; moss from stone, mixed forest; Johenn Sholl leg.; slide GR1/5; Department of Animal Taxonomy and Ecology, Faculty of Biology, Adam Mickiewicz University in Poznań. Paraneotypes GERMANY • 33 ♀♀, 22 ♂♂, 35 juvs, 15 undefined, 3 larvae; same collection data as for neotype; slides GR 1/1, 1/2, 1/4, 1/5, 1/7; Department of Animal Taxonomy and Ecology, Faculty of Biology, Adam Mickiewicz University in Poznań • 9 ♀♀, 6 ♂♂, 4 juvs, 2 undefined, 1 larvae; same collection data as for neotype; slides GR 1/3, 1/6; Institute of Systematics and Evolution of Animals, Polish Academy of Sciences. Remarks Animals were mounted on microscope slides in Hoyer’s medium, 45 animals prepared for SEM, and 15 prepared for molecular analyses. However, DNA sequences were obtained from two females and one male only (exoskeleton, No GR5/S, GR8/S and GR10/S) which was later mounted on microscope slide in Hoyer’s medium. Description of the neotypic population Females (measurements and statistics in Table 1, Supp. file 2 and Figs 1–4) Body orange in live specimens and transparent/light yellow after preparation. Eyes red. Apart from the head appendages which include internal and external cirri and cylindrical cephalic papillae (secondary clava), only appendage A with clava (primary clava) near its base is present (Fig. 1D). Dorsal and lateral appendages in the shape of short, and long filaments and/or spines are present at positions A-B-C-Cd- D-Dd- E (Fig. 1A–C). However, appendage B may be absent, at least on one side of the body, for more details see Morphological variability below. Dorsal plates well developed. Head and scapular plates not faceted. Under PCM, lateral portions of the scapular plate appear to be detached from the dorsal plate, forming small additional plates (one on each side of the body) (Fig. 1E, arrowheads) divided from the lateral margin of the scapular plate by a thin bright stripe. This division is formed by the narrow stripe without pores, as it is clearly visible in SEM along with a small additional plate (Fig. 1C, arrowhead). Paired plates I and II are divided into two parts – narrow anterior part and a wider posterior part – by smooth stripe without sculpture (Fig. 2A, arrows). Anterior parts most often divided longitudinally into two parts (Fig. 2A–B, indented arrowheads). Median plate 1 and 2 divided into anterior part (Fig. 2A, asterisks) and posterior part (Fig. 2A, filled arrowhead; which is especially visible in lateral position), m3 undivided (Fig. 2A, empty arrow). The terminal plate with two notches (Fig. 2A, empty arrowhead). All dorsal plates, covered with double sculpture under PCM (Fig. 2A–E), i.e., regular polygonal or roundish black ‘granules’, i.e., endocuticular pillars (0.6–1.0 µm in diameter on scapular plate and similar in size also on other plates) and white roundish or oval pores which especially in larger specimens may be merged (seen as white spots, 1.0–4.2 µm in diameter on scapular plate and similar in size also on other plates); but see below for more details. Pores absent on anterior parts of m1 and m2 (Fig. 2A, asterisks), with typical double sculpture present on the posterior parts (Fig. 2A, filled arrowheads). Central part of terminal and scapular plates with cross-like pattern, and on terminal plate a transverse line of the cross is an extension of notches (Fig. 2C–E). Sometimes, on the scapular plate, the transverse line is absent and then only the longitudinal line visible in the middle of this plate or an additional transverse line is present and the plate is poorly divided into six rectangles (Fig. 2C–D). Under SEM the plates with regularly distributed pores (Fig. 2F), which means that where the white pores visible under PCM are absent (i.e., neck plate, lateral portions of the scapular plate and anterior parts of paired plates) the plates appear to be smooth or with poorly visible granulation (Fig. 2F). Two poorly marked ventral rectangular plates, arranged transversally, are present below the head (Fig. 3A, C, arrowheads). Another two rounded square plates are present on lateral sides of the gonopore (Fig. 3B–C, arrows). Ventral cuticle possesses tiny and regular granulation (due to the presence of dense endocuticular pillars). Granulation is a little larger on the plates around the gonophore (0.3–0.4 µm diameter) than in other parts of the ventral cuticle, 0.1–0.2 µm diameter) (Fig. 3D). Outer cuticle of legs I–III with clearly visible stripes of tiny and regular granulation (0.1–0.4 µm in diameter): a thin frontal stripe on the upper part of the leg (Fig. 4A, empty arrow), a wide stripe in the central part of the leg covering frontal and lateral side of the leg (Fig. 4A, empty arrowhead) and the most distal, thin stripe above claws on the ventral side of the leg (Fig. 4A, filled indented arrowhead); white pores absent. On legs IV only frontal stripe on the upper part of the leg (just above the plate with dentate collar) (Fig. 4C, E, empty arrows) and thin stripe above claws on the ventral side of the leg are present. Triangular spine on leg I (Fig. 4B–C, filled indented arrowhead) and finger-like papilla on leg IV, present (Fig. 4D–E, filled arrow). Legs IV with dentate collar with seven to ten sharp, triangular teeth and the plate with the same sculpture as dorsal plates (Fig. 4D–E). External claws of all legs I–IV smooth, internal with spurs directed downwards (Fig. 4C–F)). The gonopore with the typical six-petal rosette (Fig. 3C, asterisk). Males (measurements and statistics in Table 2, Supp. file 2 and Fig. 5) Males are, in general, similar to females in the morphology of plates and chaetotaxy (Fig. 5A, C–D). However, there are differences in the lengths of some morphological structures (especially slightly shorter head appendages, i.e., cirri internal and external and body appendages) (compare values in Tables 1–2). On the dorsal side, regular polygonal or roundish black ‘granules’ 0.3–0.9 µm in diameter (on scapular plate and similar in size also on other plates) and white roundish or oval pores 1.0–3.1 µm in diameter (on scapular plate and similar in size also on other plates). Gonopore round and without the six-petal rosette (Fig. 5B). Juveniles – four-clawed (measurements and statistics in Table 3, Supp. file 2 and Fig. 6) In general, juveniles were similar to adults of both sexes in the morphology of plates (Fig. 6A). However, there are differences in chaetotaxy (Fig. 6A–D), in general, lack of filaments B. Moreover, appendages are shorter in juveniles than in adult females (compare values in Tables 1–3) (Fig. 6A–D). On the dorsal side, regular polygonal or roundish black ‘granules’ 0.4–0.9 µm in diameter (on scapular plate and similar in size also on other plates) and white roundish or oval pores 1.0–2.7 µm in diameter (on scapular plate and similar in size also on other plates). Gonopore absent. Larvae – two-clawed (measurements and statistics in Table 4, Supp. file 2 and Fig. 7G–H) All head and body appendages much shorter than in adults and juveniles (compare values in Tables 1–4) (Fig. 7G–H). Moreover, morphology and sculpture of plates are also different. Anterior parts of the paired plates not divided. Although, sculpture on dorsal plates composed of regular polygonal or roundish black ‘granules’ and white roundish or oval pores as in adults and juveniles. However, on head plate, anterior parts of paired plates and on entire median plate m3, pores completely absent and black ‘granules’ poorly marked. On scapular plate pores are distributed mainly on the margins of the plate and almost absent in the centre. On the dorsal side, regular polygonal or roundish black ‘granules’ 0.4–0.9 µm in diameter (on scapular plate) and white roundish or oval pores 0.7–1.4 µm in diameter (on scapular plate). Also, chaetotaxy different than adults and juveniles, i.e., A-Cd- Dd- E (Fig. 7G). Gonopore absent (Fig. 7H). Eggs Smooth, light orange and deposited in the exuviae up to 6 in one exuvium. DNA sequences We obtained good quality sequences for the applied molecular markers: 18S rRNA: GenBank: MZ798389 - MZ798396, 771 bp long; 28S rRNA: GenBank: MZ816972 - MZ816979, 715-747 bp long; ITS-2: GenBank: MZ816980 - MZ816987, 464 bp long; COI: GenBank: MZ798397 - MZ798404, 688 bp long. Morphological variability A strict chaetotaxy was analysed in 36 females, 23 males, 35 juveniles, 3 larvae. In all adult specimens, both females and males typical chaetotaxy, i.e., A-B-C-Cd- D-Dd- E was observed (Fig. 1A–B), whereas, in juveniles appendages B are most often absent (chaetotaxy: A-C-Cd- D-Dd- E). The other dorsal and lateral appendages were in general shorter in juveniles than in adults. In all studied larvae, chaetotaxy was always A-Cd- Dd- E and all appendages were much shorter than in juveniles and adults (compare values in Tables 1–4). Moreover, some modifications in chaetotaxy were observed in juveniles and adults. In two juveniles appendages B were present on both sides of the body and in five juveniles appendages B were present only on one side of the body (chaetotaxy: A-B-C-Cd- D-Dd- E) (Fig. 6A–B). Moreover, in one juvenile appendages Cd and Dd were present only on one side of the body, and in another one Dd was present on both sides and Cd only on one side (chaetotaxy: A-C-Cd- D-Dd- E) (Fig. 6C–D). In two females appendages B were present only on one side of the body (chaetotaxy: A-B-C-Cd- D-Dd- E) (Fig. 7A). Two females had additional small spines near the base of normally developed appendages B (chaetotaxy: A-B-C-Cd- D-Dd- E) (Fig. 7B–D). In other two females only appendage Cd on one side of the body was present and appendages Dd were completely absent (chaetotaxy: A-B-C-Cd- D-E ) (Fig. 7E–F). One female had very short appendage B on one side of the body (chaetotaxy: A-B-C-Cd- D-Dd- E). In another one a very short appendage B on one side of the body was present and appendages Cd and Dd were present only on one side of the body (chaetotaxy: A-B-C-Cd- D-Dd- E). Finally, in one female appendages Cd and Dd were present on only one side of the body (chaetotaxy: A-B-C-Cd- D-Dd- E). In four males appendages B were present only on one side of the body (chaetotaxy: A-B-C-Cd- D-Dd- E) (Fig. 5C–D) and in two males appendages B were absent on both sides (chaetotaxy: A-C-Cd- D-Dd- E) (Fig. 5A). Based on these assumptions chaetotaxy formula for adults and juveniles of this species is in general A-(B)- C-Cd- D-Dd- E and for larvae A-Cd- Dd- E. The other observed aberrations in chaetotaxy are only accidental. Genetic variability The obtained eight COI sequences (GenBank accession numbers: MZ798397 - MZ798404) of E. q. quadrispinosus consisted of four COI haplotypes. Haplotype 1 was found in the Norwegian (169/7 sequence, population code: 169) and German (GR8 and GR10 sequences, population code: GR) populations whereas haplotypes 2, 3 and 4 were identified in different Norwegian populations (haplotype2 – 169/8sequence, population code: 169; haplotype 3– 184/3 and 184/8 sequences, population code: 184; haplotype 4 –187/4 and 187/7 sequences, population code: 187). The value of uncorrected genetic p-distances between obtained COI haplotypes ranged from 0.2% to 0.8%. In turn, the analysis of the p-distances between E. q. quadrispinosus and compared 20 taxa of the genus Echiniscus ranged from the most similar 1.2% for E. quadrispinosus (GenBank accession number: JX683821, Vincente et al. 2013) to the least similar 21.8% for E. tantulus Gąsiorek, Bochnak, Vončina & Kristensen, 2020 (GenBank accession number: MT107427, Bochnak et al. 2020), with an average p-distance of 14.4%. In the conservative 18S rRNA gene fragment we observed no differences between our eight sequences from the German and Norwegian populations (GenBank accession numbers: MZ798389 - MZ798396) and sequences of E. quadrispinosus deposited in NCBI (GenBank accession number: MK529684). In turn, the uncorrected genetic p-distances between the other 21 taxa of the genus Echiniscus showed that the least similar was E. belloporus Gąsiorek & Kristensen, 2018 (GenBank accession number: MK529674, Gąsiorek et al. 2019a) with a genetic distance value of 3.1% and an average p-distance was 1.4%. The analysis of the p-distances between our eight sequences of 28S rRNA from the German and Norwegian populations (GenBank accession numbers: MZ816972 - MZ816979; two groups of sequences, i.e., the first consisted of GR8, GR10, 169/8, 169/9, 184/3, 184/8, 187/1 sequences and the second – one 187/8 sequence) indicated that the genetic distance was 1%. Comparison with other 19 taxa of the genus Echiniscus, for which GenBank sequences are available, are as follows: the most similar was E. quadrispinosus (GenBank accession number: MK529714, Gąsiorek et al. 2019a) with 1% value of the p-distance and the least similar was E. belloporus Gąsiorek & Kristensen, 2018 (GenBank accession numbers: MK529702, Gąsiorek et al. 2019a) – 5.4%, with an average p-distance of 2.5%. No genetic differences were observed between our eight ITS2 sequences from the German and Norwegian populations (GenBank accession numbers: MZ816980 - MZ816987). The ranges of uncorrected genetic p-distances between our sequences and the other 10 species of the genus Echiniscus indicated that the most similar was E. virginicus Riggin, 1962 (GenBank accession number: MN545756, Gąsiorek et al. 2019b) – 0.42% and the least similar was E. blumi Richters, 1903 (GenBank accession number: EF620383, Jørgensen et al. 2007) – 34.5%, with an average p-distance of 21.8%. There were no available ITS2 sequences of E. quadrispinosus in the GenBank database. Establishing of the neotype and paraneotypes of E. q. quadrispinosus The search for the type material of E. q. quadrispinosus in various collections did not bring positive results. We can probably assume that the type material of E. q. quadrispinosus no longer exists. Taking into consideration that accurate diagnoses of the species were poorly provided in the past, it is necessary to establish a neotype series of this species. For this reason, we designated the neotype and 108 paraneotypes of E. q. quadrispinosus which agree with the original description and were collected in the terra typica in the Taunus Mountain Range (Germany). The neotype series was deposited at the Department of Animal Taxonomy and Ecology, Adam Mickiewicz University in Poznań and Institute of Systematics and Evolution of Animals, Polish Academy of Sciences, Poland. All the above-mentioned statements are in accordance with the International Commission on Zoological Nomenclature (ICZN) acts dedicated to the establishment of neotype series., Published as part of Kaczmarek, Łukasz, Kayastha, Pushpalata, Gawlak, Magdalena, Mioduchowska, Monika & Roszkowska, Milena, 2022, An integrative redescription of Echiniscus quadrispinosus quadrispinosus Richters, 1902 (Heterotardigrada, Echiniscidae) from the terra typica in Taunus Mountain Range (Europe; Germany), pp. 102-124 in European Journal of Taxonomy 823 (1) on pages 105-118, DOI: 10.5852/ejt.2022.823.1819, http://zenodo.org/record/6658659, {"references":["Richters F. 1902. Neue Moosbewohner. Bericht der Senckenbergischen Naturforschenden Gesellschaft in Frankfurt am Main 1902: 23 - 26. Available from https: // www. biodiversitylibrary. org / page / 8478822 [accessed 4 May 2022].","Bochnak M., Voncina K., Kristensen R. M. & Gasiorek P. 2020. Continued exploration of Tanzanian rainforests reveals a new echiniscid species (Heterotardigrada). Zoological Studies 59: 18. https: // doi. org / 10.6620 / ZS. 2020.59 - 18","Gasiorek P. & Kristensen R. M. 2018. Echiniscidae (Heterotardigrada) of Tanzania and Uganda. Tropical Zoology 31 (3): 131 - 160. https: // doi. org / 10.1080 / 03946975.2018.1477350","Gasiorek P., Morek W., Stec D. & Michalczyk L. 2019 a. Untangling the Echiniscus Gordian knot: paraphyly of the \" arctomys group \" (Heterotardigrada: Echiniscidae). Cladistics 35 (6): 633 - 653. https: // doi. org / 10.1111 / cla. 12377","Gasiorek P., Jackson K. J., Meyer H. A., Zajac K., Nelson D. R., Kristensen R. M. & Michalczyk L. 2019 b. Echiniscus virginicus complex: the first case of pseudocryptic allopatry and pantropical distribution in tardigrades. Biological Journal of the Linnean Society 128 (4): 789 - 805. https: // doi. org / 10.1093 / biolinnean / blz 147","Jorgensen A., Mobjerg N. & Kristensen R. M. 2007. A molecular study of the tardigrade Echiniscus testudo (Echiniscidae) reveals low DNA sequence diversity over a large geographical area. Journal of Limnology 66 (1 s): 77 - 83. https: // doi. org / 10.4081 / jlimnol. 2007. s 1.77"]}

3. Pseudechiniscus angelusalas sp. nov. Roszkowska, Grobys, Bartylak & Kaczmarek (Tables 2 and 4, Figs 1, 7–9) Pseudechiniscus sp. 3 (Grobys et al. 2020) Material examined: 10 animals (holotype (female) and 9 paratypes (8 females, 1 male)) mounted on microscope slides in Hoyer’s medium and 5 prepared for barcoding. Description Animals (measurements and statistics in Tables 2 and 4) Females. Body (Fig. 7 A–B) yellow in living specimens (transparent after mounting on slides), black eyes present after mounting on slides. Apart from the head appendages (cirri interni and externi and elongated cephalic papillae [secondary clava]), only lateral cirrus A present (with finger-like clavae near the base [primary clava]). Cephalic papillae smaller than clava. Dorsal plates with small hemispherical granules/upper ends of cuticular pillars (dots by PCM) 0.3–0.7 μm in diameter, densely (spaces between granules 0.4–0.6 μm) and uniformly distributed and joined by very thin and delicate (poorly visible) striae forming indistinct hexagonal pattern (which are in fact a thin stripes positioned under epicuticle) (Fig. 7C). Granules/upper ends of cuticular pillars slightly larger in the centre of the plates. Dorsal plates typical for the genus Pseudechiniscus (cephalic plate (cp), neck plate (np), scapular plate (scp), median plates (m1, m2, m3), paired segmental plates I and II (s1, s2), pseudosegmental plate (psp) and the caudal plate (cap), see also Dorsal and ventral plates and sculpture in Grobys et al. 2020) well developed. The cp with W-shaped pattern divided into five parts. The scp divided by transversal fold which form a long narrow stripe in posterior part of the plate. This narrow stripe is sometimes divided by three longitudinal folds which form four plate parts/subplates (Fig. 7A). Besides, the entire scp divided into two parts by median longitudinal fold (Fig. 7A, empty arrow). Additionally, lateral portions of the scp appear detached from the dorsal plate, forming small platelike structures separated from the scp by a thin bright stripe. Plates m1 and m2 divided in two portions by transverse fold, plate m3 undivided (Fig. 7A, filled indented arrowheads). Laterally to the median plates, lateral intersegmental plates (lip) present. On the plates s1 and s2 darker stripes also visible (Fig. 7A, filled arrows). The psp divided by a longitudinal fold. Posterior margin of the psp straight, i.e. without projections, teeth or spines (Fig. 7A, empty indented arrowhead). The cap concave with two Y-shaped bifurcated ridges (Fig. 7A, filled arrowhead). Ventral cuticle with tiny granulation (formed by dense granules/upper ends of cuticular pillars, 0.1–0.2 μm) forming unique pattern (Figs 1, 7B, D). Ventral PGs present (granulation 0.2–0.4 μm in diameter, spaces between granules 0.2–0.3 μm) with configuration PG:I-II-III-IV-VI-VIII a (Figs 1, 7B). The female gonopore with the typical six-petal rosette. The outer cuticle on legs I–III with round patches of granulation (with larger granules but sparser in the centre and smaller and denser in peripheral parts), on legs IV with uniform wide stripes of granulation (slightly larger in the centre of these stripes) (Fig. 8). Triangular spine on leg I and dentate collar on leg IV absent. A finger-like papillae on leg IV present (Fig. 8B). External claws of all legs smooth, internal with spurs directed downwards, identical in legs I–IV (Fig. 8C). Male (Fig. 9). Dorsal plates and body appendages arranged identically like in females. Cirri interni and externi and cephalic papillae larger than in females (compare Tables 2 and 4). Cephalic papillae larger than clava. Dorsal granules/upper ends of cuticular pillars similar in size, shape and pattern to those in females but without striae (Fig. 9 A–B). Only one male was studied its body being in lateral position with the ventral side not well visible. Ventral granulation uniform (granules/upper ends of cuticular pillars, 0.2–0.4 μm) without visible pattern or the pattern poorly visible. Ventral PGs present (granulation 0.4–0.6 μm in diameter, spaces between granules 0.3–0.8 μm) with configuration PG:I-II-IV-VI-VIII g (Fig. 9C). Patches of granulation more pronounced than in females. The gonopore round and without the six-petal rosette and with indistinct PG in anterior and lateral parts. Leg sculpture similar to females but with larger granulation (Fig. 9D). Juveniles. Unknown. Larvae. Unknown. DNA sequences We obtained good quality sequences for the applied molecular markers: – COI sequence (GenBank: MN528468), 691 bp long; – ITS-2 sequence (GenBank: MN537864), 474 bp long. Etymology. The specific epithet ‘ angelus alas ’ means, in Latin, ‘angel wings’ and refers to the similarity of some ventral structures to the angel wings. Type locality. 22°37’04.5”S, 46°43’14.1”E, 1,201 m asl: Madagascar, Fianarantsoa Province, Ivohibory forest, mosses from dead wood, 04.06.2017, coll. Marta Kepel and Andrzej Kepel. Type depositories. Holotype: slide MAD87/10 and 9 paratypes (slides: MAD87/*, where the asterisk can be substituted by any of the following numbers: 1, 5, 10, 2/S 4/S, 5/S, 6/S, 7/S) are deposited at the Department of Animal Taxonomy and Ecology, Institute of Environmental Biology, Adam Mickiewicz University in Poznań, Uniwersytetu Poznańskiego 6, 61-614 Poznań, Poland. Morphological differential diagnosis* *only measurements of adult females are used in differential diagnosis Pseudechiniscus angelusalas sp. nov. differs specifically from: 1. Pse. beasleyi, by: the scp not divided in anterior part (the scp divided into four parts in Pse. beasleyi), shorter cirri interni (6.0–7.3 μm in Pse. angelusalas sp. nov. vs 10.4–15.7 μm in Pse. beasleyi), shorter cirri externi (9.1–10.4 μm in Pse. angelusalas sp. nov. vs 13.1–18.3 μm in Pse. beasleyi), different claws length arrangement (shortest claws II and III, and longest claws IV in Pse. angelusalas sp. nov. vs claws I and II shortest and III and IV longest in Pse. beasleyi) and shorter claws (see Table 4 in this paper and Table 2 in Li et al. 2007). 2. Pse. chengi, known only from China (Xue et al. 2017), by: dorsal granules joined by striae, plates m1 and m2 divided in two portions by transverse fold (unndivided in Pse. chengi), higher sp of cephalic papillae ([18.3–20.8] in Pse. angelusalas sp. nov. vs [11.8–16.6] in Pse. chegi), shorter cirri externi (9.1–10.4 μm in Pse. angelusalas sp. nov. vs 10.5–14.2 μm in Pse. chegi), higher sp of cirri A ([129.2–152.2] in Pse. angelusalas sp. nov. vs [96.0 –111.1] in Pse. chengi), higher cirrus A /body length ratio (19–22% in Pse. angelusalas sp. nov. vs 13–17% in Pse. chengi) and shorter claws I–IV and lower sp of claws II–III (see Table 4 herein and Table 2 in Xue et al. 2017). 3. Pse. clavatus, by: different shape of clavae (finger-like in Pse. angelusalas sp. nov. vs club-shaped in Pse. clavatus) and normally developed cephalic papillae (reduced in Pse. clavatus). 4. Pse. dastychi sp. nov., see Morphological differential diagnosis of Pse. dastychi sp. nov. above. 5. Pse. ehrenbergi sp. nov. see Morphological differential diagnosis of Pse. ehrenbergi sp. nov. above. 6. Pse. facettalis, known from distant localities throughout the world (McInnes 1994). Based on present study, an inaccurate description of this species makes it impossible to differentiate this taxon from Pse. angelusalas sp. nov.. See also Morphological differential diagnosis of Pse. suillus and Discussion in the paper Grobys et al. (2020). 7. Pse. indistinctus sp. nov., by: different ventral pattern (Figs 1, 7B for Pse. angelusalas sp. nov. vs Figs 1, 13 C–D for Pse. indistinctus sp. nov.), different ventral PG configuration (PG:I-II-III-IV-VI-VIII a in Pse. angelusalas sp. nov. vs PG:I-II-III-IV-V-VI-VIII a in Pse. indistinctus sp. nov.), granules on the cap similar in size to other dorsal plates (granules visibly larger on cap in comparison with other dorsal plates in Pse. indistinctus sp. nov.), higher sp of cephalic papillae (18.3–20.8 in Pse. angelusalas sp. nov. vs 15.5–17.3 in Pse. indistinctus sp. nov.), shorter cirri externi (9.1–10.4 μm in Pse. angelusalas sp. nov. vs 11.3–14.4 μm in Pse. indistinctus sp. nov.) and shorter spurs on all legs (1.3–1.6 μm in Pse. angelusalas sp. nov. vs 1.7–2.3 μm in Pse. indistinctus sp. nov.). 8. Pse. juanitae, known from Austria, Brazil (type locality), Italy and Galapagos Islands (McInnes 1994). Based on present study, an inaccurate description of this species makes it impossible to differentiate this taxon from Pse. angelusalas sp. nov.. See also Morphological differential diagnosis of Pse. suillus and Discussion in the paper Grobys et al. (2020). 9. Pse. lacyformis sp. nov., by: different ventral pattern Figs 1, 7B for Pse. angelusalas sp. nov. vs Figs 1, 10B for Pse. lacyformis sp. nov.), dorsal granules joined by striae, smaller body size (113.0–143.0 μm in Pse. angelusalas sp. nov. vs 152.0–201.0 μm in Pse. lacyformis sp. nov.), shorter cirri interni (6.0–7.3 μm, [sp=34.4–36.6] in Pse. angelusalas sp. nov. vs 10.6–14.0 μm, [sp=48.4–53.9] in Pse. lacyformis sp. nov.), shorter cirri externi (9.1– 10.4 μm, [sp=49.0–54.3] in Pse. angelusalas sp. nov. vs 10.6–14.0 μm, [sp=48.4–53.9] in Pse. lacyformis sp. nov.), shorter papillae on leg IV (2.0–2.4 μm, [sp=10.9–11.6] in Pse. angelusalas sp. nov. vs 3.2–4.2 μm, [sp=14.7–17.2] in Pse. lacyformis sp. nov.) and lower spur/branch length ratio of all claws (23–26% in Pse. angelusalas sp. nov. vs 15–20% in Pse. lacyformis sp. nov.). 10. Pse. megacephalus, by: different shape of cephalic papillae (elongated in Pse. angelusalas sp. nov. vs mushroom-like in Pse. megacephalus), absence of papilliform projection between external buccal cirri and cirri A. 11. Pse. suillus, by: a small papilla-like structure on leg I absent, different ventral pattern (Figs 1 and 7B for Pse. angelusalas sp. nov. vs Figs 1 herein and 4C–D in Grobys et al. (2020) for Pse. suillus), well-developed ventral patches of granulation, different ventral PG configuration (PG:I-II-III-IV-VI-VIII a in Pse. angelusalas sp. nov. vs PG:I-II-III-IV-VI-VIII g in Pse. suillus), dorsal granules joined by striae, shorter cirri interni (6.0–7.3 μm, [sp=34.4–36.6] in Pse. angelusalas sp. nov. vs 8.7–11.1 μm, [sp=44.0–49.6] in Pse. suillus), shorter cirri externi (9.1–10.4 μm, [sp=49.0–54.3] in Pse. angelusalas sp. nov. vs 12.0–16.8 μm, [sp=62.1–75.0] in Pse. suillus), shorter papillae on leg IV (2.0–2.4 μm, [sp=10.9–11.6] in Pse. angelusalas sp. nov. vs 3.3–4.3 μm, [sp=14.7–18.4] in Pse. suillus) and shorter claws (see Table 4 herein and Table 4 in Grobys et al. 2020). 12. Pse. xiai, known only from China (Wang et al. 2018), by: different ventral pattern (Figs 1 and 7B for Pse. angelusalas sp. nov. vs Figs 1B, F and 2E in Wang et al. (2018) for Pse. xiai), dorsal granules joined by striae, smaller body size (113.0–143.0 μm in Pse. angelusalas sp. nov. vs 179.0–215.0 μm in Pse. xiai), higher cirrus A / body length ratio (19–22% in Pse. angelusalas sp. nov. vs 13–16% in Pse. xiai), shorter papillae on leg IV (2.0–2.4 μm in Pse. angelusalas sp. nov. vs 2.5–5.1 μm in Pse. xiai) and shorter claws (see Table 4 herein and Table 1 in Wang et al. 2018). Genotypic differential diagnosis The ranges of genetic between Pse. angelusalas sp. nov. and species of the genus Pseudechiniscus, for which DNA sequences are available in GenBank, are as follows: COI: 24.5–28.5% (26.7% on average), with the most similar being Pse. dastychi sp. nov. (MN528469, present study) and the least similar being Pse. aff. suillus (MK804900, Cesari et al. 2020). ITS-2: 12.8–39.3% (29.1% on average), with the most similar being Pse. ehrenbergi sp. nov. (MN537866, present study) and the least similar being Pse. suillus (MN537863).

5. Pseudechiniscus indistinctus sp. nov. Roszkowska, Grobys, Bartylak & Kaczmarek (Table 7, Figs 1, 13–14) Pseudechiniscus sp. 5 (Grobys et al. 2020) Material examined: 77 animals (holotype and 76 paratypes (all females)) mounted on microscope slides in Hoyer’s medium, 97 animals prepared for SEM and 7 prepared for barcoding. Description Animals (measurements and statistics in Table 7) Females. Body (Fig. 13 A–B) orange-yellow in living specimens (transparent after mounting on slides), black eyes present after mounting on slides. Apart from the head appendages (cirri interni and externi and elongated cephalic papillae [secondary clava]), only lateral cirrus A present (with finger-like clavae near the base [primary clava]) (Fig. 13 A–B, D). Dorsal plates with small hemispherical granules/upper ends of cuticular pillars (dots by PCM) 0.4–1.0 μm in diameter (except of the cap), densely (spaces between granules 0.4–1.3 μm) and uniformly distributed and joined by very thin striae forming indistinct hexagonal pattern (which are in fact a thin stripes positioned under epicuticle) (Fig. 14A). Granules/upper ends of cuticular pillars slightly larger in the centre and anterior parts of the plates and much larger (1.1–1.3 μm in diameter) on the caudal plate (cap), especially in the central area between Y-shaped bifurcated ridges (Fig. 13A). Dorsal plates typical for the genus Pseudechiniscus (cephalic plate (cp), neck plate (np), scapular plate (scp), median plates (m1, m2, m3), paired segmental plates (s1, s2), pseudosegmental plate (psp) and caudal plate (cap), see also Dorsal and ventral plates and sculpture in Grobys et al. 2020) well developed. The cp with W-shaped pattern divided into five parts (Fig. 13A). The scp divided by transversal fold, which forms a long narrow stripe in posterior part of the plate. Narrow stripe sometimes divided by three longitudinal folds which form four plate parts/ subplates (Fig. 13A). The entire scp divided by median longitudinal fold into two parts (Fig. 13 A–B, empty arrow). Additionally, lateral portions of the scp appear detached from the dorsal plate, forming small plate-like structures separated from the scp by a thin bright stripe (Fig. 13C, indented arrowhead). Plates m1 and m2 divided in two portions by transverse fold, plate m3 undivided (Fig. 13 A–B, filled indented arrowheads). Laterally to the median plates, lateral intersegmental plates (lip) present. On the plates s1 and s2 darker stripes also visible (Fig. 13 A–B, filled arrows). The psp divided by a longitudinal fold. Posterior margin of the psp straight, i.e. without projections, teeth or spines (Fig. 13 A–B, empty indented arrowhead). The cap concave with two Y-shaped bifurcated ridges (Fig. 13 A–B, filled arrowhead). Ventral cuticle with tiny granulation (formed by dense granules/upper ends of cuticular pillars, 0.1–0.3 μm) forming unique pattern (Figs 1, 13 C–D). Ventral PGs present (granulation 0.3–0.6 μm in diameter, spaces between granules 0.4–0.7 μm) with configuration PG:I-II-III-IV-V-VI-VIII a (Figs 1, 13 C–D). The PG VI (in line of leg III) is not uniformly granulated, i.e. line of granules is present in the center of the PG and two smooth areas clearly visible below and above of this line (Fig. 1). The female gonopore with the typical six-petal rosette. (Fig. 13 C–D, asterisks). The outer cuticle on legs I–III with round patches of granulation (with larger granules but more sparse in the centre and smaller and denser in peripheral parts), on leg IV with uniform wide stripes of granulation (slightly larger in the centre of these stripes) (Fig. 14 B–C). Triangular spine on leg I and dentate collar on leg IV absent. A fingerlike papillae on leg IV present (Fig. 14B, arrow). External claws of all legs smooth, internal with spurs directed downwards, identical in legs I–IV (Fig. 14C). Males. Unknown. Juveniles. Unknown. Larvae. Unknown. DNA sequences We obtained good quality sequences for the applied molecular markers: – COI sequence (GenBank: MN528471), 622 bp long; – ITS-2 sequence (GenBank: MN537867), 434 bp long. Etymology. The specific epithet ‘ indistinctus ’ means, in Latin, ‘indistinct’ and refers to the delicate and poorly marked ventral sculpture. Type locality. 58°33’33’’N, 06°26’24’’E, 273 m asl: Norway, Rogaland Province, Lund Region, Førland Sletthei Landscape Protection Area, S Førlandsvatnet, Betula pubescens forest, lichen (Cladonia squamosa (Scop.) Hoffm., 1796), 12.08.2018, coll. Terje Meier, Elisabeth Stur, Karstein Hårsaker, Tommy Prestø, Lasse Topstad. Type depositories. Holotype: slide NR255/17 and 10 paratypes (slides: NR255/*, where the asterisk can be substituted by any of the following numbers: 17, 17/S, 18/S, 19/S) are deposited at the Department of Animal Taxonomy and Ecology, Institute of Environmental Biology, Adam Mickiewicz University in Poznań, Uniwersytetu Poznańskiego 6, 61- 614 Poznań, Poland; 66 paratypes (slides: NR255/*, where the asterisk can be substituted by any of the following numbers: 3, 4, 8, 10, 11, 14, 15, 16, 19, 21) are deposited at the collection NTNU University Museum, NO-7491 Trondheim, Norway. Morphological differential diagnosis* *only measurements of adult females are used in differential diagnosis Pseudechiniscus indistinctus sp. nov. differs specifically from: 1. Pse. angelusalas sp. nov. , see Morphological differential diagnosis of Pse. angelusalas sp. nov. above. 2. Pse. beasleyi, by: the scp not divided in anterior part (the scp divided into four parts in Pse. beasleyi), short- er cirri interni (7.0–9.2 μm in Pse. indistinctus sp. nov. vs 10.4–15.7 μm in Pse. beasleyi), longer clavae (4.2–5.4 μm in Pse. indistinctus sp. nov. vs 3.1–3.9 μm in Pse. beasleyi) and different claws length arrangement (shortest claws II and III, and longest claws IV in Pse. indistinctus sp. nov. vs claws I and II shortest and III and IV longest in Pse. beasleyi) and shorter claws (see Table 7 in this paper and Table 2 in Li et al. 2007). 3. Pse. chengi, known only from China (Xue et al. 2017), by: dorsal granules joined by striae, plates m1 and m2 divided in two portions by transverse fold (unndivided in Pse. chengi), longer cirri A (27.6–34.2 μm [sp=123.0– 144.9] in Pse. indistinctus sp. nov. vs 21.4–27.5 μm [sp=96.0–111.1] in Pse. chengi), higher cirrus A /body length ratio (18–22% in Pse. indistinctus sp. nov. vs 13–17% in Pse. chengi) and shorter claws I–III and lower sp of claws I–III (see Table 7 herein and Table 2 in Xue et al. 2017). 4. Pse. clavatus, by: different shape of clavae (finger like in Pse. indistinctus sp. nov. vs club-shaped in Pse. clavatus) and normally developed cephalic papillae (reduced in Pse. clavatus). 5. Pse. dastychi sp. nov., see Morphological differential diagnosis of Pse. dastychi sp. nov. above. 6. Pse. ehrenbergi sp. nov. see Morphological differential diagnosis of Pse. ehrenbergi sp. nov. above. 7. Pse. facettalis, known from distant localities throughout the world (McInnes 1994). Based on present study, an inaccurate description of this species makes it impossible to differentiate this taxon from Pse. indistinctus sp. nov.. See also Morphological differential diagnosis of Pse. suillus and Discussion in the paper Grobys et al. (2020). 8. Pse. juanitae, known from Austria, Brazil (type locality), Italy and Galapagos Islands (McInnes 1994). Based on present study, an inaccurate description of this species makes it impossible to differentiate this taxon from Pse. indistinctus sp. nov.. See also Morphological differential diagnosis of Pse. suillus and Discussion in the paper Grobys et al. (2020). 9. Pse. lacyformis sp. nov. , see Morphological differential diagnosis of Pse. lacyformis sp. nov. above. 10. Pse. megacephalus, by: different shape of cephalic papillae (elongated in Pse. indistinctus sp. nov. vs mushroom-like in Pse. megacephalus), absence of papilliform projection between external buccal cirri and cirri A. 11. Pse. suillus, by: a small papilla-like structure on leg I absent, different ventral pattern (Figs 1 and 13 C–D for Pse. indistinctus sp. nov. vs Figs 1 herein and 4C–D in Grobys et al. (2020) for Pse. suillus), well-developed ventral patches of granulation, different ventral PG configuration (PG:I-II-III-IV-V-VI-VIII a in Pse. indistinctus sp. nov. vs PG:I-II-III-IV-VI-VIII g in Pse. suillus), granules visibly larger on cap in comparison with other dorsal plates in Pse. indistinctus sp. nov. (granules on cap similar in size to other dorsal plates in Pse. suillus), dorsal granules joined by striae, lower sp of cirri interni (34.1–38.5 in Pse. indistinctus sp. nov. vs 44.0– 49.6 in Pse. suillus), lower sp of cephalic papillae (15.5–17.3 in Pse. indistinctus sp. nov. vs 19.1–24.3 in Pse. suillus), lower sp of cirri externi (54.3–59.3 in Pse. indistinctus sp. nov. vs 62.1–75.0 in Pse. suillus), shorter papillae on leg IV (2.2–2.9 μm, [sp=9.8–11.9] in Pse. indistinctus sp. nov. vs 3.3–4.3 μm, [sp=14.7–18.4] in Pse. suillus) and higher spur/branch length ratio (28–34% in Pse. indistinctus sp. nov. vs 21–27% in Pse. suillus). 12. Pse. xiai, known only from China (Wang et al. 2018), by: different ventral pattern (Figs 1 and 13 C–D for Pse. indistinctus sp. nov. vs Figs 1B, F and 2E in Wang et al. (2018) for Pse. xiai), dorsal granules joined by striae, smaller body size (140.0–166.0 μm in Pse. indistinctus sp. nov. vs 179.0–215.0 μm in Pse. xiai), higher cirrus A / body length ratio (18–22% in Pse. indistinctus sp. nov. vs 13–16% in Pse. xiai) and shorter claws (see Table 7 herein and Table 1 in Wang et al. 2018). Genotypic differential diagnosis The ranges of genetic distances between Pse. indistinctus sp. nov. and species of the genus Pseudechiniscus, for which DNA sequences are available in GenBank, are as follows: COI: 25.9–30.9% (27.9% on average), with the most similar being Pseudechiniscus sp. (KJ857005, Velasco- Castrillón et. al. 2015 (described in GenBank as Echiniscus sp.); for more details see Discussion section in Grobys et al. 2020) and the least similar being Pse. aff. suillus (MK804907, Cesari et al. 2020). ITS-2: 6.8–38.0% (25.3% on average), with the most similar being Pse. suillus sp. nov. (MN537863) and the least similar being Pse. angelusalas sp. nov. (MN537864, present study)., Published as part of Roszkowska, Milena, Grobys, Daria, Bartylak, Tomasz, Gawlak, Magdalena, Kmita, Han- Na, Kepel, Andrzej, Kepel, Marta, Parnikoza, Ivan & Kaczmarek, Łukasz, 2020, Integrative description of five Pseudechiniscus species (Heterotardigrada Echiniscidae: the suillus-facettalis complex), pp. 451-484 in Zootaxa 4763 (4) on pages 476-480, DOI: 10.11646/zootaxa.4763.4.1, http://zenodo.org/record/3762014, {"references":["Grobys, D., Roszkowska, M., Gawlak, M., Kmita, H., Kepel, A., Kepel, M., Parnikoza, I., Bartylak, T. & Kaczmarek, L. (2020) High diversity in the Pseudechiniscus suillus-facettalis complex (Heterotardigrada; Echiniscidae) with remarks on the morphology of the genus Pseudechiniscus. Zoological Journal of the Linnean Society, 188 (3), 733 - 752. https: // doi. org / 10.1093 / zoolinnean / zlz 171","Li, X. L., Wang, L. & Yu, D. (2007) The Tardigrada fauna of China with descriptions of three new species of Echiniscidae. Zoological Studies, 46 (2), 135 - 147.","Xue, J., Li, X., Wang, L., Xian, P. & Chen, H. (2017) Bryochoerus liupanensis sp. nov. and Pseudechiniscus chengi. sp. nov. (Tardigrada: Heterotardigrada: Echiniscidae) from China. Zootaxa, 4291 (2), 324 - 334. https: // doi. org / 10.11646 / zootaxa. 4291.2.5","McInnes, S. J. (1994) Zoogeographic distribution of terrestrial / freshwater tardigrades from current literature. Journal of Natural History, 28, 257 - 352. https: // doi. org / 10.1080 / 00222939400770131","Wang, L., Xue, J. & Li, X. (2018) A description of Pseudechiniscus xiai sp. nov., with a key to genus Pseudechiniscus in China. Zootaxa, 4388 (2), 255 - 264. https: // doi. org / 10.11646 / zootaxa. 4388.2.7","Cesari, M., Montanari, M., Kristensen, R. M., Guidetti, R., Bertolani, R. & Rebecchi, L. (2020) An integrated study of the biodiversity within Pseudechiniscus suillus - facettalis group (Heterotardigrada, Echiniscidae). Zoological Journal of the Linnean Society, 188 (3), 717 - 732. https: // doi. org / 10.1093 / zoolinnean / zlz 045"]}

1. Pseudechiniscus ehrenbergi sp. nov. Roszkowska, Grobys, Bartylak & Kaczmarek (Tables 1–2, Figs 1–4) Pseudechiniscus sp. 1 (Grobys et al. 2020) Pseudechiniscus aff. suillus [Ca6] MONGOLIA C 2595-V06 (Cesari et al. 2020) Material examined: 35 animals (holotype (female) and 34 paratypes (31 females, 3 males)) mounted on microscope slides in Hoyer’s medium, 30 animals prepared for SEM and 5 prepared for barcoding. Description Animals (measurements and statistics in Tables 1 and 2) Females. Body (Figs 1, 2) yellow-orange in living specimens (transparent after mounting on slides), black eyes present after mounting on slides. Apart from the head appendages (cirri interni and externi and spherical cephalic papillae [secondary clavae]), only lateral cirrus A present (with finger-like [primary] clavae near the base) (Fig. 2B). Cephalic papillae smaller than clavae. Dorsal plates with small hemispherical granules/upper ends of cuticular pillars (dots in LM) 0.3–0.8 μm in di- ameter, densely (spaces between granules 0.2–1.1 μm) and uniformly distributed and not joined by striae (Figs 2A, 3 A–B). Granules/upper ends of cuticular pillars slightly larger in the centre of the plates. Dorsal plates typical for the genus Pseudechiniscus (cephalic plate (cp), neck plate (np) scapular plate (scp), median plates (m1, m2, m3), paired segmental plates I and II (s1, s2), pseudosegmental plate (psp), and the caudal plate (cap), see also Dorsal and ventral plates and sculpture in Grobys et al. (2020)) well developed. The cp with W-shaped pattern divided into five parts. The scp divided by a transversal fold, forming a long narrow stripe in posterior part of the plate. This narrow stripe often divided by three longitudinal folds resulting in four parts/subplates (Fig. 2A). Entire scp divided by median longitudinal fold into two parts (Fig. 2 A–B, empty arrow). Lateral portions of the scp appear detached from the dorsal plate, forming small plate-like structures separated from the scp by a thin bright stripe (Fig. 3A, indented arrowhead). Plates m1 and m2 divided in two portions by transverse fold, plate m3 undivided (Fig. 2 A–B, filled indented arrowheads). Laterally to the median plates, lateral intersegmental plates (lip) present. On the plates s1 and s2 darker stripes (folds in SEM) visible (Fig. 2 A–B, filled arrows). The psp divided by a longitudinal fold. Posterior margin of psp straight, i.e. without projections, teeth or spines (Fig. 2 A–B, empty indented arrowheads). The cap concave with two Y-shaped bifurcated ridges (Fig. 2B, filled arrowhead). Ventral cuticle with tiny granulation (formed by dense granules/upper ends of cuticular pillars, 0.1–0.4 μm) forming a unique pattern (Figs 1, 2 C–D). Ventral patches of granulation present (granulation 0.5–0.6 μm in diameter, spaces between granules 0.5–1.0 μm) with configuration PG:I-II-III-IV-VI-VIII g (Figs 1, 2 C–D). The female gonopore with the typical six-petal rosette. (Fig. 2 C–D, asterisks). The outer cuticle on legs I–III with round patches of granulation (with larger granules but more sparse in the centre and smaller and denser in peripheral parts), on legs IV with uniform wide stripes of granulation (slightly larger in the centre of these stripes) (Fig. 3 C–D). Triangular spine on leg I absent, instead a small papilla-like structure present, but very hardly visible under LM (Figs 2D, 3C filled arrowheads). Dentate collar on leg IV absent. A finger-like papillae on leg IV present (Fig. 3D, filled arrow). External claws of all legs smooth, internal with spurs directed downwards, identical on legs I–IV (Fig. 3E). Males (Figs 1, 4). Body slenderer than in females. Dorsal plates and body appendages arranged identically like in females. Head appendages clearly longer than in females (compare Tables 1 and 2). Dorsal granules/upper ends of cuticular pillars larger than in females (0.4–1.3 μm in diameter), densely (spaces between granules 0.5–1.3 μm) and uniformly distributed, not joined by striae (Fig. 4 A–C). Ventral pattern (granules/upper ends of cuticular pil- lars, 0.1–0.4 μm) of different shape than in females (Figs 1, 4 D–E). Patches of granulation with configuration PG: I-II-III-IV-VI-VIII a (granules/upper ends of cuticular pillars, 0.4–0.7 μm) (Figs 1, 4 D–E). The gonopore round and without six-petal rosette and with more distinct PG in anterior and lateral parts (Fig. 4E). The leg sculpture similar to females but with larger granulation (Fig. 4F). Triangular spine on leg I absent, instead a small papilla-like structure present, but very hardly visible under LM (Fig. 4F, filled arrowhead). Juveniles. Unknown. Larvae. Unknown. PG PG PG PG PG PG PG PG . suillus (Ehrenberg, 1853) Pse. ehrenbergi sp. nov. Pse. ehrenbergi sp. nov. Pse. dastychi sp. nov. female female male female Pse. angelusalas sp. nov. Pse. lacyformis sp. nov. Pse. indistinctus sp. nov. female female female DNA sequences We obtained good quality sequences for the applied molecular markers: - COI sequence (GenBank: MN528470), 686 bp long; - ITS-2 sequence (GenBank: MN537866), 405 bp long. Etymology. We dedicate this species to the distinguished German zoologist Christian Gottfried Ehrenberg, who described Pse. suillus, the first species in the genus Pseudechiniscus. Type locality. 45°53’10’’N, 10°50’11’’E, 140 m asl: Italy, Trentino Province, Riva del Garda, on the way to the Bastion, moss from dead wood, 07.07.2016, coll. Daria Grobys and Hanna Kmita. Type depositories. Holotype: slide IT40/7 and 25 paratypes (slides: IT40/*, where the asterisk can be substituted by any of the following numbers: 3, 7, 9, 3/S, 4/S, 5/S, 6/S, 7/S) are deposited at the Department of Animal Taxonomy and Ecology, Institute of Environmental Biology, Adam Mickiewicz University in Poznań, Uniwersytetu Poznańskiego 6, 61-614 Poznań, Poland; 16 paratypes (slide IT40/10) are deposited at the collection of Binda and Pilato, Museum of the Department of Animal Biology ‘Marcello La Greca’, University of Catania, Italy. Morphological differential diagnosis* *only measurements of adult females are used in differential diagnosis Pseudechiniscus ehrenbergi sp. nov. differs specifically from: 1. Pse. angelusalas sp. nov., known only from Madagascar (the present study), by: a small papilla-like structure on leg I present, different ventral pattern (Figs 1, 2 C–D for Pse. ehrenbergi sp. nov. vs Figs 1, 7B for Pse. angelusalas sp. nov.), different position of PG VIII (PG VIII placed around gonopore in Pse. ehrenbergi sp. nov. vs PG VIII placed only above gonopore in Pse. angelusalas sp. nov.), lower sp of cephalic papillae (12.8–17.1 in Pse. ehrenbergi sp. nov. vs 18.3–20.8 in Pse. angelusalas sp. nov.), lower sp of clavae (17.7–20.6 in Pse. ehrenbergi sp. nov. vs 22.4–25.7 in Pse. angelusalas sp. nov.), lower sp of cirrus A (95.1–112.6 in Pse. ehrenbergi sp. nov. vs 129.2–152.2 in Pse. angelusalas sp. nov.), lower cirrus A /body length ratio (14–17% in Pse. suillus vs 19–22% in Pse. angelusalas sp. nov.), higher cirri interni / externi length ratio (72–85% in Pse. ehrenbergi sp. nov. vs 66–71% in Pse. angelusalas sp. nov.), longer papillae on leg IV (2.8–3.3 μm in Pse. ehrenbergi sp. nov. vs 2.0–2.4 μm in Pse. angelusalas sp. nov.) and longer claws (see Tables 1 and 4). 2. Pse. beasleyi, known only from China (Li et al. 2007), by: scp not divided in anterior part (scp divided into four parts in Pse. beasleyi), smaller granules of dorsal sculpture (0.3–0.8 μm in Pse. ehrenbergi sp. nov. vs up to 1.6 μm in Pse. beasleyi), shorter cirri interni and externi (7.1–8.6 and 9.0–11.7 μm respectively in Pse. ehrenbergi sp. nov. vs 10.4–15.7 and 13.1–18.3 μm respectively in Pse. beasleyi), different claw lengths arrangement (shortest claws II and III, and longest claws IV in Pse. ehrenbergi sp. nov. vs shortest claws I and II, and longest claws III and IV in Pse. beasleyi) and shorter claws (see Table 1 in this paper and Table 2 in Li et al. 2007). 3. Pse. chengi, known only from China (Xue et al. 2017), by: a small papilla-like structure on leg I present, plates m1 and m2 divided in two portions by transverse fold (unndivided in Pse. chengi), shorter claws I–III and lower sp of claws I–II (see Table 1 herein and Table 2 in Xue et al. 2017). 4. Pse. clavatus, known only from Spain (Mihelčič 1955), by: different shape of clavae (finger-like in Pse. ehrenbergi sp. nov. vs club-shaped in Pse. clavatus) and normally developed cephalic papillae (reduced in Pse. clavatus). 5. Pse. dastychi sp. nov., known only from Antarctica (Dastych 1984 and present study), by: a small papilla-like structure on leg I present, different ventral pattern (Figs 1, 2 C–D for Pse. ehrenbergi sp. nov. vs Figs 1, 5 C–D for Pse. dastychi sp. nov.), different ventral PG configuration (PG:I-II-III-IV-VI-VIII g in Pse. ehrenbergi sp. nov. vs PG:I-II-III-IV-V-VI-VII-VIII a in Pse. dastychi sp. nov.), dorsal granules not joined by striae, narrower scp (20.3–26.6 μm in Pse. ehrenbergi sp. nov. vs 27.5–33.0 μm in Pse. dastychi sp. nov.), shorter cirri interni (7.1–8.6 μm in Pse. ehrenbergi sp. nov. vs 10.4–12.7 μm in Pse. dastychi sp. nov.), shorter cirri externi (9.0–11.7 μm [sp=39.6–49.6] in Pse. ehrenbergi sp. nov. vs 15.9–19.1 μm [sp=54.7–61.4] in Pse. dastychi sp. nov.), shorter cirri A (21.6–26.8 μm [sp=95.1–112.6] in Pse. ehrenbergi sp. nov. vs 40.0–45.0 μm [sp=135.2–149.1] in Pse. dastychi sp. nov.), lower cirrus A /body length ratio (14–17% in Pse. ehrenbergi sp. nov. vs 22–26% in Pse. dastychi sp. nov.), higher cirri interni/externi length ratio (72–85% in Pse. ehrenbergi sp. nov. vs 62–69% in Pse. dastychi sp. nov.) and shorter claws (see Tables 1 and 3). 6. Pse. facettalis, known from distant localities throughout the world (McInnes 1994). Based on present study, an inaccurate description of this species makes it impossible to differentiate this taxon from Pse. ehrenbergi sp. nov.. See also Morphological differential diagnosis of Pse. suillus and Discussion in the paper Grobys et al. (2020). 7. Pse. indistinctus sp. nov., known only from Norway (present study), by: a small papilla-like structure on leg I present, different ventral pattern (Figs 1, 2 C–D for Pse. ehrenbergi sp. nov. vs Figs 1, 13 C–D for Pse. indistinctus sp. nov.), different ventral PG configuration (PG:I-II-III-IV-VI-VIII g in Pse. ehrenbergi sp. nov. vs PG: I-II-III-IV-V-VI-VIII a in Pse. indistinctus sp. nov.), granules on cap similar in size to other dorsal plates (granules visibly larger on cap in comparison with other dorsal plates in Pse. indistinctus sp. nov.), lower sp of cirri externi (39.6–49.6 in Pse. ehrenbergi sp. nov. vs 54.3–59.3 in Pse. indistinctus sp. nov.), shorter cirri A (21.6–26.8 μm [sp=95.1–112.6] in Pse. ehrenbergi sp. nov. vs 27.6–34.2 μm [sp=123.0–144.9] in Pse. indistinctus sp. nov.), lower cirrus A /body length ratio (14–17% in Pse. ehrenbergi sp. nov. vs 18–22% in Pse. indistinctus sp. nov.), higher cirri interni/externi length ratio (72–85% in Pse. ehrenbergi sp. nov. vs 62–70% in Pse. indistinctus sp. nov.) and lower spur/branch length ratio of all claws (17–24% in Pse. ehrenbergi sp. nov. vs 28–34% in Pse. indistinctus sp. nov.). 8. Pse. juanitae, known from Austria, Brazil (type locality), Italy and Galapagos Islands (McInnes 1994). Based on present study, an inaccurate description of this species makes it impossible to differentiate this taxon from Pse. ehrenbergi sp. nov.. See also Morphological differential diagnosis of Pse. suillus and Discussion in the paper Grobys et al. (2020). 9. Pse. lacyformis sp. nov., known only from Norway (the present study), by: a small papilla-like structure on leg I present, different ventral pattern (Figs 1, 2 C–D for Pse. ehrenbergi sp. nov. vs Figs 1, 10B for Pse. lacyformis sp. nov.), different position of PG VIII (PG VIII placed around gonopore in Pse. ehrenbergi sp. nov. vs PG VIII placed above gonopore in Pse. lacyformis sp. nov.), shorter cirri interni (7.1–8.6 μm [sp=32.3–37.4] in Pse. suillus vs 10.6–14.0 μm [sp=48.4–53.9] in Pse. lacyformis sp. nov.), shorter cirri externi (9.0–11.7 μm [sp=39.6–49.6] in Pse. suillus vs 14.1–19.4 μm [sp=66.5–77.8] in Pse. lacyformis sp. nov.), shorter clavae (4.0–4.7 μm [sp=17.7– 20.6] in Pse. ehrenbergi sp. nov. vs 5.1–6.4 μm [sp=23.4–26.8] in Pse. lacyformis sp. nov.), lower sp of cirrus A (95.1–112.6 in Pse. ehrenbergi sp. nov. vs 114.1–142.5 in Pse. lacyformis sp. nov.) and lower sp of papillae on leg IV (11.3–13.7 in Pse. ehrenbergi sp. nov. vs 14.7–17.2 in Pse. lacyformis sp. nov.). 10. Pse. megacephalus, known only from Austria (type locality) and Turkey (McInnes 1994), by: different shape of cephalic papillae (spherical in Pse. ehrenbergi sp. nov. vs mushroom-like in Pse. megacephalus), absence of papilliform projection between external buccal cirri and cirri A. 11. Pse. suillus, known only from Italy and Portugal (Grobys et al. 2020), by: different ventral pattern (Figs 1, 2 C–D for Pse. ehrenbergi sp. nov. vs Figs 1 herein and 4C–D in Grobys et al. (2020) for Pse. suillus), well developed ventral patches of granulation, shorter cirri interni (7.1–8.6 μm [sp=32.3–37.4] in Pse. ehrenbergi sp. nov. vs 8.7–11.1 μm [sp=44.0–49.6] in Pse. suillus), lower sp of cephalic papillae (12.8–17.1 in Pse. ehrenbergi sp. nov. vs 19.1–24.3 in Pse. suillus), shorter cirri externi (9.0–11.7 μm [sp=39.6–49.6] in Pse. ehrenbergi sp. nov. vs 12.0–16.8 μm [sp= 62.1–75.0] in Pse. suillus) lower sp of clavae (17.7–20.6 in Pse. ehrenbergi sp. nov. vs 20.9–26.8 in Pse. suillus), shorter cirrus A (21.6–26.8 μm [sp=95.1–112.6] in Pse. ehrenbergi sp. nov. vs 28.4–34.4 μm [sp=134.9–156.9] in Pse. suillus) and lower sp of papillae on leg IV (11.3–13.7 in Pse. ehrenbergi sp. nov. vs 14.7–18.4 in Pse. suillus). 12. Pse. xiai, known only from China (Wang et al. 2018), by: different ventral pattern (Figs 1, 2 C–D for Pse. ehrenbergi sp. nov. vs Figs 1B, F and 2E in Wang et al. 2018 for Pse. xiai) and higher cirrus A /body length ratio (17–21% in Pse. ehrenbergi sp. nov. vs 13–16% in Pse. xiai). Genotypic differential diagnosis The ranges of genetic distances between Pse. ehrenbergi sp. nov. and species of the genus Pseudechiniscus, for which DNA sequences are available in GenBank, are as follows: COI: 0.4–29.3% (22.4% on average), with the most similar being Pse. aff. suillus (MK804906, Cesari et al. 2020) and the least similar being Pseudechiniscus sp. (KJ857008, Velasco-Castrillón et. al. 2015 (described in Gen- Bank as Echiniscus sp.); for more details see Discussion section in Grobys et al. 2020). ITS-2: 7.7–35.5% (25.1% on average), with the most similar being Pse. dastychi sp. nov. (MN537865, present study) and the least similar being Pse. lacyformis sp. nov. (MN537868, present study).

4. Pseudechiniscus lacyformis sp. nov. Roszkowska, Grobys, Bartylak & Kaczmarek (Tables 5–6, Figs 1, 10–12) Pseudechiniscus sp. 4 (Grobys et al. 2020) Material examined: 15 animals (holotype (female) and 14 paratypes (11 females, 3 males) mounted on microscope slides in Hoyer’s medium and 4 prepared for barcoding. Description Animals (measurements and statistics in Table 5 and 6) Females. Body (Fig. 10A) orange-yellow in living specimens (transparent after mounting on slides), black eyes present after mounting on slides. Apart from the head appendages (cirri interni and externi and spherical cephalic papillae [secondary clava]), only lateral cirrus A present (with finger-like clavae near the base [primary clava]) (Fig. 10 A–C). Cephalic papillae smaller than clava. Dorsal plates with small hemispherical granules/upper ends of cuticular pillars (dots by PCM) 0.4–0.9 μm in diameter, densely (spaces between granules 0.5–0.9 μm) and uniformly distributed and not joined by striae (Fig. 10 C–D). Granules/upper ends of cuticular pillars slightly larger in the centre of the plates. On all dorsal plates, sometimes poorly visible darker stripes also present (Fig. 10A, C–D, filled arrows). Dorsal plates typical for the genus Pseudechiniscus (cephalic plate (cp), scapular plate (scp), median plates (m1, m2, m3), paired segmental plates I and II (s1, s2), pseudosegmental plate (psp) and the caudal plate (cap), see also Dorsal and ventral plates and sculpture in Grobys et al. (2020)) well developed. The cp with W-shaped pattern divided into five parts (Fig. 10A). The scp divided by transversal fold which forms a long narrow stripe in posterior part of the plate. Narrow stripe divided by three longitudinal folds forming four plate parts/subplates (Fig. 10A, C). Besides, the entire scp divided by median longitudinal fold into two parts (Fig. 10A, C, empty arrow). Additionally, lateral portions of the scp appear detached from the dorsal plate, forming small plate-like structures separated from the scp by a thin bright stripe (Fig. 10A, asterisk). Plates m1 and m2 divided in two portions by transverse fold, plate m3 undivided (Fig. 10A, filled indented arrowheads). Laterally to median plates, lateral intersegmental plates (lip) present. On the plates s1 and s2 darker stripes also visible (Fig. 10A, filled arrows). The psp divided by a longitudinal fold. Posterior margin of the psp plate straight, i.e. without projections, teeth or spines (Fig. 10A, empty indented arrowhead). The cap concave with two Y-shaped bifurcated ridges (Fig. 10A, filled arrowhead). Ventral cuticle with tiny granulation (formed by dense granules/upper ends of cuticular pillars, 0.2–0.4 μm) forming unique pattern (Figs 1, 10B, E). Ventral PGs present (granulation 0.4–0.8 μm in diameter, spaces between granules 0.4–0.8 μm) with configuration PG:I-II-III-IV-VI-VIII a (Figs 1, 10B). The female gonopore with the typical sixpetal rosette. (Fig. 10B). The outer cuticle on legs I–III with round patches of granulation (with larger granules but sparser in the centre and smaller and denser in peripheral parts), on leg IV with uniform wide stripes of granulation (slightly larger in the centre of these stripes) (Fig. 11). Triangular spine on leg I and dentate collar on leg IV absent. A finger-like papillae on leg IV present (Fig. 11A, arrow). External claws of all legs smooth, internal with spurs directed downwards, identical in legs I–IV (Fig. 11B). Males (Fig. 12). Dorsal plates and body appendages arranged identically like in females. Dorsal granules/upper ends of cuticular pillars similar in size, shape and pattern to those in females (Fig. 12 A–B). Ventral granulation pattern similar in shape and size to females, but less visible (Fig. 12 C–D). Ventral PGs present (granulation 0.3–0.6 μm in diameter, spaces between granules 0.2–0.5 μm) with configuration PG:I-II-IV-VI-VIII a (Fig. 12 C–D). The gonopore round and without the six-petal rosette and with the PG in anterior and lateral parts. Leg sculpture similar to females (Fig. 12E). Juveniles. Unknown. Larvae. Unknown. DNA sequences We obtained good quality sequences for the applied molecular markers: – COI sequence (GenBank: MN528472), 689 bp long; – ITS-2 sequence (GenBank: MN537868), 281 bp long. Etymology. The specific epithet ‘ lacy ’ means, in Latin, ‘lacy’ and refers to the complicated and lace-like structure of the ventral sculpture. Type locality. 59°46’12’’N, 10°28’17’’E, 72 m asl: Norway, Buskerud Province, Røyken Region, Bøsnipa Nature Reserve, at Slemmestadveien, calcareous Tilia cordata forest, lichen, 10.08.2018, coll. Torbjørn Ekrem, Ingemar Jönsson, Terje Meier, Tommy Prestø, Lasse Topstad. Type depositories. Holotype: slide NR135/6/S and 8 paratypes (slides: NR135/*, where the asterisk can be substituted by any of the following numbers: 2, 6/S, 8/S, 9/S, 10/S) are deposited at the Department of Animal Taxonomy and Ecology, Institute of Environmental Biology, Adam Mickiewicz University in Poznań, Uniwersytetu Poznańskiego 6, 61-614 Poznań, Poland; 6 paratypes (slides: NR255/*, where the asterisk can be substituted by any of the following num- bers: 1, 3) are deposited at the collection NTNU University Museum, NO-7491 Trondheim, Norway. Morphological differential diagnosis* *only measurements of adult females are used in differential diagnosis Pseudechiniscus lacyformis sp. nov. differs specifically from: 1. Pse. angelusalas sp. nov. , see Morphological differential diagnosis of Pse. angelusalas sp. nov. above. 2. Pse. beasleyi, by: the scp not divided in anterior part (the scp divided into four parts in Pse. beasleyi), longer clavae (5.1–6.4 μm in Pse. lacyformis sp. nov. vs 3.1–3.9 μm in Pse. beasleyi) and different claws length arrangement (shortest claws II and III, and longest claws IV in Pse. lacyformis sp. nov. vs shorter claws I and II, and longest claws III and IV in Pse. beasleyi). 3. Pse. clavatus, by: different shape of clavae (finger-like in Pse. lacyformis sp. nov. vs club-shaped in Pse. clavatus) and normally developed cephalic papillae (reduced in Pse. clavatus). 4. Pse. dastychi sp. nov., see Morphological differential diagnosis of Pse. dastychi sp. nov. above. 5. Pse. ehrenbergi sp. nov. see Morphological differential diagnosis of Pse. ehrenbergi sp. nov. above. 6. Pse. facettalis, known from distant localities throughout the world (McInnes 1994). Based on present study, an inaccurate description of this species makes it impossible to differentiate this taxon from Pse. lacyformis sp. nov.. See also Morphological differential diagnosis of Pse. suillus and Discussion in the paper Grobys et al. (2020). 7. Pse. indistinctus sp. nov., by: different ventral pattern (Figs 1, 10B for Pse. lacyformis sp. nov. vs Figs 1, 13 C–D for Pse. indistinctus sp. nov.), different ventral PG configuration (PG:I-II-III-IV-VI-VIII a in Pse. lacyformis sp. nov. vs PG:I-II-III-IV-V-VI-VIII a in Pse. indistinctus sp. nov.), granules on the cap similar in size to other dorsal plates (granules visibly larger on the cap in comparison with other dorsal plates in Pse. indistinctus sp. nov.), longer cirri interni (10.6–14.0 μm, [sp=48.4–53.9] in Pse. lacyformis sp. nov. vs 7.0–9.2 μm, [sp=34.1–38.5] in Pse. indistinctus sp. nov.), higher sp of cirri externi (66.5–77.8 in Pse. lacyformis sp. nov. vs 54.3–54.9 in Pse. indistinctus sp. nov.), higher sp of clavae (23.4–26.8 in Pse. lacyformis sp. nov. vs 19.3–22.5 in Pse. indistinctus sp. nov.), longer papillae on leg IV (3.2–4.2 μm, [sp=14.7–17.2] in Pse. lacyformis sp. nov. vs 2.2–2.9 μm, [sp=9.8–11.9] in Pse. indistinctus sp. nov.) and lower spur/branch length ratio of all claws (15–20% in Pse. lacyformis sp. nov. vs 28–34% in Pse. indistinctus sp. nov.). 8. Pse. juanitae, known from Austria, Brazil (type locality), Italy and Galapagos Islands (McInnes 1994). Based on present study, an inaccurate description of this species makes it impossible to differentiate this taxon from Pse. lacyformis sp. nov.. See also Morphological differential diagnosis of Pse. suillus and Discussion in the paper Grobys et al. (2020). 9. Pse. megacephalus, by: different shape of cephalic papillae (spherical in Pse. lacyformis sp. nov. vs mushroom-like in Pse. megacephalus), absence of papilliform projection between external buccal cirri and cirri A. 10. Pse. suillus, see by: a small papilla-like structure on leg I absent, different ventral pattern (Figs 1 and 10B for Pse. lacyformis sp. nov. vs Fig. 1 herein and 4C–D in Grobys et al. (2020) for Pse. suillus), well-developed ventral patches of granulation, different position of PG VIII (PG VIII placed above gonopore in Pse. lacyformis sp. nov. vs PG VIII placed around gonopore in Pse. suillus) and lower spur/branch length ratio (15–20% in Pse. lacyformis sp. nov. vs 21–27% in Pse. suillus). 11. Pse. xiai, known only from China (Wang et al. 2018), by: different ventral pattern (Figs 1 and 10B for Pse. lacyformis sp. nov. vs Figs 1B, F and 2E in Wang et al. (2018) for Pse. xiai). 12. Pse. chengi, known only from China (Xue et al. 2017), by: plates m1 and m2 divided in two portions by transverse fold (unndivided in Pse. chengi), longer cirri interni (10.6–14.0 μm [sp=48.4 –53.9] in Pse. lacyformis sp. nov. vs 3.2–8.5 μm [sp=13.2 –38.1] in Pse. chegi), higher sp of cirri externi ([66.5–77.8] in Pse. lacyformis sp. nov. vs [45.9–56.1] in Pse. chegi), higher sp of cirri A ([114.1–142.5] in Pse. lacyformis sp. nov. vs [96.0 –111.1] in Pse. chengi). Genotypic differential diagnosis The ranges of genetic distances between Pse. lacyformis sp. nov. and species of the genus Pseudechiniscus, for which DNA sequences are available in GenBank, are as follows: COI: 10.6–27.0% (21.1% on average), with the most similar being Pse. aff. xiai (MK804894 –7, Cesari et al. 2020) and the least similar being Pse. indistinctus sp. nov. (MN528471, present study). ITS-2: 10.3–39.3% (26.7% on average), with the most similar being Pse. indistinctus sp. nov. (MN537867, present study) and the least similar being Pse. angelusalas sp. nov. (MN537864, present study).

2. Pseudechiniscus dastychi sp. nov. Roszkowska, Grobys, Bartylak & Kaczmarek (Table 3, Figs 1, 5–6) Pseudechiniscus suillus (Ehrenberg, 1853) (Dastych (1984) Pseudechiniscus sp. 2 (Grobys et al. 2020) Material examined: 80 animals (holotype and 79 paratypes, all females) mounted on microscope slides in Hoyer’s medium, 40 animals prepared for SEM and 7 prepared for barcoding. Description Animals (measurements and statistics in Table 3) Females. Body (Fig. 5) orange in living specimens (transparent after mounting on slides), black eyes present after mounting on slides. Apart from the head appendages (cirri interni and externi and elongated cephalic papillae [secondary clavae]), only lateral cirrus A present (with finger-like clavae near the base [primary clava]) (Fig. 5 A–B,D). Cephalic papillae smaller than clavae. Dorsal plates with small hemispherical granules/upper ends of cuticular pillars (dots in PCM) 0.5–1.2 μm in diameter, densely (spaces between granules 0.5–0.8 μm) and uniformly distributed and joined by thin striae forming indistinct hexagonal pattern (which are in fact thin stripes positioned under epicuticle) (Fig. 5A). Granules/upper ends of cuticular pillars slightly larger in the anterior and posterior parts of the plates. Dorsal plates typical for the genus Pseudechiniscus (cephalic plate (cp), neck plate (np), scapular plate (scp), median plates (m1, m2, m3), paired segmental plates I and II (s1, s2), pseudosegmental plate (psp) and the caudal plate (cap), see also Dorsal and ventral plates and sculpture in Grobys et al. (2020)) well developed. The cp with Wshaped pattern divided into five parts (Fig. 6A, filled arrowhead). The scp divided by transversal fold which forms a long narrow stripe in posterior part of the plate. This narrow stripe is often divided by three longitudinal folds forming four plate parts/subplates (Fig. 5A). The entire scp divided by median longitudinal fold into two parts (Fig. 5 A–B, empty arrow). Additionally, lateral portions of the scp appear detached from the dorsal plate, forming small plate-like structures separated from the scp by a thin bright stripe. Plates m1 and m2 divided in two portions by transverse fold, plate m3 undivided (Fig. 5 A–B, filled indented arrowheads). Laterally to the median plates, lateral intersegmental plates (lip) present. The psp divided by a longitudinal fold. Posterior margin of the psp straight, i.e. without projections, teeth or spines (Fig. 5 A–B, empty indented arrowhead). The cap concave with two Y-shaped bifurcated ridges (Fig. 5 A–B, filled arrowhead). Ventral cuticle with tiny granulation (formed by dense granules/upper ends of cuticular pillars, 0.1–0.4 μm) forming unique pattern (Figs 1, 5 C–D, 6B). Ventral PGs present (granula- tion 0.4–0.5 μm in diameter, spaces between granules 0.4–0.5 μm) with configuration PG:I-II-III-IV-V-VI-VII-VIII a (Figs. 1, 5 C–D). The female gonopore with the typical six-petal rosette. (Fig. 5 C–D, asterisks). The outer cuticle on legs I–III with round patches of granulation (with larger granules but more sparse in the centre and smaller and denser in peripheral parts), on legs IV with uniform, wide stripes of granulation (slightly larger in the centre of these stripes) (Fig. 6 C–D). Triangular spine on leg I and dentate collar on leg IV absent. A finger-like papillae on leg IV present (Fig. 6D, filled arrow). External claws of all legs smooth, internal with spurs directed downwards, identical in legs I–IV (Fig. 6 E–F). Males. Unknown. Juveniles. Unknown. Larvae. Unknown. DNA sequences We obtained good quality sequences for the applied molecular markers: – COI sequence (GenBank: MN528469), 641 bp long; – ITS-2 sequence (GenBank: MN537865), 452 bp long. Etymology. We dedicated this species to the distinguished Polish tardigradologist Dr Hieronymus Dastych, who originally reported specimens of this species from Antarctica as Pse. suillus. Type locality. 65°14’57.30”S, 64°15’13.20”W, 8 m asl: maritime Antarctic, Argentine Islands, Galindez Island, Carolina Point, Charybda Tover, 15 m from sea, moss (Sanionia georgicouncinata) and lichens (Cladonia sp.), 29.01.2016, coll. Ivan Parnikoza. Type depositories. Holotype: slide AT2940/2 and 67 paratypes (slides: AT2940/*, where the asterisk can be substituted by any of the following numbers: 2, 3, 4, 6, 7, 8, 9, 10, 12, 19, 5/S) are deposited at the Department of Animal Taxonomy and Ecology, Institute of Environmental Biology, Adam Mickiewicz University in Poznań, Uniwersytetu Poznańskiego 6, 61-614 Poznań, Poland; 12 paratypes (slides: AT2940/5 and AT2940/18) are deposited at the collection of Binda and Pilato, Museum of the Department of Animal Biology ‘Marcello La Greca’, University of Catania, Italy. Morphological differential diagnosis* *only measurements of adult females are used in differential diagnosis Pseudechiniscus dastychi sp. nov. differs specifically from: 1. Pse. angelusalas sp. nov., by: different ventral pattern (Figs 1, 5 C–D for Pse. dastychi sp. nov. vs Figs 1, 7B for Pse. angelusalas sp. nov.), different ventral PG configuration (PG:I-II-III-IV-V-VI-VII-VIII a in Pse. dastychi sp. nov. vs PG:I-II-III-IV-VI-VIII a in Pse. angelusalas sp. nov.), larger body length (167.0–202.0 μm in Pse. dastychi sp. nov. vs 113.0–143.0 μm in Pse. angelusalas sp. nov.), wider scp (27.5–33.0 μm in Pse. dastychi sp. nov. vs 17.4–20.9 μm in Pse. angelusalas sp. nov.), longer cirri interni (10.4–12.7 μm, [sp=36.9–41.0] in Pse. dastychi sp. nov. vs 6.0–7.3 μm, [sp=34.4–36.6] in Pse. angelusalas sp. nov.), lower sp of cephalic papillae (14.7–16.9 in Pse. dastychi sp. nov. vs 18.3–20.8 in Pse. angelusalas sp. nov.), longer cirri externi (15.9–19.1 μm, [sp=54.7–61.4] in Pse. dastychi sp. nov. vs 9.1–10.4 μm, [sp=49.0–54.3] in Pse. angelusalas sp. nov.), lower sp of clavae (16.4–18.6 in Pse. dastychi sp. nov. vs 22.4–25.7 in Pse. angelusalas sp. nov.), longer cirri A (40.0–45.0 μm in Pse. dastychi sp. nov. vs 24.1–28.0 μm in Pse. angulusalus sp. nov.), longer papillae on leg IV (2.6–3.5 μm in Pse. dastychi sp. nov. vs 2.0–2.4 μm in Pse. angelusalas sp. nov.) and longer claws (see Tables 3 and 4). 2. Pse. beasleyi, by: the scp not divided in the anterior part (the scp divided into four parts in Pse. beasleyi), longer clavae (4.5–5.7 μm in Pse. dastychi sp. nov. vs 3.1–3.9 μm in Pse. beasleyi sp. nov.), longer cephalic papillae (4.2–5.3 μm in Pse. dastychi sp. nov. vs 2.6–3.9 μm in Pse. beasleyi sp. nov.) and longer cirri A (40.0–45.0 μm in Pse. dastychi sp. nov. vs 23.5–31.3 μm in Pse. beasleyi sp. nov.) and different claws length arrangement (shortest claws II and III, and longest claws IV in Pse. dastychi sp. nov. vs shortest claws I and II, and longest claws III and IV in Pse. beasleyi). 3. Pse. chengi, known only from China (Xue et al. 2017), by: dorsal granules joined by striae, plates m1 and m2 divided in two portions by transverse fold (unndivided in Pse. chengi), wider scp (27.5–33.0 μm in Pse. dastychi sp. nov. vs 20.8–25.7 μm in Pse. chengi), longer cephalic papillae (4.2–5.3 μm in Pse. dastychi sp. nov. vs 2.8–3.7 μm in Pse. chegi), longer cirri interni and externi (10.4–12.7 μm and 15.9–19.1 μm respectively in Pse. dastychi sp. nov. vs 3.2–8.5 μm and 10.5–14.2 μm respectively in Pse. chegi), longer cirri A (40.0–45.0 μm [sp=135.2–149.1] in Pse. dastychi sp. nov. vs 21.4–27.5 μm [sp=96.0 –111.1] in Pse. chengi) and higher cirrus A /body length ratio (22–26% in Pse. dastychi sp. nov. vs 13–17% in Pse. chengi). 4. Pse. clavatus, by: different shape of clavae (finger-like in Pse. dastychi sp. nov. vs club-shaped in Pse. clavatus) and normally developed cephalic papillae (reduced in Pse. clavatus). 5. Pse. ehrenbergi sp. nov. see Morphological differential diagnosis of Pse. ehrenbergi sp. nov. above. 6. Pse. facettalis, known from distant localities throughout the world (McInnes 1994). Based on present study, an inaccurate description of this species makes it impossible to differentiate this taxon from Pse. dastychi sp. nov.. See also Morphological differential diagnosis of Pse. suillus and Discussion in the paper by Grobys et al. (2020). 7. Pse. indistinctus sp. nov., by: different ventral pattern (Figs 1, 5 C–D for Pse. dastychi sp. nov. vs Figs 1, 13 C–D for Pse. indistinctus sp. nov.), different ventral PG configuration (PG:I-II-III-IV-V-VI-VII-VIII a in Pse. dastychi sp. nov. vs PG:I-II-III-IV-V-VI-VIII a in Pse. indistinctus sp. nov.), granules on the cap similar in size to other dorsal plates (granules visibly larger on the cap in comparison with other dorsal plates in Pse. indistinctus sp. nov.), larger body size (167.0–202.0 μm in Pse. dastychi sp. nov. vs 140.0–166.0 μm in Pse. indistinctus sp. nov.), wider scp (27.5–33.0 μm in Pse. dastychi sp. nov. vs 20.5–24.9 μm in Pse. indistinctus sp. nov.), longer cirri interni (10.4–12.7 μm in Pse. dastychi sp. nov. vs 7.0–9.2 μm in Pse. indistinctus sp. nov.), longer cirri externi (15.9–19.1 μm in Pse. dastychi sp. nov. vs 11.3–14.4 μm in Pse. indistinctus sp. nov.), lower sp of clavae (16.4–18.6 in Pse. dastychi sp. nov. vs 19.3–22.5 in Pse. indistinctus sp. nov.), longer cirri A (40.0–45.0 μm in Pse. dastychi sp. nov. vs 27.6–34.2 μm in Pse. indistinctus sp. nov.) and longer claws (see Tables 3 and 7). 8. Pse. juanitae, known from Austria, Brazil (type locality), Italy and Galapagos Islands (McInnes 1994). Based on present study, an inaccurate description of this species makes it impossible to differentiate this taxon from Pse. dastychi sp. nov.. See also Morphological differential diagnosis of Pse. suillus and Discussion in the paper by Grobys et al. (2020). 9. Pse. lacyformis sp. nov., by: different ventral pattern (Figs 1, 5 C–D for Pse. dastychi sp. nov. vs Figs 1, 10B for Pse. lacyformis sp. nov.), different ventral PG configuration (PG:I-II-III-IV-V-VI-VII-VIII a in Pse. dastychi sp. nov. vs PG:I-II-III-IV-VI-VIII a in Pse. lacyformis sp. nov.), dorsal granules joined by striae, wider scp (27.5–33.0 μm in Pse. dastychi sp. nov. vs 20.4–26.0 μm in Pse. lacyformis sp. nov.), lower sp of cirri interni (36.9–41.0 in Pse. dastychi sp. nov. vs 48.4–53.9 in Pse. lacyformis sp. nov.), lower sp of cirri externi (54.7–61.4 in Pse. dastychi sp. nov. vs 66.5–77.8 in Pse. lacyformis sp. nov.), lower sp of clavae (16.4–18.6 in Pse. dastychi sp. nov. vs 23.4–26.8 in Pse. lacyformis sp. nov.), longer cirri A (40.0–45.0 μm in Pse. dastychi sp. nov. vs 26.5–35.0 μm in Pse. lacyformis sp. nov.), higher cirrus A /body length ratio (22–26% in Pse. dastychi sp. nov. vs 15–20% in Pse. lacyformis sp. nov.), lower sp of papillae on leg IV (8.9–11.6 in Pse. dastychi sp. nov. vs 14.7–17.2 in Pse. lacyformis sp. nov.), longer spurs on all legs (2.1–2.8 μm in Pse. dastychi sp. nov. vs 1.3–1.7 μm in Pse. lacyformis sp. nov.) and lower spur/branch length ratio of all claws (23–30% in Pse. dastychi sp. nov. vs 15–20% in Pse. lacyformis sp. nov.). 10. Pse. megacephalus, by: different shape of cephalic papillae (elongated in Pse. dastychi sp. nov. vs mushroom-like in Pse. megacephalus), absence of papilliform projection between external buccal cirri and cirri A. 11. Pse. suillus, by: a small papilla-like structure on leg I absent, different ventral pattern (Figs 1, 5 C–D for Pse. dastychi sp. nov. vs Figs 1 herein and 4C–D in Grobys et al. (2020) for Pse. suillus), well-developed ventral patches of granulation, different ventral PG configuration (PG:I-II-III-IV-V-VI-VII-VIIIa in Pse. dastychi sp. nov. vs PG: I-II-III-IV-VI-VIIIg in Pse. suillus), dorsal granules joined by striae, wider scp (27.5–33.0 μm in Pse. dastychi sp. nov. vs 18.8–23.5 μm in Pse. suillus), lower sp of cirri interni (36.9–41.0 in Pse. dastychi sp. nov. vs 44.0– 49.6 in Pse. suillus), lower sp of cephalic papillae (14.7–16.9 in Pse. dastychi sp. nov. vs 19.1–24.3 in Pse. suillus), lower sp of cirri externi (54.7–61.4 in Pse. dastychi sp. nov. vs 62.1–75.0 in Pse. suillus), lower sp of clavae (16.4–18.6 in Pse. dastychi sp. nov. vs 20.9–26.8 in Pse. suillus), higher cirrus A /body length ratio (22–26% in Pse. dastychi sp. nov. vs 17–21% in Pse. suillus), lower sp of papillae on leg IV (8.9–11.6 in Pse. dastychi sp. nov. vs 14.7–18.4 in Pse. suillus) and longer claws (see Table 3 herein and Table 4 in Grobys et al. 2020). 12. Pse. xiai, known only from China (Wang et al. 2018), by: different ventral pattern (Figs 1, 5 C–D for Pse. dastychi sp. nov. vs Figs 1B, F and 2E in Wang et al. (2018) for Pse. xiai), dorsal granules joined by striae, wider scp (27.5–33.0 μm in Pse. dastychi sp. nov. vs 11.3–26.7 μm in Pse. xiai), longer cephalic papillae (4.2–5.3 μm in Pse. dastychi sp. nov. vs 1.4–3.9 μm in Pse. xiai), longer cirri A (40.0–45.0 μm in Pse. dastychi sp. nov. vs 24.7–31.9 μm in Pse. xiai), and higher cirrus A /body length ratio (22–26% in Pse. dastychi sp. nov. vs 13–16% in Pse. xiai). Genotypic differential diagnosis The ranges of genetic distances between Pse. dastychi sp. nov. and species of the genus Pseudechiniscus, for which DNA sequences are available in GenBank, are as follows: COI: 2.0–28.5% (24.0% on average), with the most similar being Pseudechiniscus sp. (KJ857005, Velasco- Castrillón et. al. 2015 (described in GenBank as Echiniscus sp.); for more details see Discussion in Grobys et al. 2020) and the least similar being Pse. aff. suillus (MK804907, Cesari et al., 2020). ITS-2: 7.7–36.8% (26.5% on average), with the most similar being Pse. ehrenbergi sp. nov. (MN537866, present study) and the least similar being Pse. lacyformis sp. nov. (MN537868, present study)., Published as part of Roszkowska, Milena, Grobys, Daria, Bartylak, Tomasz, Gawlak, Magdalena, Kmita, Han- Na, Kepel, Andrzej, Kepel, Marta, Parnikoza, Ivan & Kaczmarek, Łukasz, 2020, Integrative description of five Pseudechiniscus species (Heterotardigrada Echiniscidae: the suillus-facettalis complex), pp. 451-484 in Zootaxa 4763 (4) on pages 461-466, DOI: 10.11646/zootaxa.4763.4.1, http://zenodo.org/record/3762014, {"references":["Ehrenberg, C. G. (1853) Diagnoses novarum formarum. Monatsberichte der Koniglich preussischen Akademie der Wissenschaften zu Berlin, 8, 526 - 533.","Dastych, H. (1984) The Tardigrada from Antartica with description of several new species. Acta Zoologica Cracoviensia, 27, 377 - 436.","Grobys, D., Roszkowska, M., Gawlak, M., Kmita, H., Kepel, A., Kepel, M., Parnikoza, I., Bartylak, T. & Kaczmarek, L. (2020) High diversity in the Pseudechiniscus suillus-facettalis complex (Heterotardigrada; Echiniscidae) with remarks on the morphology of the genus Pseudechiniscus. Zoological Journal of the Linnean Society, 188 (3), 733 - 752. https: // doi. org / 10.1093 / zoolinnean / zlz 171","Xue, J., Li, X., Wang, L., Xian, P. & Chen, H. (2017) Bryochoerus liupanensis sp. nov. and Pseudechiniscus chengi. sp. nov. (Tardigrada: Heterotardigrada: Echiniscidae) from China. Zootaxa, 4291 (2), 324 - 334. https: // doi. org / 10.11646 / zootaxa. 4291.2.5","McInnes, S. J. (1994) Zoogeographic distribution of terrestrial / freshwater tardigrades from current literature. Journal of Natural History, 28, 257 - 352. https: // doi. org / 10.1080 / 00222939400770131","Wang, L., Xue, J. & Li, X. (2018) A description of Pseudechiniscus xiai sp. nov., with a key to genus Pseudechiniscus in China. Zootaxa, 4388 (2), 255 - 264. https: // doi. org / 10.11646 / zootaxa. 4388.2.7","Cesari, M., Montanari, M., Kristensen, R. M., Guidetti, R., Bertolani, R. & Rebecchi, L. (2020) An integrated study of the biodiversity within Pseudechiniscus suillus - facettalis group (Heterotardigrada, Echiniscidae). Zoological Journal of the Linnean Society, 188 (3), 717 - 732. https: // doi. org / 10.1093 / zoolinnean / zlz 045"]}

Family: Echiniscidae Thulin, 1928 Subfamily: Echiniscinae Guil, Jørgensen & Kristensen, 2019, Published as part of Bartylak, Tomasz, Kulpa, Adam, Grobys, Daria, Kepel, Marta, Kepel, Andrzej, Kmita, Hanna, Gawlak, Magdalena, Grabiński, Wojciech, Roszkowska, Milena & Kaczmarek, Łukasz, 2019, Variability of Echiniscus tristis Gąsiorek & Kristensen, 2018 - is morphology sufficient for taxonomic differentiation of Echiniscidae?, pp. 1-24 in Zootaxa 4701 (1) on page 5, DOI: 10.11646/zootaxa.4701.1.1, http://zenodo.org/record/3557773, {"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","Guil, N., Jorgensen, A. & Kristensen R. M. (2019) An upgraded comprehensive multilocus phylogeny of the Tardigrada tree of life. Zoologica Scripta, 48, 120 - 137. https: // doi. org / 10.1111 / zsc. 12321"]}

Bartylak, Tomasz, Kulpa, Adam, Grobys, Daria, Kepel, Marta, Kepel, Andrzej, Kmita, Hanna, Gawlak, Magdalena, Grabiński, Wojciech, Roszkowska, Milena, Kaczmarek, Łukasz (2019): Variability of Echiniscus tristis Gąsiorek & Kristensen, 2018 - is morphology sufficient for taxonomic differentiation of Echiniscidae? Zootaxa 4701 (1): 1-24, DOI: https://doi.org/10.11646/zootaxa.4701.1.1

Echiniscus tristis Gąsiorek & Kristensen, 2018 (Tables 4–10, Figs 2–9) Material examined: 320 animals mounted on microscope slides in Hoyer’s medium, 20 animals prepared for SEM and 6 prepared for molecular analysis from Madagascar, 22°37’07.7”S, 46°43’14.5”E, ca. 1,187 m asl, Fianarantsoa Province, Ivohibory forest, lichen on quartz rock, coll. Marta Kepel and Andrzej Kepel. Description of the Madagascan population. Animals (measurements and statistics in Tables 4-8) Females. Body light orange in live specimens and transparent/white after preparation (Fig. 2). Eyes absent or not visible after the preparation. Apart from the head appendages (internal and external cirri and cylindrical cephalic papillae [secondary clavae], the appendage A with clava [primary clava] near its base) present (Fig. 2). Dorsal and lateral appendages in the shape of long and short spines present in positions A-C-Dd- E (mainly in adult females; morphotype 1) or A-Dd- E (mainly in juveniles and larvae, morphotype 2). This means that the general chaetotaxy of adult females of this species is A-(C)- Dd- E . However, single females in the studied population possessed also different chaetotaxy configurations, i.e. A-E, A-C-E, A-C-Cd- Dd- E, A-C-D-Dd- E , but with some spines being present only on one side of the body (Fig. 2–3). See also Morphological variability below. Dorsal plates covered by smaller and larger pores (Figs 4–6). Pores 0.6–2.2 µm in diameter visible as bright/ white dots when focusing down through cuticle (Figs 4 A–B, E–F; 5A, C–F; 6A). Although pores may differ significantly in the same individual and between specimens, in general they are largest on the head plate, narrow anterior stripe of paired plates and in the place where median plate 3 should be localised (Figs 5A, 6B arrow). Dorsal plates well developed. Head and scapular plates not faceted. In PCM, lateral portions of the scapular plate appear to be detached from the dorsal plate, forming small additional plates (one on each side of the body) divided from the scapular plate by a thin bright stripe. This false division is caused by a bend of the plate where the cuticle is thinner; in SEM, this pseudo-division is not visible. Paired plates I and II are divided into two parts—narrow anterior stripe and a wider posterior part—by smooth stripe without pores (Fig. 5 B–D). Median plates 1 and 2 undivided, median plate 3 absent (but cuticle in this place sculptured with pores) (Fig. 6B). The terminal plate possesses two notches and in some specimens a characteristic pattern (Figs 6 A–B). Narrow stripes form four rectangular areas in central part of terminal plate (Figs 6 A–B). Ventral plates absent, but cuticle possesses tiny and regular granulation (caused by dense top endings of cuticular pillars). Granulation is little larger between the legs (0.9–1.0 µm diameter) than in other parts of the ventral side, 0.4–0.7 µm diameter) (Fig. 6C). A-C-Dd- E A-Dd- E A-C-Cd- D-Dd- E A-C-D-Dd- E A-C-Cd- Dd- E A-C-Dd- E A-C-Dd- E A-C-E A-E On outer cuticle of legs I–III two clearly visible patches, composed of tiny and regular granulation similar to this on ventral side, are present: a narrow stripe (cuticular fold) in upper part (Fig. 7 A–C, filled arrowhead) and a large wide stripe below (plate) (Fig. 7 A–C, filled arrow). Triangular spine on leg I and finger-like papilla on leg IV present (Fig. 7C, E–F, H, indented filled arrowhead and empty arrow). Dentate collar on legs IV with sculpture similar to those on dorsal plates but with smaller pores, and with eight to twelve sharp, triangular teeth (Fig. 7 D–F, empty arrowhead). Above the dentate collar a cuticular fold also present (Fig. 7 D–F, filled arrowhead). External claws of all legs smooth, internal with spurs directed downwards (Fig. 7 F–H). Juveniles (Fig. 8). In general they are similar to adult females. The chaetotaxy types in juveniles are A-C-Dd- E, A-Dd- E, A-C-D-Dd- E, A-C-E , with the most frequent A-Dd- E . For more details see Table 9 and Morphological variability below. Terminal plate without characteristic pattern or with pattern only poorly visible. Gonopore absent. Larvae (Fig. 9). In general they are similar to adult females, but dorsal cuticular pores seem to be larger in comparison to the body size. The chaetotaxy in larvae is always A-Dd- E . For more details see Table 9 and Morphological variability below. Terminal plate without the characteristic pattern. All legs with two internal claws. Gonopore and anus absent. Males. Not found, indicating the species is most likely parthenogenetic. Eggs. Smooth, light orange and deposited in the exuviae up to 5 in one exuvium. DNA sequences. We obtained good quality sequences for the applied molecular markers (voucher numbers marked in bold indicate specimens which were analysed using all three markers)

Moreno, Antonio, Roszkowska, Milena, García, Mario Alberto, Flores, José Juan, Kaczmarek, Łukasz (2019): Current knowledge on Mexican tardigrades with a description of Milnesium cassandrae sp. nov. (Eutardigrada: Milnesiidae) and discussion on the taxonomic value of dorsal pseudoplates in the genus Milnesium Doyère, 1840. Zootaxa 4691 (5): 501-524, DOI: https://doi.org/10.11646/zootaxa.4691.5.5

Echiniscus cf. tamus Material examined: One specimen: Las Crucitas (24°30′46.62′′N, 99°59′03.91′′W, ca. 2499 m asl), Aramberri, Nuevo Leon. Mexico. Lichen on tree, coll. Miguel Angel Gonzalez Botello, 3 May 2014. Comments: Echiniscus tamus Mehlen, 1969 was described from North America (Texas, USA). Our single specimen is similar to the Texan specimen (Mehlen 1969; Miller & Mehlen 2007), but differs by having a basal spur on the internal claws. This is possibly a new species, but for final identification a larger number of specimens are required., Published as part of Moreno, Antonio, Roszkowska, Milena, García, Mario Alberto, Flores, José Juan & Kaczmarek, Łukasz, 2019, Current knowledge on Mexican tardigrades with a description of Milnesium cassandrae sp. nov. (Eutardigrada: Milnesiidae) and discussion on the taxonomic value of dorsal pseudoplates in the genus Milnesium Doyère, 1840, pp. 501-524 in Zootaxa 4691 (5) on page 505, DOI: 10.11646/zootaxa.4691.5.5, http://zenodo.org/record/3527562, {"references":["Mehlen, R. H. (1969) New Tardigrada From Texas. The American Midland Naturalist, 81, 395 - 404. https: // doi. org / 10.2307 / 2423979","Miller, W. R. & Mehlen, R. H. (2007) Tardigrades of North America: re - description of Echiniscus tamus Mehlen, 1969 (Tardigrada: Heterotardigrada: Echiniscoidae: Echiniscidae). Proceedings of the Biological Society of Washington, 120, 184 - 188. https: // doi. org / 10.2988 / 0006 - 324 X (2007) 120 [184: TONARO] 2.0. CO; 2"]}

Echiniscus manuelae da Cunha & du Nascimento Ribeiro, 1962 Terra typica: Madera Island (Portugal, Atlantic Ocean). Material examined: One specimen: El Vado (25°21′54.81′′N, 100°14′02.27′′W, ca. 1411 m asl), Cienega de Gonzalez, Santiago, Nuevo Leon, Mexico. Moss on tree (Quercus sp.), coll. Mario Alberto Garcia Aranda, 17 January 2015. Comments: Our specimens correspond well with the original description (da Cunha & du Nascimento Ribeiro 1962). This species has been reported from the type locality, Madera Island (da Cunha & du Nascimento 1962; McInnes 1994), Corrientes Province, Bella Vista in Argentina (Claps & Rossi 1988) and Costa Rica (Kaczmarek & Michalczyk 2010). Kaczmarek & Michalczyk (2010) and Kaczmarek et al. (2014) suggest the possibility that the Costa Rican record, found in a big city, was a non - indigenous to the region, but had been brought to Costa Rica by humans. Our record for this study corresponds to a small town, remote from big cities. This is a new record for Mexico., Published as part of Moreno, Antonio, Roszkowska, Milena, García, Mario Alberto, Flores, José Juan & Kaczmarek, Łukasz, 2019, Current knowledge on Mexican tardigrades with a description of Milnesium cassandrae sp. nov. (Eutardigrada: Milnesiidae) and discussion on the taxonomic value of dorsal pseudoplates in the genus Milnesium Doyère, 1840, pp. 501-524 in Zootaxa 4691 (5) on page 504, DOI: 10.11646/zootaxa.4691.5.5, http://zenodo.org/record/3527562, {"references":["Da Cunha, A. X. & du Nascimento R. F. (1962) A fauna de tardigrados da Ilha da Madeira. Memorias e estudos do Museu Zoologico da Universidade de Coimbra, 279, 1 - 24.","McInnes, S. J. (1994) Zoogeographic distribution of terrestrial / freshwater tardigrades from current literature. Journal of Natural History, 28, 257 - 352. https: // doi. org / 10.1080 / 00222939400770131","Claps, M. C. & Rossi, G. C. (1988). Tardigrados de Argentina VI. Iheringia, 67, 3 - 11.","Michalczyk, L. & Kaczmarek, L. (2010) Description of Doryphoribius dawkinsi, a new species of Tardigrada (Eutardigrada: Hypsibiidae) from the Costa Rican highlands, with the key to the genus Doryphoribius. Zootaxa, 2393 (1), 46 - 58. https: // doi. org / 10.11646 / zootaxa. 2393.1.4","Kaczmarek, L., Michalczyk, L. & McInnes, S. J. (2014) Annotated zoogeography of non - marine Tardigrada. Part I: Central America. Zootaxa, 3763 (1), 1 - 62. https: // doi. org / 10.11646 / zootaxa. 3763.1.1"]}

Cornechiniscus lobatus (Ramazzotti, 1943) Terra typica: Italy (Europe). Material examined: 24 specimens: 1 km S of Puerto Genovevo (25°43′02.98′′N, 100°04′18.71′′W, ca. 1103 m asl), Santiago, Nuevo Leon, Mexico. Lichen on rock, coll. Antonio Moreno - Talamantes, 30 August 2014. Comments: Our specimens correspond well with to the original description. This species has been reported in many localities in the northern hemisphere (McInnes 1994). This species was reported from Sinaloa, Mexico (Beasley 1972), and this is the second record for Mexico., Published as part of Moreno, Antonio, Roszkowska, Milena, García, Mario Alberto, Flores, José Juan & Kaczmarek, Łukasz, 2019, Current knowledge on Mexican tardigrades with a description of Milnesium cassandrae sp. nov. (Eutardigrada: Milnesiidae) and discussion on the taxonomic value of dorsal pseudoplates in the genus Milnesium Doyère, 1840, pp. 501-524 in Zootaxa 4691 (5) on page 505, DOI: 10.11646/zootaxa.4691.5.5, http://zenodo.org/record/3527562, {"references":["Ramazzotti, G. (1943) Nuova varieta del Tardigrado Pseudechiniscus cornutus. Rivista di Scienze Naturali ' Natura', 34, 89 - 90.","McInnes, S. J. (1994) Zoogeographic distribution of terrestrial / freshwater tardigrades from current literature. Journal of Natural History, 28, 257 - 352. https: // doi. org / 10.1080 / 00222939400770131","Beasley, C. W. (1972) Some tardigrades from Mexico. The Southwestern Naturalist, 17, 21 - 29. https: // doi. org / 10.2307 / 3669835"]}

Pseudechiniscus cf. juanitae Material examined: One specimen: Piedmont of Sierra Las Mitras (25°44′55.57′′N, 100°24′11.19′′W, ca. 632 m asl), Monterrey, Nuevo Leon, Mexico. Lichen (Physcia stellaris) on trees (Parkinsonia aculeta and Celtis pallida), coll. Antonio Moreno - Talamantes, 7 August 2014. Comments: Pseudechiniscus juanitae de Barros, 1939 was described from Brazil and later reported from Costa Rica and Mexico (Kaczmarek et al. 2014, 2015), but the diagnosis for this species is very unclear, and this is why we report our specimen as confer to the nominal taxon., Published as part of Moreno, Antonio, Roszkowska, Milena, García, Mario Alberto, Flores, José Juan & Kaczmarek, Łukasz, 2019, Current knowledge on Mexican tardigrades with a description of Milnesium cassandrae sp. nov. (Eutardigrada: Milnesiidae) and discussion on the taxonomic value of dorsal pseudoplates in the genus Milnesium Doyère, 1840, pp. 501-524 in Zootaxa 4691 (5) on page 505, DOI: 10.11646/zootaxa.4691.5.5, http://zenodo.org/record/3527562, {"references":["De Barros, R. (1939) Pseudechiniscus juanitae nova especie de Tardigrado. Boletim Biologico, New Series, 4, 367 - 368.","Kaczmarek, L., Michalczyk, L. & McInnes, S. J. (2014) Annotated zoogeography of non - marine Tardigrada. Part I: Central America. Zootaxa, 3763 (1), 1 - 62. https: // doi. org / 10.11646 / zootaxa. 3763.1.1"]}

Family: Echiniscidae Thulin, 1928 Subfamily: Echiniscinae Guil, Jørgensen & Kristensen, 2018, Published as part of Moreno, Antonio, Roszkowska, Milena, García, Mario Alberto, Flores, José Juan & Kaczmarek, Łukasz, 2019, Current knowledge on Mexican tardigrades with a description of Milnesium cassandrae sp. nov. (Eutardigrada: Milnesiidae) and discussion on the taxonomic value of dorsal pseudoplates in the genus Milnesium Doyère, 1840, pp. 501-524 in Zootaxa 4691 (5) on page 504, DOI: 10.11646/zootaxa.4691.5.5, http://zenodo.org/record/3527562, {"references":["Thulin, G. (1928) Uber die Phylogie und das System der Tardigraden. Heriditas, 11, 207 - 266. https: // doi. org / 10.1111 / j. 1601 - 5223.1928. tb 02488. x","Guil, N., Jorgensen, A. & Kristensen, R. (2018) An upgraded comprehensive multilocus phylogeny of the Tardigrada tree of life. Zoologica Scripta, 48 (1), 1 - 18. https: // doi. org / 10.1111 / zsc. 12321"]}

15. Echiniscus carusoi Pilato, 1972 Terra typica: Italy (Europe) Libya: • 32°29′N, 20°50′E; 350 m asl: Marj District, Barce [Marj], moss. Pilato & Pennisi (1976) Record numbers. Libya: 1, total: 1. Remarks. Species with a limited distribution of Southern Italy and Libya (McInnes 1994).

12. Echiniscus bisetosus Heinis, 1908a [T] Echiniscus (E) bisetosus Heinis, 1908 (Binda 1971) Terra typica: Switzerland (Europe) Algeria: • 34°40′N, 06°11′E; 0 m asl: Biskra Province, 30 km from Biskra, El Oued [Zeribet el Oued]. Binda & Pilato (1987) Morocco: • 31°12′N, 07°52′W; 2,500 m asl [2,600 m asl]: Marrakesz-Tansift-Al-Hauz Region, Oukaïmeden, moss. Binda (1971) Record numbers. Algeria: 1, Morocco: 1; total: 2. Remarks. Species with typical Holarctic distribution recorded from many localities in Europe and fewer from Canada, Afghanistan, Algeria and Morocco (McInnes 1994).

9. Echiniscus arctomys Ehrenberg, 1853 sensu lato [T] E. aRctomys EHR (Murray 1907b) Terra typica: Switzerland (Europe) Uganda: • 01°22′N, 32°17′E: Undefined locality, moss. Murray (1907b) Record numbers. Uganda: 1; total: 1. Remarks. This reference to cysts in moss samples from Uganda (Murray 1907b), is questionable.

6. Cornechiniscus lobatus (Ramazzotti, 1943) [T] CoRnechiniscus lobatus (Ramløv & Kristensen 1985) Terra typica: Italy (Europe) Jordan: • 31°50′N, 36°49′E; 500 m asl: Zarqa Governorate, Zarqa [Azarq Wetland Reserve], moss. Kaczmarek & Michalczyk (2004a) Morocco: • 30°27′N, 07°32′W; 1,850 m asl: Souss-Massa-Drâa Region, Tizi-n-Taghatine, sparse vegetation, mosses and lichens on isolated basalt cliffs. Ramløv & Kristensen (1985) Record numbers. Jordan: 1, Morocco: 1; total: 2. Remarks. A northern hemisphere species (McInnes 1994a), also known from Nearctic Mexico and Neotropic Peru (Kaczmarek et al. 2014, 2015)., Published as part of Kaczmarek, Łukasz, 2017, Annotated zoogeography of non-marine Tardigrada. Part IV: Africa, pp. 1-74 in Zootaxa 4284 (1) on page 9, DOI: 10.11646/zootaxa.4284.1.1, http://zenodo.org/record/1010400, {"references":["Ramazzotti, G. (1943) Nuova varieta del Tardigrado Pseudechinsicus coRnutus. Rivista di Scienze NatuRali \" NatuRa \", 34, 89 - 90.","Ramlov, H. & Kristensen, R. M. (1985) Bjornedyr, Tardigrada. Marokko - 1985. Dansk NatuRhistoRisk FoRenings Rejse til MaRokko, 8 - 22 ApRil 1985, pp. 27 - 32.","Kaczmarek, L. & Michalczyk, L. (2004 a) The first record of water bears (Tardigrada) from Jordan. DiRasat (PuRe Sciences), 31 (2), 154 - 157.","McInnes, S. J. (1994) Zoogeographic distribution of terrestrial / freshwater tardigrades from current literature. JouRnal of NatuRal HistoRy, 28, 257 - 352. https: // doi. org / 10.1080 / 00222939400770131"]}

22. Echiniscus limai da Cunha & do Nascimento Ribeiro, 1964 [T] Echiniscus limai sp. n. (da Cunha & do Nascimento Ribeiro 1964) Terra typica: Angola (Africa) Angola: • 07°25′S, 15°06′E; 850 m asl: Type Locality: Uíge Province, Mahondo, lichens. da Cunha & do Nascimento (1964) Record numbers. Angola: 1; total: 1. Remarks. The species has a disjunct Africa (type locale) and Asia distribution (Beasley & Cleveland 1996; Kaczmarek & Beasley 2002; Li et al. 2008), suggesting that the Asian records need re-assessment.

4. Bryodelphax tatrensis (Węglarska, 1959) Terra typica: Poland (Europe) Libya: • 32°29′N, 20°50′E; 350 m asl: Marj District, Barce [Marj], moss. Pilato & Pennisi (1976) Record numbers. Libya: 1, total: 1. Remarks. A largely Palaearctic distribution and relatively rare species., Published as part of Kaczmarek, Łukasz, 2017, Annotated zoogeography of non-marine Tardigrada. Part IV: Africa, pp. 1-74 in Zootaxa 4284 (1) on pages 8-9, DOI: 10.11646/zootaxa.4284.1.1, http://zenodo.org/record/1010400, {"references":["Weglarska, B. (1959) Tardigraden Polens II. Vestnik Ceskoslovenske Zoologicke Spolecnosti, 23, 354 - 357.","Pilato, G. & Pennisi, G. (1976) Prime notizie sui Tardigradi della Cirenaica. Animalia, 3, 243 - 258."]}

19. Echiniscus duboisi Richters, 1902 [T] Echiniscus duboisii Richters (Murray 1913) Echiniscus duboisi F. Richters, 1902 (Teunissen 1938) Terra typica: Java (Indonesia) Republic of South Africa: • 29°44′S, 22°44′E: Undefined locality, Cape Colony [Cape Province], moss. Murray (1913) • 29°44′S, 30°32′E; 750 m asl: KwaZulu-Natal Province, Tala Private Game Reserve, lichen on tree (Acacia). Meyer & Hinton (2009) Rwanda: • 01°24′S, 29°39′E; 3,000 m asl: Northern Province, Albert National Park [Volcanoes National Park / Mgahinga Gorilla National Park], W slope of Gahinga Volcano, bamboo forest, soil. Teunissen (1938) Record numbers. Republic of South Africa: 2; Rwanda: 1; total: 3. Remarks. Species with a limited tropical and subtropical distribution from Africa, Australia, Indonesia and South America (McInnes 1994).

3. Bryodelphax parvulus Thulin, 1928 sensu lato [T] BRyodelphax paRvulus Thulin 1928 (da Cunha & do Nascimento 1964) Echiniscus (B.) paRvulus (Thulin 1928) (Binda 1971) BRyodelphax paRvulus (Ramløv & Kristensen 1985) Terra typica: Norway (Europe) Angola: • 07°47′S, 15°01′E; 650 m asl: Uíge Province, near Carmona [Uíge], Estate at Pumba Loge, lichens. da Cunha & do Nascimento (1964) Libya: • 32°29′N, 20°50′E; 350 m asl: Marj District, Barce [Marj], moss. Pilato & Pennisi (1976) Morocco: • 32°56′N, 05°40′W; 1,700 m asl [850 m asl]: Meknès-Tafilalet Region, Khénifra, moss. Binda (1971) • 31°08′N, 07°55′W; 1,700 m asl: Tadla-Azilal Region, Imlil, walnut forest, moss on limestone cliffs. Ramløv & Kristensen (1985) • 31°06′N, 07°55′W; 2,300–2,500 m asl: Tadla-Azilal Region, path to the Jebel Toubkal, scattered tussock vegetation, moss on basalt and mica slate. Ramløv & Kristensen (1985) Record numbers. Angola: 1, Morocco: 3, Libya: 1; total: 5. Remarks. The considerable morphological variation of B. parvulus (e.g. the presence or absence of the spine and papilla on legs, etc.) suggests that this taxon may be a complex of closely related species, each with limited geographic ranges, rather than a single cosmopolitan species (see also Pilato et al. 2010). The actual geographic range and the presence of B. parvulus sensu lato in Africa must be carefully verified.

5. Cornechiniscus cornutus (Richters, 1907a) [T] CoRnechiniscus coRnutus Richters, 1906 (Pilato & Pennisi 1976) CoRnechiniscus coRnutus (Richters, 1906) (Binda & Pilato 1987) Terra typica: Germany (Europe) Algeria: • 31°56′N, 05°24′E; 150 m asl: Ouargla Province, Ain Beida. Binda & Pilato (1987) Libya: • 32°29′N, 20°50′E; 350 m asl: Marj District, Barce [Marj], moss. Pilato & Pennisi (1976) Record numbers. Algeria: 1, Libya: 1; total: 2. Remarks. A mainly northern hemisphere (Palaearctic) species with a Nearctic record (McInnes 1994, Kaczmarek et al. 2016)., Published as part of Kaczmarek, Łukasz, 2017, Annotated zoogeography of non-marine Tardigrada. Part IV: Africa, pp. 1-74 in Zootaxa 4284 (1) on page 9, DOI: 10.11646/zootaxa.4284.1.1, http://zenodo.org/record/1010400, {"references":["Richters, F. (1907 a) Zwei neue Echiniscus Arten. ZoologischeR AnzeigeR, 31, 197 - 202.","Pilato, G. & Pennisi, G. (1976) Prime notizie sui Tardigradi della Cirenaica. Animalia, 3, 243 - 258.","McInnes, S. J. (1994) Zoogeographic distribution of terrestrial / freshwater tardigrades from current literature. JouRnal of NatuRal HistoRy, 28, 257 - 352. https: // doi. org / 10.1080 / 00222939400770131"]}

33. Echiniscus rugospinosus Marcus, 1928 [T] Echiniscus sp. (Murray 1913) Echiniscus Rugospinosus nom nov. (Marcus 1928) Terra typica: East Africa Uganda / Kenya: • 00°47′N, 34°22′E: Type Locality: Undefined locality, tropical East Africa. (Murray 1913; as an unnamed species), Marcus (1928) Record numbers. Uganda / Kenya: 1; total: 1. Remarks. This species has not been reported since the original reference (Murray 1 913), and although currently endemic to Africa, it will be difficult to locate and verify the type locality.

41. Pseudechiniscus facettalis Petersen, 1951 sensu lato [T] Pseudechiniscus suillus (Ehrenberg, 1853) forma facettalis Petersen, 1951 (Haspeslagh 1982) Terra typica: Greenland Kenya: • 00°10′S, 37°18′E; 4,270 m asl: Nyeri County, Mount Kenya National Park, Mount Kenya, Teleki Tarn, small effluent and shallow pond, silt, sediments and sandy gravel. Haspeslagh (1982) Tunisia: • 36°43'19''N, 08°43'30''E; 474 m asl: Jendouba Governorate, Beni M'tir, forest, moss on soil. Gąsiorek et al. (2017) Record numbers. Kenya: 1, Tunisia: 1; total: 2. Remarks. A widespread taxon (McInnes 1994a), in the suillus species complex. Only two individuals were identified in Kenyan samples and from a non-typical (aquatic) habitat, which coupled with the type locality (Greenland) indicates this African record requires re- examination.

42. Pseudechiniscus jiroveci Bartoš, 1963 [T] Pseudechiniscus jiRoveci Bartos, 1963 (Binda 1984) Pseudechiniscus jiRoveci Bartoš 1963 (Pilato et al. 1991, Binda & Pilato 1995a) Terra typica: China (Asia) Republic of South Africa: • 34°01′S, 23°55′E; 200 m asl: Eastern Cape Province, Tizitzikama / Tzitzikama [Tsitsikamma National Park], moss. Binda (1984), Pilato et al. (1991) Tanzania: • 03°09′S, 35°29′E: Undefined locality, Arusha Region, Ngorongoro Volcano, moss. Pilato et al. (1991) • 03°12′S, 37°31′E; 2,200–2,400 m asl: Kilimanjaro Region, near Marangu [gate], outskirts of the entrance of Kilimanjaro National Park, mosses. Binda & Pilato (1995a) Record numbers. Republic of South Africa: 1, Tanzania: 2; total: 3. Remarks. Originally described from five juvenile (two clawed) individuals (Bartoš, 1963), this rare species has a restricted distribution including the type locale (China), Mongolia (Kaczmarek & Michalczyk 2006) and Africa.

25. Echiniscus merokensis Richters, 1904b sensu lato [T] Echiniscus meRokensis Richt (Marcus 1928) Terra typica: Norway (Europe) ?Africa: • 23°30′N, 12°00′E: Undefined locality, N Africa. Marcus (1928), Rahm (1928) Algeria: • 36°53′N, 07°39′E; 800 m asl: Annaba Province, Massif of Eldough [Edough Mountains]. Binda & Pilato (1987) • 36°40′N, 04°36′E; 1,400 m asl: Tizi Ouzou Province, Akfadou Forest. Binda & Pilato (1987) • 36°38′N, 05°45′E; 1,200 m asl: Jijel Province, M’Sid-Ech-Cheta (Texenna Pass). Binda & Pilato (1987) Record numbers. ?Africa: 1, Algeria: 3; total 4. Remarks. The morphological descriptions of E. merokensis in various reports exhibit a great deal of variation, suggestive of a species complex (see: Ramazzotti & Maucci 1983). However, most of the records (including the type locality) are Holarctic (McInnes 1994), which implies the records from other regions need confirmation.

44. Pseudechiniscus suillus (Ehrenberg, 1853) sensu lato [T] E. aRctomys Ehr. (Murray 1907a) Echiniscus aRctomys Ehrenbg. (Richters 1908) Echiniscus suillus Ehrenberg. (Richters 1911) Pseudechiniscus suillus (Ehr.) (Murray 1913) Pseudechiniscus suillus (Ehrenbg.) Marcus (1928) P. suillus (Ehrbg.) (Marcus 1936) Pseudechiniscus suillus Ehrenberg, 1853 (Rahm 1936) Pseudechiniscus suillus (Ehrbg.) forma Zavattarii n. (Pardi 1941) Pseudechiniscus suillus (Ehrenberg) 1853 (da Cunha & do Nascimento 1964) Pseudechiniscus suillus (Iharos 1969a) Pseudechiniscus suillus Ehrbg. (Iharos (1978) Pseudechiniscus suillus (Ramløv & Kristensen 1985) Pseudechiniscus suillus (Ehrbg) (Maucci 1993) Terra typica: Switzerland (Europe) Angola: • 07°47′S, 15°01′E; 650 m asl: Uíge Province, near Carmona [Uíge], Estate at Pumba Loge, mosses. da Cunha & do Nascimento (1964) Cameroon: • 04°11′N, 09°12′E; 1,900–1,950 m asl: Southwest Region, Mount Cameroon National Park, Mount Cameroon, primeval forest, above bothy No. 1, moss on trees and lava. Iharos (1969a) Ethiopia: • 04°03′N, 38°19′E; 1,700 m asl: Oromia Region, Mēga, moss (NeckeRa sp.) on tree (JunipeRus pRoceRa). Pardi (1941) • Israel and Palestinian National Authority: • 31°51′N, 35°25′E; - 100 m asl: near Jericho, St. George's Monastery, Wadi Qelt, extreme arid climate, lichen (XanthoRia paRietina L.). Rahm (1936) Kenya: • 00°01′S, 37°54′E: Undefined locality, British East Africa [Kenya], moss. Murray (1913) Morocco: • 32°25′N, 05°13′W; 1,750 m asl: Meknès-Tafilalet Region, S of Azrou, cedar forest, moss on limestone boulders in the forest glade. Ramløv & Kristensen (1985) • 31°08′N, 07°55′W; 1,700 m asl: Tadla-Azilal Region, Imlil, walnut forest, moss on limestone cliffs. Ramløv & Kristensen (1985) • 31°06′N, 07°55′W; 2,300–2,500 m asl: Tadla-Azilal Region, path to the Jebel Toubkal, scattered tussock vegetation, moss on basalt and mica slate. Ramløv & Kristensen (1985) Republic of South Africa: • 23°50′S, 29°59′E; 1,450 m asl: N Transvaal [Limpopo Province], Woodbush [Woodbush Forest Reserve]. Murray (1913) • 26°03′S, 27°49′E; 1,500 m asl: Gauteng Province, NE Usumbara. Richters (1911) • 29°44′S, 22°44′E: Undefined locality, Cape Colony [Cape Province]. Murray (1907a, 1913) (Marcus 1928) Tanzania: • 03°07′S, 37°21′E; ca. 4,000 m asl: Kilimanjaro Region, Kilimanjaro National Park, Barranco trail, above tree line, slow running ice covered brooklet, pH 5.5, water temperature 5°C, benthos sample. Van Rompu et al. (1991a) Tunisia: • 36°46′N, 08°41′E; 700 m asl: Jendouba Governorate, surroundings of Ain Draham, moss and lichen on tree (QueRcus saubeR). Iharos (1978) Uganda: • 01°22′N, 32°17′E: Undefined locality. Murray (1907b, 1913) Record numbers. Angola: 1, Cameroon: 1, Ethiopia: 1, Israel: 1, Kenya: 1, Morocco: 3, Republic of South Africa: 3, Tanzania: 1, Tunisia: 1, Uganda: 1; total: 14. Remarks. There is confusion with some early records of Echiniscus arctomys (e.g. Murray 1907a, b; Richters 1908), which were identified as in error by Richters (1911), corrected to Echiniscus suillus and in Marcus (1928) to Pseudechiniscus suillus. The suillus species complex (see: Fontoura & Morais 2011) was originally described from Europe (Ehrenberg, 1853), and requires a re-description. Other species within this complex (e.g. P. facettalis and P. santomensis Fontoura, Pilato & Lisi, 2010) have been described from the African continent and neighbouring islands, though some descriptions (e.g. Pardi 1941) do not conform to the more standard definition of P. suillus. The presence of P. suillus sensu stricto (in Africa) requires confirmation., Published as part of Kaczmarek, Łukasz, 2017, Annotated zoogeography of non-marine Tardigrada. Part IV: Africa, pp. 1-74 in Zootaxa 4284 (1) on pages 19-20, DOI: 10.11646/zootaxa.4284.1.1, http://zenodo.org/record/1010400, {"references":["Ehrenberg, C. G. (1853) Diagnoses novarum formarum. VeRhandlungen deR Koniglich PReussische Akademie deR Wissenschaften zu BeRlin, 8, 526 - 533.","Murray, J. (1907 a) Some South African Tardigrada. JouRnal of the Royal MicRoscopical Society, 5, 515 - 524. https: // doi. org / 10.1111 / j. 1365 - 2818.1907. tb 01665. x","Richters, F. (1908) Moosbewohner. Wissenschaftliche ERgebnisse deR Schwedischen SudpolaR-Expedition (1901 - 1903), Stockholm, 6, 1 - 16.","Richters, F. (1911) Faune des mousses. TaRdigRades. Duc d'ORleans. Campagne aRctique de 1907. Impr. Sci. C. Buelens, Bruxelles, 20 pp.","Murray, J. (1913) African Tardigrada. JouRnal of the Royal MicRoscopical Society, 2, 136 - 144. https: // doi. org / 10.1111 / j. 1365 - 2818.1913. tb 01014. x","Marcus, E. (1928) Spinnentiere oder Arachoides. IV Bartierchen (Tardigrada). TieRwelt Deutschlands und deR angRenzenden MeeResteile Jena, 12, 1 - 230.","Marcus, E. (1936) TaRdigRada. Das TieRReich, 66, 1 - 340.","Rahm, G. (1936) Tardigraden aus Palastina. ZoologischeR AnzeigeR, 115, 65 - 76.","Pardi, L. (1941) Tardigrada. Missione Biologica Sagan-Omo, Reale Accademia d'Italia, Roma Zoologia, 6, 221 - 232.","Da Cunha, A. X. da & Nascimento Ribeiro, F. (1964) Tardigrados de Angola. GaRcia de ORta, Lisboa, 12, 397 - 406.","Iharos, G. (1969 a) Tardigraden aus Mittelwestafrika. Opuscula Zoologica (Budapest), 9, 115 - 120.","Iharos, G. (1978) Data to the knowledge of the Tardigrada fauna of Tunisia. Folia Entomologica HungaRica, 21, 175 - 177.","Ramlov, H. & Kristensen, R. M. (1985) Bjornedyr, Tardigrada. Marokko - 1985. Dansk NatuRhistoRisk FoRenings Rejse til MaRokko, 8 - 22 ApRil 1985, pp. 27 - 32.","Maucci, W. (1993) Prime notizie su Tardigradi \" Terrestri \" del Madagascar con descrizione di tre specie nuove. Bollettino del Museo Civico StoRia NatuRale di VeRona, 17, 381 - 392.","Van Rompu, E. A., De Smet, W. H. & Bafort, J. M. (1991 a) Some freshwater tardigrades from the Kilimanjaro. Naturwetenschappelijk Tijdschrift, 73, 55 - 62.","Murray, J. (1907 b) Encystment of Tardigrada. TRansactions of the Royal Society of EdinbuRgh, 45, 837 - 854. https: // doi. org / 10.1017 / S 0080456800022869","Fontoura, P. & Morais, P. (2011) Assessment of traditional and geometric morphometrics for discriminating cryptic species of the Pseudechiniscus suillus complex (Tardigrada, Echiniscidae). JouRnal of Zoological Systematics and EvolutionaRy ReseaRch, 49 (Supplement 1), 26 - 33. https: // doi. org / 10.1111 / j. 1439 - 0469.2010.00594. x","Fontoura, P., Pilato, G. & Lisi, O. (2010) First record of Tardigrada from Sao Tome (Gulf of Guinea, Western Equatorial Africa) and description of Pseudechiniscus santomensis sp. nov. (Heterotardigrada: Echiniscidae). Zootaxa, 2564, 31 - 42."]}

35. Echiniscus spiniger Richters, 1904c [T] E. spinigeR (Ramløv & Kristensen 1985) Terra typica: Sweden (Europe) Morocco: • 31°08′N, 07°55′W; 1,700 m asl: Tadla-Azilal Region, Imlil, walnut forest, moss on limestone cliffs. Ramløv & Kristensen (1985) Record numbers. Morocco: 1; total: 1. Remarks. This species recorded from numerous European localities and single sites in Asia, Indonesia, Australia, New Zealand, North and South America (McInnes 1994a). Such a wide geographic distribution suggests a species complex or misidentifications. Nominal species should be considered Palaearctic.

26. Echiniscus merokensis suecicus Thulin, 1911 [T] Echiniscus meRokensis Richters forma suecica Thulin, 1911 (da Cunha & do Nascimento 1964) Terra typica: Sweden (Europe) Angola: • 07°25′S, 15°06′E; 850 m asl: Uíge Province, Mahondo, lichens. da Cunha & do Nascimento (1964) Record numbers. Angola: 1; total: 1. Remarks. A subspecies, which with further analysis may be elevated to species is recorded mainly from European localities, Greenland and Angola (McInnes 1994). Probably a Palaearctic taxon.

36. Echiniscus testudo (Doyère, 1840) [T] [m] Echiniscus testudo Doyère, 1840, Echiniscus testudo Doyère, 1840 subsp. tRifilis Rahm 1925 (Rahm 1936) Echiniscus muscicola (Iharos 1969a) Echiniscus testudo Doy. (Iharos 1978) E. testudo (Ramløv & Kristensen 1985) Echiniscus testudo (Doyére, 1840) (Kaczmarek & Michalczyk 2004a) Terra typica: France (Europe) Cameroon: • 04°12′N, 09°12′E; 2,900–3,000 m asl: Southwest Region, Mount Cameroon National Park, Mount Cameroon, moss on tree and lava blocks. Iharos (1969a) Israel and Palestinian National Authority: • 32°59′N, 35°45′E: Undefined locality, Golan Heights, moss. Jørgensen et al. (2007, 2013) • 31°53′N, 35°27′E; - 200 m asl: old Jericho, excavations, mosses (SyntRichium sp., Hypnum sp. or ToRtula sp.), extreme arid, semi-arid or semi-humid climate, lichens or liverworts (FRullania sp.) on rock, stone walls or trees. Rahm (1936) • 31°52′N, 35°26′E; 0 m asl: Jericho, Quarantania Mt. (The Mount of Temptation), rock, extreme arid, semi-arid or semihumid climate, mosses (SyntRichium sp., Hypnum sp. or ToRtula sp.), lichens or liverworts (FRullania sp.) on rock, stone walls or trees. Rahm (1936) • 31°51′N, 35°25′E; - 100 m asl: near Jericho, St. George's Monastery, Wadi Qelt, extreme arid, semi-arid or semi-humid climate, mosses (SyntRichium sp., Hypnum sp. or ToRtula sp.), lichens or liverworts (FRullania sp.) on rock, stone walls or trees. Rahm (1936) • 31°50′N, 34°59′E; 250 m asl: Judean Mountains, Emmaus (Emmaus Nicopolis), extreme arid, semi-arid or semi-humid climate, mosses (SyntRichium sp., Hypnum sp. or ToRtula sp.), lichens or liverworts (FRullania sp.) on rock, stone walls or trees. Rahm (1936) • 31°50′N, 35°11′E; 900 m asl: Judean Mountains, Nabi Samwil (al-Nabi Samuil), extreme arid, semi-arid or semi-humid climate, mosses (SyntRichium sp., Hypnum sp. or ToRtula sp.), lichens or liverworts (FRullania sp.) on rock, stone walls or trees. Rahm (1936) • 31°50′N, 35°23′E; - 50 m asl: on the road to Jericho, Chan of the Samaritan, extreme arid, semi-arid or semi-humid climate, mosses (SyntRichium sp., Hypnum sp. or ToRtula sp.), lichens or liverworts (FRullania sp.) on rock, stone walls or trees. Rahm (1936) • 31°48′N, 35°13′E; 800 m asl: Judean Mountains, Jerusalem, Tombs of the Sanhedrin [Tombs of the Judges], extreme arid, semi-arid or semi-humid climate, mosses (SyntRichium sp., Hypnum sp. or ToRtula sp.), lichens or liverworts (FRullania sp.) on rock, stone walls or trees. Rahm (1936) • 31°48′N, 35°15′E; 800 m asl: Judean Mountains, Jerusalem, Scopus Mt., The Hebrew University, rocks, extreme arid, semi-arid or semi-humid climate, mosses (SyntRichium sp., Hypnum sp. or ToRtula sp.), lichens or liverworts (FRullania sp.) on rock, stone walls or trees. Rahm (1936) • 31°47′N, 35°14′E; 750 m asl: Judean Mountains, Jerusalem, temples walls, extreme arid, semi-arid or semi-humid climate, mosses (SyntRichium sp., Hypnum sp. or ToRtula sp.), lichens or liverworts (FRullania sp.) on rock, stone walls or trees. Rahm (1936) • 31°47′N, 35°14′E; 750 m asl: Judean Mountains, Jerusalem, Gethsemane, bark of olive trees, extreme arid, semi-arid or semi-humid climate, mosses (SyntRichium sp., Hypnum sp. or ToRtula sp.), lichens or liverworts (FRullania sp.) on rock, stone walls or trees. Rahm (1936) • 31°47′N, 35°14′E; 750 m asl: Judean Mountains, Jerusalem, Gethsemane, rocks from grave of Mary [Church of the Assumption (Mary's Tomb)], extreme arid, semi-arid or semi-humid climate, mosses (SyntRichium sp., Hypnum sp. or ToRtula sp.), lichens or liverworts (FRullania sp.) on rock, stone walls or trees. Rahm (1936) • 31°47′N, 35°14′E; 750 m asl: Judean Mountains, Jerusalem, Scopus Mt., E [W] from The Hebrew University, Wadi et Tur [Wadi al-Joz], extreme arid, semi-arid or semi-humid climate, mosses (SyntRichium sp., Hypnum sp. or ToRtula sp.), lichens or liverworts (FRullania sp.) on rock, stone walls or trees. Rahm (1936) • 31°47′N, 35°15′E; 800 m asl: Judean Mountains, Jerusalem, Olives Mt., Pater Noster church, Russians tower, wall of church, extreme arid, semi-arid or semi-humid climate, mosses (SyntRichium sp., Hypnum sp. or ToRtula sp.), lichens or liverworts (FRullania sp.) on rock, stone walls or trees. Rahm (1936) • 31°46′N, 35°10′E; 650 m asl: Judean Mountains, Jerusalem, Ein Kerem, extreme arid, semi-arid or semi-humid climate, mosses (SyntRichium sp., Hypnum sp. or ToRtula sp.), lichens or liverworts (FRullania sp.) on rock, stone walls or trees. Rahm (1936) • 31°46′N, 35°12′E; 750 m asl: Judean Mountains, Jerusalem, Monastery of the Cross, extreme arid, semi-arid or semihumid climate, mosses (SyntRichium sp., Hypnum sp. or ToRtula sp.), lichens or liverworts (FRullania sp.) on rock, stone walls or trees. Rahm (1936) • 31°46′N, 35°12′E; 750 m asl: Judean Mountains, Jerusalem, Monastery of the Cross, garden in Katamin?, extreme arid, semi-arid or semi-humid climate, mosses (SyntRichium sp., Hypnum sp. or ToRtula sp.), lichens or liverworts (FRullania sp.) on rock, stone walls or trees. Rahm (1936) • 31°46′N, 35°13′E; 800 m asl: Undefined locality, Betschar bei Kilometer 22, extreme arid, semi-arid or semi-humid climate, mosses (SyntRichium sp., Hypnum sp. or ToRtula sp.), lichens or liverworts (FRullania sp.) on rock, stone walls or trees. Rahm (1936) • 31°46′N, 35°14′E; 650 m asl: Judean Mountains, Jerusalem, Aceldama, rock, extreme arid, semi-arid or semi-humid climate, mosses (SyntRichium sp., Hypnum sp. or ToRtula sp.), lichens or liverworts (FRullania sp.) on rock, stone walls or trees. Rahm (1936) • 31°46′N, 35°14′E; 750 m asl: Judean Mountains, Jerusalem, Zion Mt., wall behind the Church of Zion, extreme arid, semiarid or semi-humid climate, mosses (SyntRichium sp., Hypnum sp. or ToRtula sp.), lichens or liverworts (FRullania sp.) on rock, stone walls or trees. Rahm (1936) • 31°46′N, 35°14′E; 750 m asl: Judean Mountains, Jerusalem, Church of Saint Peter in Gallicantu, rock, extreme arid, semiarid or semi-humid climate, mosses (SyntRichium sp., Hypnum sp. or ToRtula sp.), lichens or liverworts (FRullania sp.) on rock, stone walls or trees. Rahm (1936) • 31°43′N, 35°17′E; 600 m asl: Judean Mountains, al-Ubeidiya, Monastery Deir Dosi (Monastery of St. Theodosius), extreme arid, semi-arid or semi-humid climate, mosses (SyntRichium sp., Hypnum sp. or ToRtula sp.), lichens or liverworts (FRullania sp.) on rock, stone walls or trees. Rahm (1936) • 31°42′N, 35°12′E; 800 m asl: Judean Mountains, Bethlehem, extreme arid, semi-arid or semi-humid climate, mosses (SyntRichium sp., Hypnum sp. or ToRtula sp.), lichens or liverworts (FRullania sp.) on rock, stone walls or trees. Rahm (1936) • 31°42′N, 35°20′E; 250 m asl: Great Lavra of St. Sabbas the Sanctified (Mar Saba), extreme arid, semi-arid or semi-humid climate, mosses (SyntRichium sp., Hypnum sp. or ToRtula sp.), lichens or liverworts (FRullania sp.) on rock, stone walls or trees. Rahm (1936) • 31°40′N, 35°15′E; 750 m asl: Judean Mountains, Herodyon National Park, Herodion, extreme arid, semi-arid or semihumid climate, mosses (SyntRichium sp., Hypnum sp. or ToRtula sp.), lichens or liverworts (FRullania sp.) on rock, stone walls or trees. Rahm (1936) • 31°33′N, 35°06′E; 1,000 m asl: Judean Mountains, Hebron, Mamre, extreme arid, semi-arid or semi-humid climate, mosses (SyntRichium sp., Hypnum sp. or ToRtula sp.), lichens or liverworts (FRullania sp.) on rock, stone walls or trees. Rahm (1936) Jordan: • 32°02′N, 35°44′E; 800 m asl: Balqa Governorate, As-Salt, extreme arid, semi-arid or semi-humid climate, mosses (SyntRichium sp., Hypnum sp. or ToRtula sp.), lichens or liverworts (FRullania sp.) on rock, stone walls or trees. Rahm (1936) • 31°50′N, 36°49′E; 500 m asl: Zarqa Governorate, Zarqa [Azarq Wetland Reserve], moss. Kaczmarek & Michalczyk (2004a) Libya: • 32°29′N, 20°50′E; 350 m asl: Marj District, Barce [Marj], moss. Pilato & Pennisi (1976) Morocco: • 32°36′N, 04°32′W; 1,900 m asl: Meknès-Tafilalet Region, Tizi-n-Talrhemt, moss or lichen. Jorgensen et al. (2007) • 31°35′N, 05°36′W; 1,650–1800 m asl: Souss-Massa-Drâa Region, Todra [Todgha] Gorge, sparse grass vegetation and few low bushes, moss on limestone cliffs (orthoceratite lime). Ramløv & Kristensen (1985) • 30°27′N, 07°32′W; 1,850 m asl: Souss-Massa-Drâa Region, Tizi-n-Taghatine, sparse vegetation, mosses and lichens on isolated basalt cliffs. Ramløv & Kristensen (1985), Jorgensen et al. (2007) Tunisia: • 36°46′N, 08°41′E; 700 m asl: Jendouba Governorate, surroundings of Ain Draham, moss on rock. Iharos (1978) • 36°07′N, 08°32′E; 450 m asl: Kef Governorate, along road from El Kef and Sakiet, 21 km W El Kef, “strongly insolated area”, lichen on soil. Iharos (1978) Record numbers. Cameroon: 1, Israel and Palestinian National Authority: 26, Jordan: 2, Libya: 1, Morocco: 3, Tunisia: 2; total: 35. Remarks. The very wide biogeographic range (McInnes 1994a, Jørgensen et al. 2007), suggests a species complex, however most of the reports of this species are Holarctic. Echiniscus testudo has mainly been reported from northern Saharan (= Palaearctic) Africa., Published as part of Kaczmarek, Łukasz, 2017, Annotated zoogeography of non-marine Tardigrada. Part IV: Africa, pp. 1-74 in Zootaxa 4284 (1) on pages 16-17, DOI: 10.11646/zootaxa.4284.1.1, http://zenodo.org/record/1010400, {"references":["Doyere, P. L. N. (1840) Memoire sur les Tardigrades. Annales des Sciences NatuRelles, Series 2 (Zoologie), 14, 269 - 362.","Rahm, G. (1936) Tardigraden aus Palastina. ZoologischeR AnzeigeR, 115, 65 - 76.","Iharos, G. (1969 a) Tardigraden aus Mittelwestafrika. Opuscula Zoologica (Budapest), 9, 115 - 120.","Iharos, G. (1978) Data to the knowledge of the Tardigrada fauna of Tunisia. Folia Entomologica HungaRica, 21, 175 - 177.","Ramlov, H. & Kristensen, R. M. (1985) Bjornedyr, Tardigrada. Marokko - 1985. Dansk NatuRhistoRisk FoRenings Rejse til MaRokko, 8 - 22 ApRil 1985, pp. 27 - 32.","Kaczmarek, L. & Michalczyk, L. (2004 a) The first record of water bears (Tardigrada) from Jordan. DiRasat (PuRe Sciences), 31 (2), 154 - 157.","Jorgensen, A., Mobjerg, N. & Kristensen, R. M. (2007) A molecular study of the tardigrade Echiniscus testudo (Echiniscidae) reveals low DNA sequence diversity over a large geographic area. JouRnal of Limnology, 66, 77 - 83. https: // doi. org / 10.4081 / jlimnol. 2007. s 1.77","Jorgensen, A., Faurby, S., Persson, D. K., Halberg, K. A., Kristensen, R. M. & Mobjerg, N. (2013) Genetic diversity in the parthenogenetic reproducing tardigrade Echiniscus testudo (Heterotardigrada: Echiniscoidea). Journal of Limnology, 72 (Supp 1 ment), 136 - 143. https: // doi. org / 10.4081 / jlimnol. 2013. s 1. e 17","Pilato, G. & Pennisi, G. (1976) Prime notizie sui Tardigradi della Cirenaica. Animalia, 3, 243 - 258.","McInnes, S. J. (1994) Zoogeographic distribution of terrestrial / freshwater tardigrades from current literature. JouRnal of NatuRal HistoRy, 28, 257 - 352. https: // doi. org / 10.1080 / 00222939400770131"]}

17. Echiniscus crassispinosus Murray, 1907a sensu stricto [T] E. cRassispinosus sp. n. (Murray 1907a) Echiniscus cRassispinosus Murray (?) (Murray 1913) Echiniscus cRassispinosus J. Murray, 1907 (Teunissen 1938) Terra typica: Republic of South Africa (Africa) Democratic Republic of Congo: • 01°17′S, 29°31′E; 1,750 m asl: North Kivu Province, Albert National Park [Virunga National Park], Tshengelero [Tshengere] (swamp N of Munagana), soil. Teunissen (1938) Kenya: • 01°15′S, 36°41′E; 2,000 m asl: Kiambu County, Kikuyu, moss. Murray (1913) Republic of South Africa: • 29°44′S, 22°44′E: Type Locality: Undefined locality, Cape Colony [Cape Province]. Murray (1907a) Record numbers. Democratic Republic of Congo: 1, Kenya: 1, Republic of South Africa: 1; total: 3. Remarks. Originally described from Africa this species also has single reports from Asia, Central and South America (McInnes 1994a, Kaczmarek et al. 2014a, Kaczmarek et al. 2015). The limited early description of E. crassispinosus (Murray, 1907a) requires new material from the type locale to confirm both the species and its biogeographic description., Published as part of Kaczmarek, Łukasz, 2017, Annotated zoogeography of non-marine Tardigrada. Part IV: Africa, pp. 1-74 in Zootaxa 4284 (1) on page 12, DOI: 10.11646/zootaxa.4284.1.1, http://zenodo.org/record/1010400, {"references":["Murray, J. (1907 a) Some South African Tardigrada. JouRnal of the Royal MicRoscopical Society, 5, 515 - 524. https: // doi. org / 10.1111 / j. 1365 - 2818.1907. tb 01665. x","Murray, J. (1913) African Tardigrada. JouRnal of the Royal MicRoscopical Society, 2, 136 - 144. https: // doi. org / 10.1111 / j. 1365 - 2818.1913. tb 01014. x","Teunissen, R. J. H. (1938) Tardigraden. ExploRation du PaRc National AlbeRt, Mission de Witte, 16, 1 - 21.","McInnes, S. J. (1994) Zoogeographic distribution of terrestrial / freshwater tardigrades from current literature. JouRnal of NatuRal HistoRy, 28, 257 - 352. https: // doi. org / 10.1080 / 00222939400770131","Kaczmarek, L, Michalczyk, L. & McInnes, S. J. (2015) Annotated zoogeography of non-marine Tardigrada. Part II: South America. Zootaxa, 3923 (1), 1 - 107. https: // doi. org / 10.11646 / zootaxa. 3923.1.1"]}

11. Echiniscus bisculptus Maucci, 1983 [T] Echiniscus bisculptus sp. nov. (Maucci 1983) Terra typica: Morocco (Africa) Morocco: • 34°04′N, 04°07′W; 1,500 m asl: Taza [Taza-Al Hoceima-Taounate Region], Bab Bou Idir, in the sun, moss on rock. Maucci (1983) Record numbers. Morocco: 1; total: 1. Remarks. Only recorded from Portugal and Morocco (McInnes 1994); a possible Palaearctic taxon.

1. Echiniscoides sigismundi (M. Schultze, 1865) sensu lato Terra typica: North Sea (Belgium, Europe) Democratic Republic of Congo: • 00°47′S, 29°18′E; 1,000 m asl: South Kivu Province, Albert National Park [Virunga National Park], Rwindi, soil sample. Teunissen (1938) Record numbers. Democratic Republic of Congo: 1; total: 1. Remarks. This is an unusual report of a coastal marine genus that was described from an inland soil environment. The description of 12 females, one male and 19 juveniles plus eggs suggests this is a valid identification. Whether the original samples were the result of contaminated sampling gear or this species truly occurs within the African Rift, requires further exploration.

30. Echiniscus pusae Marcus, 1928 [T] Echiniscus sp. (Murray 1907a) Echiniscus pusae nom. nov. (Marcus 1928) Echiniscus pusae Marcus, 1928 (Pilato et al. 2003) Terra typica: Republic of South Africa (Africa) Democratic Republic of Congo: • 01°10′S, 28°28′E; 1,000 m asl: North Kivu Province, outskirts of Mohanga, liverwort (Plagiochila poRelloides). Pilato et al. (2003) Republic of South Africa: • 29°44′S, 22°44′E: Type Locality: Undefined locality, Cape Colony [Cape Province]. (Murray 1907a), Marcus (1928) Record numbers. Democratic Republic of Congo: 1, Republic of South Africa: 1; total: 2. Remarks. Only reported from Africa and Indonesia (Lombok) (McInnes 1994); a possible tropical/ subtropical distribution.

20. Echiniscus granulatus (Doyère, 1840) [T] Echiniscus (E.) gRanulatus (Doy., 1840) (Binda 1971) E. gRanulatus (Ramløv & Kristensen 1985) E. (E.) gRanulatus (Doyère, 1840) (Séméria (1986) Terra typica: France (Europe) Algeria: • 36°28′N, 04°14′E; 1,700 m asl: Tizi Ouzou Province, Massif of Djurdjura (Assouille depression). Binda & Pilato (1987) Morocco: • 33°31′N, 05°07′W; 1,650 m asl: Meknès-Tafilalet Region, Ras al Ma [Res el Ma], mosses on trees (Cedrus) and rocks. Séméria (1986) • 32°56′N, 05°40′W; 1,700 m asl [850 m asl]: Meknès-Tafilalet Region, Khénifra, moss. Binda (1971) • 32°25′N, 05°13′W; 1,750 m asl: Meknès-Tafilalet Region, S of Azrou, cedar forest, moss on limestone boulders in the forest glade and moss on soil, fallen branches and stems (2 samples). Ramløv & Kristensen (1985) • 31°08′N, 07°55′W; 1,700 m asl: Tadla-Azilal Region, Imlil, walnut forest, moss on limestone cliffs. Ramløv & Kristensen (1985) Record numbers. Algeria: 1, Morocco: 5; total: 6. Remarks. Originally described from France, this species has mainly been reported from Europe and Asia, indicating a Palaearctic distribution. The few records outside this area (see: McInnes 1994), need confirmation., Published as part of Kaczmarek, Łukasz, 2017, Annotated zoogeography of non-marine Tardigrada. Part IV: Africa, pp. 1-74 in Zootaxa 4284 (1) on pages 12-13, DOI: 10.11646/zootaxa.4284.1.1, http://zenodo.org/record/1010400, {"references":["Doyere, P. L. N. (1840) Memoire sur les Tardigrades. Annales des Sciences NatuRelles, Series 2 (Zoologie), 14, 269 - 362.","Ramlov, H. & Kristensen, R. M. (1985) Bjornedyr, Tardigrada. Marokko - 1985. Dansk NatuRhistoRisk FoRenings Rejse til MaRokko, 8 - 22 ApRil 1985, pp. 27 - 32.","Semeria, Y. (1986) Notes sur les Tardigrades du Maroc. Biocosme mesogeen, Nice, 3, 117 - 119.","McInnes, S. J. (1994) Zoogeographic distribution of terrestrial / freshwater tardigrades from current literature. JouRnal of NatuRal HistoRy, 28, 257 - 352. https: // doi. org / 10.1080 / 00222939400770131"]}

43. Pseudechiniscus pseudoconifer Ramazzotti, 1943 Terra typica: Italy (Europe) Uganda: • 01°22′N, 32°17′E: Undefined locality. Binda (1974) Record numbers. Uganda: 1; total: 1. Remarks. A passing comment by Binda (1974) places this species in Uganda.

39. Parechiniscus chitonides Cuénot, 1926 [T] Terra typica: France (Europe) Libya: • 32°29′N, 20°50′E; 350 m asl: Marj District, Barce [Marj], moss. Pilato & Pennisi (1976) Record numbers. Libya: 1; total: 1. Remarks. Palaearctic.

16. Echiniscus cirinoi Binda & Pilato, 1993 [T] Echiniscus ciRinoi n. sp. (Binda & Pilato 1993) Terra typica: Tanzania (Africa) Tanzania: • 03°12′S, 37°31′E; 2,200–2,400 m asl: Type Locality: Kilimanjaro Region, near Marangu [gate], outskirts of the entrance of Kilimanjaro National Park, mosses. Binda & Pilato (1993) Record numbers. Tanzania: 1; total: 1. Remarks. This species is currently endemic to Tanzania.

8. Echiniscus angolensis da Cunha & do Nascimento Ribeiro, 1964 [T] Echiniscus angolensis da Cunha & Nascimento Ribeiro, 1964 (Van Rompu et al. 1991a) Echiniscus angolensis da Cunha & Ribeiro, 1964 (Middleton 2003) Terra typica: Angola (Africa) Angola: • 15°01′S, 13°22′E; 1,950 m asl: Type Locality: Huíla Province, Sá da Bandeira [Lubango], Boca da Humpata, lichens. da Cunha & do Nascimento (1964) Botswana: • 19°10′S, 23°25′E; 950 m asl: North West District, Moremi, lichen on tree. Middleton (2003) Tanzania: • 03°17′S, 37°31′E; ca. 1,500 m asl: Kilimanjaro Region, Kilimanjaro National Park, Marangu gate, rain forest, below tree line, river, pH 5.5, water temperature 12°C, benthos sample. Van Rompu et al. (1991a) Record numbers. Angola: 1, Botswana: 1, Tanzania: 1; total: 3. Remarks. A species with a disjunct distribution, reported from Africa and South and Central America (McInnes 1994, Kaczmarek et al. 2014, 2015), which potentially hides either more than one cryptic species, a rare species with very specific habitat requirements, or a species with Pan-tropical distribution.

40. Pseudechiniscus bispinosus (Murray, 1907a) [T] E. bispinosus sp. n. (Murray 1907a) Terra typica: Republic of South Africa (Africa) Republic of South Africa: • 29°44′S, 22°44′E: Type Locality: Undefined locality, Cape Colony [Cape Province]. Murray (1907a) Record numbers. Republic of South Africa: 1; total: 1. Remarks. This rare species is based on the single original reference from Africa (Murray 1907a) and two early reports from South America (Kaczmarek et al. 2015). As no type material is known it may prove difficult to find neotype material., Published as part of Kaczmarek, Łukasz, 2017, Annotated zoogeography of non-marine Tardigrada. Part IV: Africa, pp. 1-74 in Zootaxa 4284 (1) on page 18, DOI: 10.11646/zootaxa.4284.1.1, http://zenodo.org/record/1010400, {"references":["Murray, J. (1907 a) Some South African Tardigrada. JouRnal of the Royal MicRoscopical Society, 5, 515 - 524. https: // doi. org / 10.1111 / j. 1365 - 2818.1907. tb 01665. x","Kaczmarek, L, Michalczyk, L. & McInnes, S. J. (2015) Annotated zoogeography of non-marine Tardigrada. Part II: South America. Zootaxa, 3923 (1), 1 - 107. https: // doi. org / 10.11646 / zootaxa. 3923.1.1"]}

24. Echiniscus mediantus Marcus, 1930 [T] E. mediantus (Ramløv & Kristensen 1985) Terra typica: Scotland (Europe) Morocco: • 33°04′N, 05°00′W; 2,000 m asl [2,100 m asl]: Meknès-Tafilalet Region, near Aguelmane de Si-Ali (Col du Zad) [Aguelmame Sidi Ali lake], scattered basalt blocks (lava field) lying on the shores of the lake, mosses taken on basalt blocks. Ramløv & Kristensen (1985) Record numbers. Morocco: 1; total: 1. Remarks. With reports mainly from Europe, Greenland and Morocco (McInnes 1994), this species has a Palaearctic distribution., Published as part of Kaczmarek, Łukasz, 2017, Annotated zoogeography of non-marine Tardigrada. Part IV: Africa, pp. 1-74 in Zootaxa 4284 (1) on page 13, DOI: 10.11646/zootaxa.4284.1.1, http://zenodo.org/record/1010400, {"references":["Marcus, E. (1930) Beitrage zur Tardigradensystematik. Zoologische JahRbucheR Abteilung fuR Systematik, Okologie und GeogRaphie deR TieRe, 59, 363 - 386.","Ramlov, H. & Kristensen, R. M. (1985) Bjornedyr, Tardigrada. Marokko - 1985. Dansk NatuRhistoRisk FoRenings Rejse til MaRokko, 8 - 22 ApRil 1985, pp. 27 - 32.","McInnes, S. J. (1994) Zoogeographic distribution of terrestrial / freshwater tardigrades from current literature. JouRnal of NatuRal HistoRy, 28, 257 - 352. https: // doi. org / 10.1080 / 00222939400770131"]}

7. Echiniscus africanus Murray, 1907a [T] E. afRicanus sp. n. (Murray 1907a) Echiniscus afRicanus Murr. (Murray 1913) Echiniscus afRicanus John Murray 1907 (da Cunha & do Nascimento 1964) Echiniscus afRicanus Murray 1907 (Binda & Pilato 1995a) Echiniscus cf. afRicanus (Murray, 1907) (Middleton 2003) Terra typica: Republic of South Africa (Africa) Angola: • 07°47′S, 15°01′E; 650 m asl: Uíge Province, near Carmona [Uíge], Estate at Pumba Loge, mosses. da Cunha & do Nascimento (1964) Lesotho: • 29°19′S, 27°29′E; 1,550 m asl: Maseru District, Maseru, moss or lichen on tree or rock. Middleton (2003) • 29°38′S, 27°31′E; 1,750 m asl: Maseru District, Morija, moss or lichen on tree or rock. Middleton (2003) Republic of South Africa: • 23°50′S, 29°59′E; 1,450 m asl: N Transvaal [Limpopo Province], Woodbush [Woodbush Forest Reserve]. Murray (1913) • 29°44′S, 22°44′E: Type Locality: Undefined locality, Cape Colony [Cape Province]. Murray (1907a) Tanzania: • 03°12′S, 37°31′E; 2,200–2,400 m asl: Kilimanjaro Region, near Marangu [gate], outskirts of the entrance of Kilimanjaro National Park, mosses. Binda & Pilato (1995a) Record numbers. Angola: 1, Lesotho: 2, Republic of South Africa: 2, Tanzania: 1; total: 6. Remarks. Species only recorded from continental Africa and an uncertain record from Vietnam (Węglarska 1962).

13. Echiniscus blumi Richters, 1903 [T] [m] Echiniscus blumi (Ramløv & Kristensen 1985) Echiniscus (Echiniscus) blumi Richters, 1903 (Séméria 1986) Terra typica: Svalbard (Spitsbergen) Israel and Palestinian National Authority: • 31°42′N, 35°12′E; 800 m asl: Judean Mountains, Bethlehem, steppe, semi-arid climate, moss (BRyum sp.) on rock. Rahm (1936) Libya: • 32°29′N, 20°50′E; 350 m asl: Marj District, Barce [Marj], moss. Pilato & Pennisi (1976) Morocco: • 33°31′N, 05°07′W; 1,650 m asl: Meknès-Tafilalet Region, Ras al Ma [Res el Ma], mosses and lichens on trees (CedRus) and rocks. Séméria (1986) • 33°27′N, 05°13′W; 1,250 m asl: Meknès-Tafilalet Region, ca. 30 [60] km S of Meknes, Azrou, mosses and lichens. Séméria (1986) • 33°04′N, 05°00′W; 2,000 m asl [2,100 m asl]: Meknès-Tafilalet Region, near Aguelmane de Si-Ali (Col du Zad) [Aguelmame Sidi Ali lake], scattered basalt blocks (lava field) lying on the shores of the lake, mosses on basalt blocks. Ramløv & Kristensen (1985) • 32°25′N, 05°13′W; 1,750 m asl: Meknès-Tafilalet Region, S of Azrou, cedar forest, moss on basalt boulders in the forest glade. Ramløv & Kristensen (1985) • 31°06′N, 07°55′W; 2,300–2,500 m asl: Tadla-Azilal Region, path to the Jebel Toubkal, scattered tussock vegetation, moss on basalt and mica slate. Ramløv & Kristensen (1985) • 30°27′N, 07°32′W; 1,850 m asl: Souss-Massa-Drâa Region, Tizi-n-Taghatine, sparse vegetation, mosses and lichens on isolated basalt cliffs. Ramløv & Kristensen (1985) Tunisia: • 36°46′N, 08°41′E; 650 m asl: Jendouba Governorate, surroundings of Ain Draham [Ayn Drahim]. Binda & Pilato (1987) Record numbers. Israel and Palestinian National Authority: 1, Libya: 1, Morocco: 6, Tunisia: 1; total: 9. Remarks. This species is believed to be almost cosmopolitan, however, the locus typicus is in the Arctic and most of the records are from the Northern Hemisphere (Europe and North America) (McInnes 1994). Echiniscus blumi should perhaps be considered Holarctic, with records from other parts of the world requiring confirmation (Kaczmarek et al. 2015)., Published as part of Kaczmarek, Łukasz, 2017, Annotated zoogeography of non-marine Tardigrada. Part IV: Africa, pp. 1-74 in Zootaxa 4284 (1) on page 11, DOI: 10.11646/zootaxa.4284.1.1, http://zenodo.org/record/1010400, {"references":["Ramlov, H. & Kristensen, R. M. (1985) Bjornedyr, Tardigrada. Marokko - 1985. Dansk NatuRhistoRisk FoRenings Rejse til MaRokko, 8 - 22 ApRil 1985, pp. 27 - 32.","Semeria, Y. (1986) Notes sur les Tardigrades du Maroc. Biocosme mesogeen, Nice, 3, 117 - 119.","Rahm, G. (1936) Tardigraden aus Palastina. ZoologischeR AnzeigeR, 115, 65 - 76.","Pilato, G. & Pennisi, G. (1976) Prime notizie sui Tardigradi della Cirenaica. Animalia, 3, 243 - 258.","McInnes, S. J. (1994) Zoogeographic distribution of terrestrial / freshwater tardigrades from current literature. JouRnal of NatuRal HistoRy, 28, 257 - 352. https: // doi. org / 10.1080 / 00222939400770131","Kaczmarek, L, Michalczyk, L. & McInnes, S. J. (2015) Annotated zoogeography of non-marine Tardigrada. Part II: South America. Zootaxa, 3923 (1), 1 - 107. https: // doi. org / 10.11646 / zootaxa. 3923.1.1"]}

38. Echiniscus wendti Richters, 1903 [T] Echiniscus wendti (Iharos 1969a) Terra typica: Svalbard (Spitsbergen) Cameroon: • 04°10′N, 09°13′E; 1,300–1,500 m asl: Southwest Region, Mount Cameroon National Park, Mount Cameroon, rainforest, moss on tree. Iharos (1969a) Record numbers. Cameroon: 1; total: 1. Remarks. This member of the arctomys group (see E. reticulatus above) has a largely Holarctic distribution (McInnes 1994; Kaczmarek et al. 2016); all other records need confirmation (Kaczmarek et al. 2015)., Published as part of Kaczmarek, Łukasz, 2017, Annotated zoogeography of non-marine Tardigrada. Part IV: Africa, pp. 1-74 in Zootaxa 4284 (1) on page 18, DOI: 10.11646/zootaxa.4284.1.1, http://zenodo.org/record/1010400, {"references":["Iharos, G. (1969 a) Tardigraden aus Mittelwestafrika. Opuscula Zoologica (Budapest), 9, 115 - 120.","McInnes, S. J. (1994) Zoogeographic distribution of terrestrial / freshwater tardigrades from current literature. JouRnal of NatuRal HistoRy, 28, 257 - 352. https: // doi. org / 10.1080 / 00222939400770131","Kaczmarek, L, Michalczyk, L. & McInnes, S. J. (2015) Annotated zoogeography of non-marine Tardigrada. Part II: South America. Zootaxa, 3923 (1), 1 - 107. https: // doi. org / 10.11646 / zootaxa. 3923.1.1"]}

14. Echiniscus canadensis Murray, 1910 [T] E. canadensis (Ramløv & Kristensen 1985) Terra typica: Canada (North America) Algeria: • 34°40′N, 06°11′E; 0 m asl: Biskra Province, 30 km from Biskra, El Oued [Zeribet el Oued]. Binda & Pilato (1987) Morocco: • 33°04′N, 05°00′W; 2,000 m asl [2,100 m asl]: Meknès-Tafilalet Region, near Aguelmane de Si-Ali (Col du Zad) [Aguelmame Sidi Ali Lake], scattered basalt blocks (lava field) lying on the shores of the lake, mosses on basalt blocks. Ramløv & Kristensen (1985) • 30°27′N, 07°32′W; 1,850 m asl: Souss-Massa-Drâa Region, Tizi-n-Taghatine, sparse vegetation, mosses and lichens on isolated basalt cliffs. Ramløv & Kristensen (1985) Record numbers. Algeria: 1, Morocco: 2; total: 3. Remarks. Species with a typical Holarctic distribution, widely distributed in Europe and North America (McInnes 1994) (the uncertain records from Argentina need confirmation – Kaczmarek et al. 2015)., Published as part of Kaczmarek, Łukasz, 2017, Annotated zoogeography of non-marine Tardigrada. Part IV: Africa, pp. 1-74 in Zootaxa 4284 (1) on page 11, DOI: 10.11646/zootaxa.4284.1.1, http://zenodo.org/record/1010400, {"references":["Murray, J. (1910) Tardigrada. British Antarctic Expedition 1907 - 1909. RepoRts on the Scientific Investigations, 1 (Biology, Part V), 83 - 187.","Ramlov, H. & Kristensen, R. M. (1985) Bjornedyr, Tardigrada. Marokko - 1985. Dansk NatuRhistoRisk FoRenings Rejse til MaRokko, 8 - 22 ApRil 1985, pp. 27 - 32.","McInnes, S. J. (1994) Zoogeographic distribution of terrestrial / freshwater tardigrades from current literature. JouRnal of NatuRal HistoRy, 28, 257 - 352. https: // doi. org / 10.1080 / 00222939400770131","Kaczmarek, L, Michalczyk, L. & McInnes, S. J. (2015) Annotated zoogeography of non-marine Tardigrada. Part II: South America. Zootaxa, 3923 (1), 1 - 107. https: // doi. org / 10.11646 / zootaxa. 3923.1.1"]}

27. Echiniscus migiurtinus Franceschi, 1957 [T] Echiniscus (Echiniscus) migiuRtinus sp. nov. (Franceschi 1957) Terra typica: Somalia (Africa) Somalia: • 11°20′N, 49°45′E; 2,000 m asl: Bari Region, Migiurtinia, Uar Medò mountain group [Warmadoow] lichen. Franceschi (1957) Record numbers. Somalia: 1; total: 1. Remarks. This species has only two reports from the type locale and China (Yang 2002a), but the disjunct distribution suggests an error. Further analysis is required to verify this species and whether it is endemic to Somalia.

34. Echiniscus scabrospinosus Fontoura, 1982 [T] Echiniscus scabRospinosus Fontoura 1982 (Binda & Pilato 1995a) Terra typica: Portugal (Europe) Tanzania: • 03°12′S, 37°31′E; 2,200–2,400 m asl: Kilimanjaro Region, near Marangu [gate], outskirts of the entrance of Kilimanjaro National Park, mosses. Binda & Pilato (1995a) Record numbers. Tanzania: 1; total: 1. Remarks. This rare species has only been reported as single specimens from its type locale (Portugal), the Azores (Fontoura et al. 2008a) and Tanzania.

Kaczmarek, Łukasz (2017): Annotated zoogeography of non-marine Tardigrada. Part IV: Africa. Zootaxa 4284 (1): 1-74, DOI: https://doi.org/10.11646/zootaxa.4284.1.1