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4.3. Re-defining Echiniscus Currently, the definition says “Echiniscids with red eyes composed of pigment granules. Rigid buccal tube lacking stylet supports or with fine, fibrillar stylet supports. Two pairs of segmental plates. Two or three median plates, sometimes transversally subdivided. Incisions(notches) on caudal (terminal) plate. No pseudosegmental plates. Ventral subcephalic and/or genital plates may be present.” (G&aogon;siorek et al., 2017). The necessary modification obviously concerns the last sentence, which is rephrased as “Ventral plates may be present, usually only in the form of subcephalic and/or genital plates.”. The affinity of E. becki (Schuster and Grigarick, 1966) remains problematic and thus, consequently, I left its classification unchanged. However, it is clear that this species does not belong in Echiniscus because of the presence of black crystalline eyes (Fig. 9B), which, together with the dorsal plate sculpturing dominated by polygonal endocuticular pillars, make it most similar to Diploechiniscus. Ultimately, E. becki does not exhibit dorsolateral spicules and is comparatively larger (adults 290–420 μm in length) than Diploechiniscus oihonnae (adults < 275 μm in length). The last record of this species, endemic to the Western Nearctic, came from Mexico (Nú˜ nez et al., 2021), yet it is clearly erroneous as the reported individuals had reticulum in their dorsal plates, absent in the paratypes of E. becki (Fig. 9A). Therefore, re-sampling the Californian localities is necessary to obtain DNA barcodes and clarify where this species should be classified., Published as part of Gąsiorek, Piotr, 2023, Eventually tested: Phylogenetic position of " Testechiniscus " meridionalis (Murray, 1906) (Heterotardigrada) revealed, pp. 49-60 in Zoologischer Anzeiger 304 on page 58, DOI: 10.1016/j.jcz.2023.02.004, http://zenodo.org/record/8164274, {"references":["Gasiorek, P., Stec, D., Morek, W., Michalczyk, L., 2017. An integrative redescription of Echiniscus testudo (Doy`ere, 1840), the nominal taxon for the class Heterotardigrada (Ecdysozoa: Panarthropoda: Tardigrada). Zool. Anz. 270, 107 - 122. https: // doi. org / 10.1016 / j. jcz. 2017.09.006.","Schuster, R. O., Grigarick, A. A., 1966. New Tardigrada from Western North America: II, Echiniscus. Proc. Biol. Soc. Wash. 79, 127 - 130."]}

3.2. Redescription of Echiniscus meridionalis stat. rev. Figs. 2 – 7, Tables 2–4, Supplementary Material 4 with raw morphometric data. ZooBank registration number: urn:lsid:zoobank.org:pub:8A36A0BF- AA4F–427F–980B-E70DE960695E. 3.2.1. Terra typica South Orkney Islands. Material examined. South Shetland Islands (see Table 1 for detailed data). 3.2.2. Etymology From Latin meridionalis = southern, referring to the distribution of the species in Antarctic and sub-Antarctic archipelagos. An adjective in nominative singular. 3.2.3. Adult females (i.e. from the third instar onwards; measurements and statistics in Table 2) Body small and plump (Figs. 2A and 3A), light yellow to dark orange in alive specimens. Eyes dark red, minute and dissolving immediately in Hoyer’ s medium. Peribuccal cirri relatively long with respect to the body length (Figs. 2A, 3A and 4A); cirrus A of a moderate length, i.e. reaching approximately 1/3 of the body length; all cephalic cirri with well-delineated cirrophores. Cephalic papillae (secondary clavae) slightly longer than (primary) clavae, both barrel-shaped (Fig. 4A). Body appendage configuration A-C-C d -D-D d -E , with most appendages formed as long, flexible cirri. Short, but robust spines present in positions D and D d (Figs. 2A and 3A). Dorsal plate sculpturing comprises: (I) poorly elevated epicuticular granules, with a strong tendency to merging in all plate portions but the anterior portions of paired segmental plates I–II, median plate 2, and the much reduced median plate 3, where they are separate (Figs. 2A and 3A), (II) pseudopores in the form of delicate depressions distributed irregularly throughout the dorsum, and (III) pores, present mainly in the cephalic, scapular (Figs. 2A and 3A), and occasionally in the caudal (terminal) plates. Pores may be absent as well. Cephalic plate narrow, with the usual chalice-like anterior incision. Cervical plate even narrower, but clearly present and rectangular, tightly joined with the scapular plate. Scapular plate with a distinct faceting-like pattern formed by a system of poorly elevated ridges (Figs. 2A, 3A and 4A); lateralmost plate portions weakly demarcated from the central portion by a similar ridge (Fig. 4A) that replaced a usual suture/incision. Median plate 1 unipartite, m2 bipartite and much larger, m3 developed as a triangular sculptured surface dominated by the caudal plate. Paired segmental plates I–II identical in size and shape. Caudal plate with large triangular posterior incisions (Fig. 2A) that are joined by a central semicircular ridge (Figs. 2A, 3A and 4A). A weakly marked ridge perpendicular to the former may be developed, delineating a cross-like epicuticular structure. Ventral plate armature variously developed, as plates in some individuals may be almost indiscernible, but the general plate formula stands as VIII:2-1-1-2-2-2-2-2 (Figs. 4B and 5A). Particularly the plate in the second row can be hardly identifiable in PCM (Fig. 5A). Ventral plates are smooth (Fig. 4B), but the endocuticular pillars are easily observable in PCM (Fig. 5A). Besides, no granulation present on venter. Hexapartite gonopore and tripartite, large anus (Figs. 4B and 5B). Legs with evident pulvini embedded in proximal portions of limbs and pedal plates in median portions, developed as wrinkled areas with sparse and occasional pores (Figs. 4A and 6B). Long spine I present (Figs. 4A and 6A); papilla IV large and barrel-shaped, dentate fringe on leg IV with numerous irregular teeth, typically interconnected at their bases (Figs. 2A, 4A and 6B). Claws with branches typically curved, those of legs IV slightly higher than the remaining ones. Claw spurs heteromorphic, as spurs IV are more divergent from the branches compared with spurs I– III. The size and position of spurs on claw branches (ca. 1/3–2/5 of the claw height) identical on all claws (Fig. 6). 3.2.4. Adult males (i.e. from the second/third instar onwards, see remarks; measurements and statistics in Table 3) Sexual dimorphism evinces in several characters of males: thinner and shorter body (Figs. 2B and C, 3B, 4C–D), prominent and swollen clavae (Figs. 2B and C, 3B, 4C–D and 5B), and often better visible (darker in PCM) ventral plates (Fig. 5B). Rarely, all appendages but A and E may be formed as spines (Fig. 2B). Gonopore circular. 3.2.5. Juveniles (i.e. the second instar; measurements and statistics in Table 4) Qualitatively similar to sexually mature females, and smaller than both sexes. Gonopore absent. 3.2.6. Larvae (i.e. the first instar; measurements and statistics in Table 4) Qualitatively similar to later life stages, but smaller than juveniles. Body appendage formula: A-Cd- E , all formed as cirri (Fig. 7). Two claws per leg. Gonopore and anus absent. 3.2.7. DNA markers Single haplotype was found in 18S rRNA (OQ 472387), 28S rRNA (OQ 472389), ITS-1 (OQ 476878), ITS-2 (OQ 476879), and two in COI (OQ 472391–2; p -distance = 0.2%). 3.2.8. Differential diagnosis A strong sexual dimorphism, the unusual dorsal plate sculpturing (in some specimens, epicuticular granules may be barely elevated above the surface of plates, and in an initial observation in PCM such exemplars seem to be sculptureless), and the ventral armature covering an entire venter, replacing the endocuticular granulation typical for Echiniscus, together with the geographic range embracing solely Antarctic and sub-Antarctic islands (Fig. 8), make E. meridionalis stat. rev. easily recognisable and well-differentiated morphologically from all other Echiniscus spp. 3.2.9. Remarks The species can be frequently encountered on King George Island, being one of the two commonest echiniscids in that location (the other is Barbaria jenningsi, often co-inhabiting the same mosses and lichens, see Table 1). An undescribed Pseudechiniscus species, eutardigrades Acutuncus antarcticus, Diphascon rudnickii, and unidentified species of the Mesobiotus furciger group (Short et al., 2022) are other relatively common taxa that co-occur with E. meridionalis stat. rev. However, irrespective of its ubiquity in the area, the population density must be low given that typically few representatives were found in samples (Table 1). Ventral armature is already developed in E. meridionalis stat. rev. at sexually immature life stages (i.e. in larvae and juveniles), which often does not take place in the Echiniscidae., Published as part of Gąsiorek, Piotr, 2023, Eventually tested: Phylogenetic position of " Testechiniscus " meridionalis (Murray, 1906) (Heterotardigrada) revealed, pp. 49-60 in Zoologischer Anzeiger 304 on pages 52-54, DOI: 10.1016/j.jcz.2023.02.004, http://zenodo.org/record/8164274, {"references":["Short, K. A., Sands, C. J., McInnes, S. J., Pisani, D., Stevens, M. I., Convey, P., 2022. An ancient, Antarctic-specific species complex: large divergences between multiple Antarctic lineages of the tardigrade genus Mesobiotus. Mol. Phylogenet. Evol. 170, 107429 https: // doi. org / 10.1016 / j. ympev. 2022.107429."]}

3.3. Revised morphological characters for Echiniscus becki and Echiniscus laterosetosus An analysis of the type material of E. becki revealed new traits crucial in determining its phylogenetic position. The dorsal plate sculpturing comprises tightly arranged, polygonal endocuticular pillars (Fig. 9A); they lack centrally positioned pores typical for the blumi-canadensis complex of species and Testechiniscus (G&aogon;siorek et al., 2018). In contrary to the original description (Schuster and Grigarick, 1966), the species does not have dark red eye spots, which always dissolve in Hoyer’ s medium, but large black crystalline eyes identifiable in all examined paratypes (Fig. 9B). An erroneous ascribing of the red colour most probably resulted from a strong orange-reddish glow enclosing eyes. In fact, already the description in Schuster and Grigarick (1966) raised suspicions as the authors drew well-identifiable eyes in the mounted holotypic female, whereas red carotenoid eyes in echiniscids in the vast majority of cases vanish within an hour from preparing a permanent slide with a tardigrade. Echiniscus becki exhibits also a pair of exceptionally shaped subcephalic plates, which extend from the lateral portions of the head and twist in a boomerang-like manner at the level of pharynx (Fig. 9B). Representatives of E. laterosetosus, collected in the high mountains of Honshu, allowed for providing the morphological details relevant to its taxonomy. Eye spots, drawn in the original description without stating their colour (Ito, 1993), are small and black (Fig. 10A). The dorsal plate sculpturing consists of well-developed endocuticular pillars, each joined by striae with four or five neighbouring pillars, and pores distributed between them (Fig. 10B)., Published as part of Gąsiorek, Piotr, 2023, Eventually tested: Phylogenetic position of " Testechiniscus " meridionalis (Murray, 1906) (Heterotardigrada) revealed, pp. 49-60 in Zoologischer Anzeiger 304 on page 55, DOI: 10.1016/j.jcz.2023.02.004, http://zenodo.org/record/8164274, {"references":["Gasiorek, P., Stec, D., Zawierucha, K., Kristensen, R. M., Michalczyk, L., 2018. Revision of Testechiniscus Kristensen, 1987 (Heterotardigrada: Echiniscidae) refutes the polartemperate distribution of the genus. Zootaxa 4472, 261 - 297. https: // doi. org / 10.11646 / zootaxa. 4472.2.3.","Schuster, R. O., Grigarick, A. A., 1966. New Tardigrada from Western North America: II, Echiniscus. Proc. Biol. Soc. Wash. 79, 127 - 130.","Ito, M., 1993. Taxonomic study on the class Heterotardigrada (Tardigrada) from the northern slope of Mt. Fuji, Central Japan. Edaphologia 50, 1 - 13."]}

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 Pozna&nacute;. Paraneotypes GERMANY • 33 &female;&female;, 22 &male;&male;, 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 Pozna&nacute; • 9 &female;&female;, 6 &male;&male;, 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 G&aogon;siorek, Bochnak, Von&ccaron;ina & 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 G&aogon;siorek & Kristensen, 2018 (GenBank accession number: MK529674, G&aogon;siorek 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, G&aogon;siorek et al. 2019a) with 1% value of the p-distance and the least similar was E. belloporus G&aogon;siorek & Kristensen, 2018 (GenBank accession numbers: MK529702, G&aogon;siorek 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, G&aogon;siorek 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 Pozna&nacute; 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"]}

Members of the genus Echiniscus C.A.S. Schultze, 1840 are mostly unisexual, with thelytokously reproducing females. Therefore, every newly described dioecious species in the genus is particularly interesting. Here, we describe Echiniscus masculinussp. nov. from Gunung Kinabalu, the highest peak of Borneo and the entire Southeast Asia. The new species belongs in the predominantly parthenogenetic E. virginicus complex, and its females are confusingly similar to females of the pantropical E. lineatus Pilato et al., 2008, another member of this group. However, genetic evidence and noticeable sexual dimorphism clearly delineate the new species. Males of E. masculinussp. nov. are unlike females in the body proportions, cuticular sculpturing, and appendage configuration. The new discoveries provide a justification to review the current knowledge about evolution and forms of sexual dimorphism within Echiniscus.

There are no published records of water bears (Phylum Tardigrada) from Swaziland. Two samples of foliose lichen collected in 2010 contained nine tardigrade specimens and one egg belonging to five genera and seven species: Echiniscuscf.quadrispinosus,Milnesiumsp., Milnesiumcf.bohleberi, Hypsibiuscf.convergens,Ramazzottiussp., Macrobiotuscf.pallarii andMinibiotusharrylewisi. Milnesiumsp. resemblesMilnesiumlagniappe, a species from southeastern USA, in its cuticle, and possibly in the number of peribuccal lamellae. Specimens from Lesotho and South Africa previously identified asMilnesiumtardigradumare in fact Milnesiumcf.bohleberi. The habitus ofRamazzottiussp. is consistent withR.theroni, a southern African species, but due to the condition of the specimen the presence of cuticular sculpture cannot be definitively ruled out. Macrobiotuscf.pallarii differs fromM.pallariisensu strictoin some structural details of the egg processes. This is the first record ofMinibiotusharrylewisioutside of its type location in KwaZulu-Natal, South Africa.

Echiniscus tristis G&aogon;siorek & 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)

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"]}

Mainly because of the problems with species delineation, the biogeography of microscopic organisms is notoriously difficult to elucidate. In this contribution, variable nuclear and mitochondrial DNA markers were sequenced from individual specimens representing the Echiniscus virginicus complex that are morphologically indistinguishable under light microscopy (five populations from the temperate Eastern Nearctic and 13 populations from the subtropical and tropical zone). A range of methods was used to dissect components of variability within the complex (Bayesian inference, haplotype networks, Poisson tree processes, automatic barcode gap discovery delineations, principal components analysis and ANOVA). We found deep divergence between the temperate Eastern Nearctic E. virginicus and pantropical Echiniscus lineatus in all three genetic markers. In contrast, intraspecific genetic variation was very low, regardless of the geographical distance between the populations. Moreover, for the first time, statistical predictions of tardigrade geographical distributions were modelled. The factor determining the allopatric geographical ranges of deceptively similar species analysed in this study is most likely to be the type of climate. Our study shows that widespread tardigrade species exist, and both geographical distribution modelling and the genetic structure of populations of the pantropical E. lineatus suggest wind-mediated (aeolian) passive long-distance dispersal.

Echiniscus militaris Murray, 1911 (Figs 28���29) Locus typicus. Castlebar, Ireland. Material examined: One four-clawed juvenile, found together with very numerous Echiniscus granulatus (Doy��re, 1840), E. reticulatus Murray, 1905, Pseudechiniscus suillus, Testechiniscus spitsbergensis, and Richtersius coronifer (Richters, 1903) (sample PL.273; Kobylarzowy Coulouir). Description: Very small (162 ��m), conspicuously pink coloured. Cirrus internus / externus length ratio 67%. Cirrus A 48 ��m long and very thick, lateral appendages B and E of equal lengths (20 ��m), lateral cirri C and D markedly longer (60 and 76 ��m, respectively). All dorsal appendages present, of which C d are the longest (44 ��m vs B d 18 ��m long and D d 33 ��m long). In general, cirri in all B-positions are visibly thinner than other appendages. The difference is especially evident in the width of the dorsal appendage bases (1.9 ��m in B d, 2.8 ��m in C d, and 4.9 ��m in D d, respectively). Plates with well-marked borders, lacking cervical (neck) and third median plates. Sculpture composed of polygons (blumi-canadensis type), with the largest polygons found on the small median plates and the central part of the caudal plate. Ventral cuticle smooth, except for the genital zone, where two large, trapezoid plates, in line with legs IV, occur (Fig. 29). Pedal (leg) plates invisible in PCM. Dentate collar on legs IV comprises seven long, parallel teeth. Large, slender claws; claws I���III markedly shorter than claws IV (11.9���13.1 ��m, 32.5���35.8% vs 16.6 ��m, 45.4%). Small spurs (2.1 ��m, 5.7%) positioned very close to the internal claw bases, and almost perpendicular to the claw branch. Remarks: New record for Poland. E. militaris is without doubt one of the most elusive Palaearctic echiniscid species (Ramazzotti & Maucci 1983). Since its original description in 1911, it has been recorded only a few times and in very small numbers from various European countries (McInnes 1994). Pilato (1977) suppressed all subspecies after analysing different populations of the species. As there is a very high variability among recorded specimens, it is very likely that a number of independent species are captured under the one specific name. Unfortunately, difficulties in obtaining an adequate number of specimens for a detailed taxonomic study, including DNA sequencing, will hinder clarification of the taxonomic status of this species and resolving whether this is a single species exhibiting considerable intraspecific variability or a group of morphologically similar species., Published as part of G��siorek, Piotr & Degma, Peter, 2018, Three Echiniscidae species (Tardigrada: Heterotardigrada) new to the Polish fauna, with the description of a new gonochoristic Bryodelphax Thulin, 1928, pp. 77-96 in Zootaxa 4410 (1) on pages 88-89, DOI: 10.11646/zootaxa.4410.1.4, http://zenodo.org/record/1221257, {"references":["Murray, J. (1911) Arctiscoida. Proceedings of the Royal Irish Academy, 31, 1 - 16.","Doyere, M. L. (1840) Memorie sur les Tardigrades. Annales des Sciences Naturelles. Zoologie, Serie 2, 14, 269 - 362.","Murray, J. (1905) The Tardigrada of the Scottish Lochs. Transactions of the Royal Society of Edinburgh, 41 (27), 677 - 698. https: // doi. org / 10.1017 / S 0080456800035547","Richters, F. (1903) Nordische Tardigraden. Zoologischer Anzeiger, 27, 168 - 172.","Ramazzotti, G. & Maucci, w. (1983) Il Phylum Tardigrada. III edizione riveduta e aggiornata. Memorie dell'Istituto Italiano di Idrobiologia, Pallanza, 41, 1 - 1012.","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","Pilato, G. (1977) Variabilita' in Echiniscus militaris Murray, 1911 (Heterotardigrada) e valore delle sue sottospecie. Animalia, 4 (1 - 2), 5 - 21."]}

Echiniscus spiniger Richters, 1904 (Fig. 34) Locus typicus. Visby, Gotland, Sweden. Material examined. Three adult specimens and one juvenile, found together with exceptionally abundant Echiniscus granulatus, E. testudo (Doy��re, 1840), and Richtersius coronifer (PL.247; Ko&sacute;cieliska Valley. Description. Medium-sized (160���313 ��m), dark-yellow. Cirrus A long (104 ��m, cirrus A/ body length ratio 33%). Cirrus internus / externus length ratio 60���65%. Plate sculpture composed of small and medium-sized pores (spinulosus type), with the largest pores found on the large median plates and on the caudal plate. Lateral and dorsal appendages at B, C, C d, D, D d, and E in the shape of long spines, with dorsal spines slightly longer than the lateral ones (38���48 ��m vs 21���36 ��m). Pedal (leg) plates very well-developed and large, those on leg IV forming a collar composed of eight small, blunt teeth. Claws I���III moderately shorter than claws IV (12.5���20.6 ��m, 27.3��� 31.1% vs 16.9���23.6 ��m, 30.3���38.3%). Spurs (1.5���3.9 ��m, 3.4���5.8%) positioned at ~30% of the claw length. Remarks. New record for Poland. Dastych (1988) expressed the opinion that the so-called spinulosus - quadrispinosus group was in need of a complete revision, and produced a schematic illustration of E. spinulosus (Doy��re, 1840) with long lateral appendages. He also remarked that the lateral appendages of Polish E. spinulosus specimens varied significantly in length (Dastych 1988). Presented findings suggest that his records could embrace two distinct species: E. spinulosus s. s., which is a predominantly lowland species, and E. spiniger, more frequently found in colder and upland habitats. In E. spinulosus specimens from the locus typicus (Saint-Maur-des-Foss��s, France), dorsal spines C d and D d are longer than the extremely short lateral spines (similar to the differences in length recorded for E. scabrospinosus Fontoura, 1982, see Figs 31���32). In E. spiniger, the lateral spines are longer and of similar or equal development to the dorsal appendages (Fig. 34). Scottish specimens of E. quadrispinosus Richters, 1902 of identical body size to Polish E. spiniger specimens have dorsal spines C d and D d distinctly shorter than in E. spiniger, and unlike in E. spiniger, spines D d are typically shorter than C d (Fig. 35). Moreover, all lateral appendages in E. quadrispinosus are developed as long cirri with broad bases (Ramazzotti & Maucci 1983). Pilato et al. (2008) showed that different species within spinulosus group can be separated on the basis of differences in plate sculpturing. An interesting pattern can be observed within members of the group: generally, species with short appendages exhibit large and sparsely distributed cuticular pores, whereas those with long spines or cirri possess comparatively smaller, more densely arranged pores (compare Figs 31���32 for E. spinulosus and E. scabrospinosus, and 33���35 for E. lichenorum Maucci, 1983, E. spiniger and E. quadrispinosus). Curiously, E. migiurtinus Franceschi, 1957 with reduced set of trunk appendages (exclusively stout spines D d present) seems to deviate from this rule, since it exhibits small, densely arranged pores (Fig. 30). However, there are also differences in the degree of pedal (leg) plate development between the representatives of the spinulosus group. Such differences could be a stable specific trait. Pedal (leg) plates can be: invisible in PCM on all legs (Fig. 32); visible only on legs IV (Figs 30, 33, 35, arrowheads); or present on all legs, and with strongly developed sculpture similar to that on the dorsal plates, which so far was detected only in E. spinulosus and E. spiniger (Figs 31, 34, arrowheads). It is clear that additional studies are needed to better understand the phylogeny and evolution of morphological traits within this species complex. For the Polish fauna, both E. spiniger and E. spinulosus should be considered as its indigenous components. The presence of E. testudo, a typical lowland species, within this sample, represents the first record for this species from the Polish Tatras (Dastych 1980)., Published as part of G��siorek, Piotr & Degma, Peter, 2018, Three Echiniscidae species (Tardigrada: Heterotardigrada) new to the Polish fauna, with the description of a new gonochoristic Bryodelphax Thulin, 1928, pp. 77-96 in Zootaxa 4410 (1) on pages 89-91, DOI: 10.11646/zootaxa.4410.1.4, http://zenodo.org/record/1221257, {"references":["Richters, F. (1904) Arktische Tardigraden. Fauna Arctica, 3, 495 - 511.","Doyere, M. L. (1840) Memorie sur les Tardigrades. Annales des Sciences Naturelles. Zoologie, Serie 2, 14, 269 - 362.","Dastych, H. (1988) The Tardigrada of Poland. Monografie Fauny Polski, 16, 1 - 255.","Fontoura, P. (1982) Deux nouvelles especes de tardigrades muscicoles du Portugal. Publicacoes do Instituto de Zoologia ' Dr Augusto Nobre', Faculdade de Ciencias do Porto, 165, 5 - 19.","Richters, F. (1902) Neue Moosbewohner. Bericht der Senckenbergischen Naturforschenden Geselschaft in Frankfurt am Main, II. Teil, 23 - 26.","Ramazzotti, G. & Maucci, w. (1983) Il Phylum Tardigrada. III edizione riveduta e aggiornata. Memorie dell'Istituto Italiano di Idrobiologia, Pallanza, 41, 1 - 1012.","Pilato, G., Fontoura, P. & Beasley, C. (2008) New description of Echiniscus scabrospinosus Fontoura, 1982, and description of a new species of Echiniscus (Heterotardigrada) from China. Zootaxa, 1856, 41 - 54.","Maucci, w. (1983) Echiniscus bisculptus n. sp., del Marocco, ed E. lichenorum n. sp., del Portogallo. Atti della Societa Italiana di Scienze Naturali e del Museo Civico di Storia Naturale in Milano, 124 (3 - 4), 257 - 261.","Franceschi, T. (1957) Una nuova specie di Echiniscus. Annali del Museo Civico di Storia Naturale di Genova, 69, 223 - 225.","Dastych, H. (1980) Niesporczaki (Tardigrada) Tatrzanskiego Parku Narodowego. Monografie Fauny Polski, 9, 1 - 232."]}

During a faunistic survey in the Pieniny and Tatra Mountains three species of Echiniscidae new to Poland, including one new to science, were found. Bryodelphax instabilis sp. nov. is characterised by an instable number of ventral plates, deep faceting of the scapular plate, having dorsal plates covered either with pseudopores or pores, and striking sexual dimorphism. Two first records, namely Echiniscus militaris and E. spiniger, are rare taxa of upland-mountain character, which have already been reported from a few European countries since the original description at the beginning of the XXth century. The interspecific appendage length variability and development of pedal plates in the spinulosus group, to which E. spiniger belongs, and their taxonomic importance is discussed. Notes relating to sexual dimorphism within the newly recorded Pseudechiniscus facettalis, are also presented. Echiniscus testudo is reported from the Polish part of the Tatra Mountains for the first time. Succeeding findings confirm the high tardigrade α-diversity in the Polish mountain ranges. An amended key for Polish Heterotardigrada is provided.

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).

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 Med�� 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., Published as part of Kaczmarek, ��ukasz, 2017, Annotated zoogeography of non-marine Tardigrada. Part IV: Africa, pp. 1-74 in Zootaxa 4284 (1) on page 14, DOI: 10.11646/zootaxa.4284.1.1, http://zenodo.org/record/1010400, {"references":["Franceschi, T. (1957) Una nuova specie di Echiniscus. Annali del Museo Civico di StoRia NatuRale di Genova, 69, 223 - 225.","Yang, T. (2002) The tardigrades from some mosses of Lijiang County in Yunnan province (Heterotardigrada: Echiniscidae: Eutardigrada: Parachela: Macrobiotidae, Hypsibiidae). Acta Zootaxonomica Sinica, 27, 53 - 64."]}

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.

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.

35. Echiniscus spiniger Richters, 1904c [T] E. spinigeR (Raml��v & 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. Raml��v & 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., Published as part of Kaczmarek, ��ukasz, 2017, Annotated zoogeography of non-marine Tardigrada. Part IV: Africa, pp. 1-74 in Zootaxa 4284 (1) on page 16, DOI: 10.11646/zootaxa.4284.1.1, http://zenodo.org/record/1010400, {"references":["Richters, F. (1904 c) Beitrag zur Verbreitungen der Tardigraden im sudlichen Scandinavien und an der mecklenburgischen Kuste. ZoologischeR AnzeigeR, 28, 347 - 352.","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"]}

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).

29. Echiniscus pooensis Rodriguez-Roda, 1947 [T] Echiniscus (E.) pooensis sp. n. (Rodriguez-Roda 1947) Terra typica: Equatorial Guinea (Africa) Equatorial Guinea: ��� 03��36���N, 08��46���E; 2,500 m asl: Type Locality: Fernando P��o [Bioko Island], Bioko Norte Province, Pico Basil��, mosses and liverworts between Lycopodium sp. Rodriguez-Roda (1947) Record numbers. Equatorial Guinea: 1; total: 1. Remarks. This species has not been reported again since Rodriguez-Roda (1947), but is currently endemic to Equatorial Guinea., Published as part of Kaczmarek, ��ukasz, 2017, Annotated zoogeography of non-marine Tardigrada. Part IV: Africa, pp. 1-74 in Zootaxa 4284 (1) on page 14, DOI: 10.11646/zootaxa.4284.1.1, http://zenodo.org/record/1010400, {"references":["Rodriguez-Roda, J. (1947) Algunos Tardigrados de Fernando Poo. Publicaciones del Instituto de Biologia Aplicada, 4, 149 - 159."]}

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.

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., Published as part of Kaczmarek, ��ukasz, 2017, Annotated zoogeography of non-marine Tardigrada. Part IV: Africa, pp. 1-74 in Zootaxa 4284 (1) on pages 14-15, DOI: 10.11646/zootaxa.4284.1.1, http://zenodo.org/record/1010400, {"references":["Marcus, E. (1928) Spinnentiere oder Arachoides. IV Bartierchen (Tardigrada). TieRwelt Deutschlands und deR angRenzenden MeeResteile Jena, 12, 1 - 230.","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","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"]}

18. Echiniscus crassispinosus fasciatus Marcus, 1928 [T] Echiniscus cRassispinosus Murray (?) (Murray 1913) E. (E.) cRassispinosus John Murr. forma fasciatus nom. nov. (Marcus 1928) Terra typica: Republic of South Africa (Africa) Republic of South Africa: ��� 01��15���S, 36��41���E; 2,000 m asl: Type Locality: Kiambu County, Kikuyu, moss. Murray (1913), Marcus (1928) Record numbers. Republic of South Africa: 1; total: 1. Remarks. Species recorded from type locality and from three uncertain records from Brazil (for more details see Kaczmarek et al. 2015)., Published as part of Kaczmarek, ��ukasz, 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":["Marcus, E. (1928) Spinnentiere oder Arachoides. IV Bartierchen (Tardigrada). TieRwelt Deutschlands und deR angRenzenden MeeResteile Jena, 12, 1 - 230.","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","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"]}

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)., Published as part of Kaczmarek, ��ukasz, 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":["Richters, F. (1902) Neue Moosbewohner. BeRicht deR SenckenbeRgischen NatuRfoRschenden gesellschaft in FRankfoRt am Main, 1902, 23 - 26.","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.","Meyer, H. A. & Hinton, J. G. (2009) The Tardigrada of southern Africa, with the description of Minibiotus haRRylewisi, a new species from KwaZulu-Natal, South Africa (Eutardigrada: Macrobiotidae). AfRican InveRtebRates, 50 (2), 255 - 268. https: // doi. org / 10.5733 / afin. 050.0203","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"]}

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.

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.

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)., Published as part of Kaczmarek, ��ukasz, 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":["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"]}

23. Echiniscus longispinosus Murray, 1907a [T] E. longispinosus sp. n. (Murray 1907a) Echiniscus longispinosus Murray 1907 (Pilato et al. 1991) 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) ��� 34��01���S, 23��55���E; 200 m asl: Eastern Cape Province, Tzitzikama [Tsitsikamma National Park], moss. Binda (1984), Pilato et al. (1991) Record numbers. Republic of South Africa: 2; total: 2. Remarks. The species is currently endemic to Republic of South Africa., Published as part of Kaczmarek, ��ukasz, 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":["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"]}

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, Ouka��meden, 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)., Published as part of Kaczmarek, ��ukasz, 2017, Annotated zoogeography of non-marine Tardigrada. Part IV: Africa, pp. 1-74 in Zootaxa 4284 (1) on pages 10-11, DOI: 10.11646/zootaxa.4284.1.1, http://zenodo.org/record/1010400, {"references":["Heinis, F. (1908 a) Tardigraden der Schweitz. ZoologischeR AnzeigeR, 32, 633 - 638.","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"]}

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.

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., Published as part of Kaczmarek, ��ukasz, 2017, Annotated zoogeography of non-marine Tardigrada. Part IV: Africa, pp. 1-74 in Zootaxa 4284 (1) on page 15, DOI: 10.11646/zootaxa.4284.1.1, http://zenodo.org/record/1010400, {"references":["Fontoura, P. (1982) Deux nouvelles especes de tardigrades muscicoles du Portugal. Publicacoes do Instituto de Zoologia \" DR. Augusto NobRe \", Faculdade Ciencas do PoRto, 165, 5 - 19.","Binda, M. G. & Pilato, G. (1995 a) Some notes on African tardigrades with description of two new species. TRopical Zoology, 8, 367 - 372. https: // doi. org / 10.1080 / 03946975.1995.10539294"]}

10. Echiniscus bigranulatus Richters, 1908 sensu lato [T] [m] Echiniscus bigRanulatus Richters. (Heinis 1928) Echiniscus bigRanulatus (Iharos 1969a) Echiniscus bigRanulatus Richters, 1907 (Middleton 2003) Terra typica: Argentina (South America) Botswana: ��� 24��25���S, 25��31���E; 1,150 m asl: Kweneng District, Molepolole, lichen on tree. Middleton (2003) Cameroon: ��� 04��57���N, 09��56���E; 800 m asl: Littoral Region, Kongsamba [Nkongsamba], moss on tree. Iharos (1969a) Republic of South Africa: ��� 30��34���S, 22��56���E: Undefined locality. Heinis (1928) Record numbers. Botswana: 1, Cameroon: 1, Republic of South Africa: 1; total: 3. Remarks. This bona fide species belongs to the Neotropical and Antarctic bigranulatus group but is easily mistaken for other siblings (see: Michalczyk & Kaczmarek 2006a, 2007). All records prior to 2006 require confirmation, while the three uncertain records from Africa require re-examination and confirmation., Published as part of Kaczmarek, ��ukasz, 2017, Annotated zoogeography of non-marine Tardigrada. Part IV: Africa, pp. 1-74 in Zootaxa 4284 (1) on page 10, DOI: 10.11646/zootaxa.4284.1.1, http://zenodo.org/record/1010400, {"references":["Richters, F. (1908) Moosbewohner. Wissenschaftliche ERgebnisse deR Schwedischen SudpolaR-Expedition (1901 - 1903), Stockholm, 6, 1 - 16.","Heinis, F. (1928) Beitrag zur Moosfauna des Krakatau. VeRhandlungen deR NatuRfoRschenden Gesellschaft in Basel, 39, 57 - 65.","Iharos, G. (1969 a) Tardigraden aus Mittelwestafrika. Opuscula Zoologica (Budapest), 9, 115 - 120.","Middleton, R. C. (2003) Tardigrades in southern Africa. AfRican JouRnal of Ecology, 41, 280 - 282. https: // doi. org / 10.1046 / j. 1365 - 2028.2003.00439. x","Michalczyk, L. & Kaczmarek, L. (2006 a) Revision of the Echiniscus bigRanulatus group with a description of a new species Echiniscus madonnae (Tardigrada: Heterotardigrada: Echiniscidae) from South America. Zootaxa, 1154, 1 - 26.","Michalczyk, L. & Kaczmarek, L. (2007) Echiniscus ganczaReki, a new species of Tardigrada (Heterotardigrada: Echiniscidae, bigRanulatus group) from Costa Rica. Zootaxa, 1471, 15 - 25."]}

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: U��ge 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., Published as part of Kaczmarek, ��ukasz, 2017, Annotated zoogeography of non-marine Tardigrada. Part IV: Africa, pp. 1-74 in Zootaxa 4284 (1) on page 14, DOI: 10.11646/zootaxa.4284.1.1, http://zenodo.org/record/1010400, {"references":["Thulin, G. (1911) Beitrag zur Kenntnis der Tardigradenfauna Schwedens. ARkiv foR Zoologi, 7, 1 - 60. https: // doi. org / 10.5962 / bhl. part. 1270","Da Cunha, A. X. da & Nascimento Ribeiro, F. (1964) Tardigrados de Angola. GaRcia de ORta, Lisboa, 12, 397 - 406.","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"]}

31. Echiniscus quadrispinosus Richters, 1902 sensu lato [T] Echiniscus quadRispinosus Richters (Murray 1913) Terra typica: Germany (Europe) Kenya: ��� 00��01���S, 37��54���E: Undefined locality, British East Africa [Kenya], moss. Murray (1913) Record numbers. Kenya: 1; total: 1. Remarks. There is considerable morphological variation in the global records of E. quadrispinosus, indicative of a species complex (see: Ramazzotti & Maucci 1983). However, most of the species records are (like the type locality) Holarctic (McInnes 1994), which suggest that African reports may be correct but, these early reports, require confirmation., Published as part of Kaczmarek, ��ukasz, 2017, Annotated zoogeography of non-marine Tardigrada. Part IV: Africa, pp. 1-74 in Zootaxa 4284 (1) on page 15, DOI: 10.11646/zootaxa.4284.1.1, http://zenodo.org/record/1010400, {"references":["Richters, F. (1902) Neue Moosbewohner. BeRicht deR SenckenbeRgischen NatuRfoRschenden gesellschaft in FRankfoRt am Main, 1902, 23 - 26.","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","Ramazzotti, G. & Maucci, W. (1983) Il Phylum Tardigrada. MemoRie dell'Istituto Italiano di IdRobiologia, 41, 1 - 1012.","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"]}

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"]}

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: U��ge Province, near Carmona [U��ge], 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 (W&eogon;glarska 1962)., Published as part of Kaczmarek, ��ukasz, 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":["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","Da Cunha, A. X. da & Nascimento Ribeiro, F. (1964) Tardigrados de Angola. GaRcia de ORta, Lisboa, 12, 397 - 406.","Binda, M. G. & Pilato, G. (1995 a) Some notes on African tardigrades with description of two new species. TRopical Zoology, 8, 367 - 372. https: // doi. org / 10.1080 / 03946975.1995.10539294","Middleton, R. C. (2003) Tardigrades in southern Africa. AfRican JouRnal of Ecology, 41, 280 - 282. https: // doi. org / 10.1046 / j. 1365 - 2028.2003.00439. x","Weglarska, B. (1962) Die Tardigraden Vietnams. Acta Societatis Zoologicae Bohemoslovenicae, 26, 300 - 307."]}

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: Hu��la Province, S�� 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., Published as part of Kaczmarek, ��ukasz, 2017, Annotated zoogeography of non-marine Tardigrada. Part IV: Africa, pp. 1-74 in Zootaxa 4284 (1) on page 10, DOI: 10.11646/zootaxa.4284.1.1, http://zenodo.org/record/1010400, {"references":["Da Cunha, A. X. da & Nascimento Ribeiro, F. (1964) Tardigrados de Angola. GaRcia de ORta, Lisboa, 12, 397 - 406.","Van Rompu, E. A., De Smet, W. H. & Bafort, J. M. (1991 a) Some freshwater tardigrades from the Kilimanjaro. Naturwetenschappelijk Tijdschrift, 73, 55 - 62.","Middleton, R. C. (2003) Tardigrades in southern Africa. AfRican JouRnal of Ecology, 41, 280 - 282. https: // doi. org / 10.1046 / j. 1365 - 2028.2003.00439. x","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"]}