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Species Paramacrobiotus metropolitanus sp. nov. Sugiura, Matsumoto & Kunieda, 2022 (Figs 1–7, Tables 2–3) Paramacrobiotus sp. TYO in: Sugiura et al. (2019), Hara et al. (2021), Sugiura & Matsumoto (2021), and Sugiura et al. (2022). Material examined Forty animals (NMST-Tg-281–310, NHMD-918038–918047, holotype: NMST-Tg-299) and 33 eggs (NMST-Tg-311–333, NHMD-918048–918057) were examined under light microscopes and were deposited in National Museum of Nature and Science (NMST), Tsukuba, Japan, and Natural History Museum (NHMD), Copenhagen, Denmark, respectively. In addition, over 30 animals and eggs were observed with the scanning electron microscope and stored in the laboratory of Keio University. Description Measurements and statistics of animals are given in Table 2. The Holotype is illustrated in Fig. 1A and a paratype observed with SEM in Figs 1B–C. Adult P. metropolitanus sp. nov. specimens whitish under dark-field illumination, the juvenile transparent (data not shown). Eyes absent in live animals. Mouth anteroventral (Figs 2A–C). Buccopharyngeal apparatus the Macrobiotus type, with ten peribuccal lamellae around the mouth (Fig. 2A). Numerous conical teeth present, forming the first band of teeth posterior to the peribuccal lamellae. Conical teeth present at just behind the ring fold, forming the second band, and ridges parallel to the main axis of the buccal tube as the third band (Figs 2A–C). Approximately 20–30 small and sharp ridges on the second band, and two to four larger ridges on the third band (Fig. 2A–C). Three macroplacoids present in the pharynx, the length sequence 2hufelandi type (Y-shaped, Figs 3–4). Smooth cuticle with a cuticular bulge on the inner side of legs I–III (Fig. 3), whereas granulation on the inner surface of leg IV visible under both PCM and SEM (Figs 4A–B). Primary branches with two distinct accessory points (Fig. 4C). A long common tract with a stalk connects the claw to the lunule. Lunules smooth (Fig. 4C). Smooth body cuticle without pores, sculpturing or granulation (Figs 1 and 5). Muscle attachments (apodemes/cribriform areas) clearly visible on the cuticle, with two holes, each with densely packed pores (Fig. 5). Eggs laid freely, yellow yolk seen internally and transparent conical processes on the surface present in PCM (Figs 6A–B). Reticulated pattern with annual rings present on the surface of the processes, with slightly rough tops (Figs 6C–D). Approximately 10 areolae around each process on the surface of the egg (Fig. 6B). Smooth surface of areoles (Fig. 6E). Morphometric data are available in Table 3. All morphometric data are listed in Supporting Materials SM.01. Type locality 35.742983°N, 139.549757°E; 55 m asl: Japan, Honshu mainland, Kanto region, Tokyo, Nishi-Tokyo city, Tozenji temple; urban cemetery; bamboo leaf litter. The locality described in Sugiura et al. (2019) was corrected in Hara et al. (2021). Etymology The species name of “ metropolitanus ” refers to the Tokyo Metropolitan Area (Tokyo Cho-mei mushi in Japanese), the region where the species was collected. Sequence data We confirmed 100% matches between the sequence of PCR-amplicon and the sequence retrieved from the published genome of the species for 18S rDNA, 28S rDNA and ITS-2 sequences. As for COI sequence, one mismatched base was found between the sequence of 658 bp PCR-amplicon and the sequence retrieved from the genome. The sequences were deposited in GenBank with IDs: LC637243 (18S rDNA), LC649795 (28S rDNA of the PCR amplicon using primer-set of 28S_Eutar_F and 28SR0090), LC649797 (28S rDNA of the PCR amplicon using primer-set of 28SF0001 and 28SR0990), LC649794 (ITS-2), LC637242 (COI sequence deduced from the genome), and LC649796 (COI sequence of PCR-amplicon). Phenotypic differential diagnosis The new species P. metropolitanus sp. nov. with microplacoid, smooth lunules, and its egg without pored surface on the areolae is similar to the following seven species but differs specifically from (data from original descriptions of the compared species): • P. danielae Pilato, Binda, Napolitano & Moncada 2001, known from Ecuador and Peru, by the lack of eyes, and the lack of body granulation, larger body length (200–390 μm in P. danielae vs 393–711 μm in the new species). • P. experimentalis, known only from Madagascar, by the lack of wrinkled surface on the egg areolae, the presence of cuticular bulge on inner surface of claws I–III and granulation on inner surface of leg IV. • P. garynahi Kaczmarek, Michalczyk & Diduszko, 2005, known only from Russia, by the lack of oval pores on the cuticle, the lack of granulation in the first three pairs of legs, the lack of a cap-like structure on the top of the egg process, smaller egg bare diameter (96.0–132.0 μm in P. garynahi vs 58.1–90.1 μm in the new species), and by a smaller egg diameter with processes (142.0–180.0 μm in P. garynahi vs 81.7–112.1 μm in the new species). • P. hapukuensis Pilato, Bind & Lisi, 2006, known only from New Zealand, by the lack of finger-shaped apices of egg processes, and the presence of leg granulation. • P. peteri Pilato, Claxton & Binda, 1989, known only from Australia and Indonesia (Pilato et al. 1989; Pilato and Binda, 1990; Mackness, 2003), by the lack of a cuticular thickening near the lunules I–III, the lack of subdivided apices of egg processes, a lower number of processes on the egg circumference (16–18 in P. peteri vs 10–15 in the new species), and by a higher number of areolae around egg processes (6–7 in P. peteri, approximately 10 in the new species). • P. rioplatensis Claps & Rossi, 1997, known only from Uruguay, by the lack of eyespots, a larger body (227– 372 μm in P. rioplatensis vs 392.5–710.8 μm in the new species), a longer buccal tube (38 μm in P. rioplatensis, over 40–58.4 μm in the new species), a larger egg full diameter (68–79 μm in P. rioplatensis vs 81.7–112.1 μm in the new species), the lack of the a filament-shaped apex of egg processes, higher egg processes (4.6 μm height in P. rioplatensis vs over 7.4 μm in the new species), and by a lower number of processes on the egg circumference (17–19 in P. rioplatensis vs 10–15 in the new species). • P. savai Binda & Pilato, 2001, known from only Sri Lanka, by the lack of eyespots, the lack of a cuticular thickening near the lunules of the first three legs, a different egg process shapes (blunt in P. savai vs conical in the new species), and by a slightly smaller egg full diameter (115 μm in P. savai vs 81.7–112.1 μm in the new species). Genetic differential diagnosis Below is a summary of genetic p-distances, whereas detailed matrices are available in Supporting Material SM.02. • 18S rDNA: 0.5%–3.6%, with the most similar species being P. experimentalis from Madagascar (MN073467 –8 and MH664938) and the least similar being P. areolatus NO. 385 from Svalbard (MH664931). • 28S rDNA: 0.98–7.5%, with the most similar species being P.experimentalis (MN073465 –6and MH664956) and the least being Paramacrobiotus tonollii Ramazzotti, 1956 strain US.035 from USA (MH664963). • ITS-2: 9.9–33.9%, with the most similar species being P. experimentalis (MH073463 –4 and MH666087) and the least similar species being P. tonollii (GQ403679, MH666096). • COI: 21.2–32.6%, with the most similar species being Paramacrobiotus sp. strain TZ.018 from Tanzania (MH676017) and with the least similar species being Paramacrobiotus depressus Guidetti, Cesari, Bertolani, Altiero & Rebecchi, 2019 from Italy (MK041013). Phylogeny The supported bootstrap values of ML trees were usually low (P. kenianus, P. experimentalis, and an undescribed species from Tanzania) and the Neotropics (two unidentified species from Brazil). Genetic species delineation Both, the ITS-2 and the COI BI trees indicated that the cluster of P. metropolitanus sp. nov. is a separate species with>93% probabilities by both maximum likelihood and Bayesian supported solutions. (A) SEM image of the postero-dorsal surface of P. metropolitanus sp. nov. (B) Magnified image of dorsal cuticle apodemes corresponding to the white dashed box in A. (C) Magnified image of the left apodemes in B. White-filled indented arrowheads indicate the apodemes. Scale bars = μm.

Species Paramacrobiotus metropolitanus sp. nov. Sugiura, Matsumoto & Kunieda, 2022 (Figs 1–7, Tables 2–3) Paramacrobiotus sp. TYO in: Sugiura et al. (2019), Hara et al. (2021), Sugiura & Matsumoto (2021), and Sugiura et al. (2022). Material examined Forty animals (NMST-Tg-281–310, NHMD-918038–918047, holotype: NMST-Tg-299) and 33 eggs (NMST-Tg-311–333, NHMD-918048–918057) were examined under light microscopes and were deposited in National Museum of Nature and Science (NMST), Tsukuba, Japan, and Natural History Museum (NHMD), Copenhagen, Denmark, respectively. In addition, over 30 animals and eggs were observed with the scanning electron microscope and stored in the laboratory of Keio University. Description Measurements and statistics of animals are given in Table 2. The Holotype is illustrated in Fig. 1A and a paratype observed with SEM in Figs 1B–C. Adult P. metropolitanus sp. nov. specimens whitish under dark-field illumination, the juvenile transparent (data not shown). Eyes absent in live animals. Mouth anteroventral (Figs 2A–C). Buccopharyngeal apparatus the Macrobiotus type, with ten peribuccal lamellae around the mouth (Fig. 2A). Numerous conical teeth present, forming the first band of teeth posterior to the peribuccal lamellae. Conical teeth present at just behind the ring fold, forming the second band, and ridges parallel to the main axis of the buccal tube as the third band (Figs 2A–C). Approximately 20–30 small and sharp ridges on the second band, and two to four larger ridges on the third band (Fig. 2A–C). Three macroplacoids present in the pharynx, the length sequence 2hufelandi type (Y-shaped, Figs 3–4). Smooth cuticle with a cuticular bulge on the inner side of legs I–III (Fig. 3), whereas granulation on the inner surface of leg IV visible under both PCM and SEM (Figs 4A–B). Primary branches with two distinct accessory points (Fig. 4C). A long common tract with a stalk connects the claw to the lunule. Lunules smooth (Fig. 4C). Smooth body cuticle without pores, sculpturing or granulation (Figs 1 and 5). Muscle attachments (apodemes/cribriform areas) clearly visible on the cuticle, with two holes, each with densely packed pores (Fig. 5). Eggs laid freely, yellow yolk seen internally and transparent conical processes on the surface present in PCM (Figs 6A–B). Reticulated pattern with annual rings present on the surface of the processes, with slightly rough tops (Figs 6C–D). Approximately 10 areolae around each process on the surface of the egg (Fig. 6B). Smooth surface of areoles (Fig. 6E). Morphometric data are available in Table 3. All morphometric data are listed in Supporting Materials SM.01. Type locality 35.742983°N, 139.549757°E; 55 m asl: Japan, Honshu mainland, Kanto region, Tokyo, Nishi-Tokyo city, Tozenji temple; urban cemetery; bamboo leaf litter. The locality described in Sugiura et al. (2019) was corrected in Hara et al. (2021). Etymology The species name of “ metropolitanus ” refers to the Tokyo Metropolitan Area (Tokyo Cho-mei mushi in Japanese), the region where the species was collected. Sequence data We confirmed 100% matches between the sequence of PCR-amplicon and the sequence retrieved from the published genome of the species for 18S rDNA, 28S rDNA and ITS-2 sequences. As for COI sequence, one mismatched base was found between the sequence of 658 bp PCR-amplicon and the sequence retrieved from the genome. The sequences were deposited in GenBank with IDs: LC637243 (18S rDNA), LC649795 (28S rDNA of the PCR amplicon using primer-set of 28S_Eutar_F and 28SR0090), LC649797 (28S rDNA of the PCR amplicon using primer-set of 28SF0001 and 28SR0990), LC649794 (ITS-2), LC637242 (COI sequence deduced from the genome), and LC649796 (COI sequence of PCR-amplicon). Phenotypic differential diagnosis The new species P. metropolitanus sp. nov. with microplacoid, smooth lunules, and its egg without pored surface on the areolae is similar to the following seven species but differs specifically from (data from original descriptions of the compared species): • P. danielae Pilato, Binda, Napolitano & Moncada 2001, known from Ecuador and Peru, by the lack of eyes, and the lack of body granulation, larger body length (200–390 μm in P. danielae vs 393–711 μm in the new species). • P. experimentalis, known only from Madagascar, by the lack of wrinkled surface on the egg areolae, the presence of cuticular bulge on inner surface of claws I–III and granulation on inner surface of leg IV. • P. garynahi Kaczmarek, Michalczyk & Diduszko, 2005, known only from Russia, by the lack of oval pores on the cuticle, the lack of granulation in the first three pairs of legs, the lack of a cap-like structure on the top of the egg process, smaller egg bare diameter (96.0–132.0 μm in P. garynahi vs 58.1–90.1 μm in the new species), and by a smaller egg diameter with processes (142.0–180.0 μm in P. garynahi vs 81.7–112.1 μm in the new species). • P. hapukuensis Pilato, Bind & Lisi, 2006, known only from New Zealand, by the lack of finger-shaped apices of egg processes, and the presence of leg granulation. • P. peteri Pilato, Claxton & Binda, 1989, known only from Australia and Indonesia (Pilato et al. 1989; Pilato and Binda, 1990; Mackness, 2003), by the lack of a cuticular thickening near the lunules I–III, the lack of subdivided apices of egg processes, a lower number of processes on the egg circumference (16–18 in P. peteri vs 10–15 in the new species), and by a higher number of areolae around egg processes (6–7 in P. peteri, approximately 10 in the new species). • P. rioplatensis Claps & Rossi, 1997, known only from Uruguay, by the lack of eyespots, a larger body (227– 372 μm in P. rioplatensis vs 392.5–710.8 μm in the new species), a longer buccal tube (38 μm in P. rioplatensis, over 40–58.4 μm in the new species), a larger egg full diameter (68–79 μm in P. rioplatensis vs 81.7–112.1 μm in the new species), the lack of the a filament-shaped apex of egg processes, higher egg processes (4.6 μm height in P. rioplatensis vs over 7.4 μm in the new species), and by a lower number of processes on the egg circumference (17–19 in P. rioplatensis vs 10–15 in the new species). • P. savai Binda & Pilato, 2001, known from only Sri Lanka, by the lack of eyespots, the lack of a cuticular thickening near the lunules of the first three legs, a different egg process shapes (blunt in P. savai vs conical in the new species), and by a slightly smaller egg full diameter (115 μm in P. savai vs 81.7–112.1 μm in the new species). Genetic differential diagnosis Below is a summary of genetic p-distances, whereas detailed matrices are available in Supporting Material SM.02. • 18S rDNA: 0.5%–3.6%, with the most similar species being P. experimentalis from Madagascar (MN073467 –8 and MH664938) and the least similar being P. areolatus NO. 385 from Svalbard (MH664931). • 28S rDNA: 0.98–7.5%, with the most similar species being P.experimentalis (MN073465 –6and MH664956) and the least being Paramacrobiotus tonollii Ramazzotti, 1956 strain US.035 from USA (MH664963). • ITS-2: 9.9–33.9%, with the most similar species being P. experimentalis (MH073463 –4 and MH666087) and the least similar species being P. tonollii (GQ403679, MH666096). • COI: 21.2–32.6%, with the most similar species being Paramacrobiotus sp. strain TZ.018 from Tanzania (MH676017) and with the least similar species being Paramacrobiotus depressus Guidetti, Cesari, Bertolani, Altiero & Rebecchi, 2019 from Italy (MK041013). Phylogeny The supported bootstrap values of ML trees were usually low (P. kenianus, P. experimentalis, and an undescribed species from Tanzania) and the Neotropics (two unidentified species from Brazil). Genetic species delineation Both, the ITS-2 and the COI BI trees indicated that the cluster of P. metropolitanus sp. nov. is a separate species with>93% probabilities by both maximum likelihood and Bayesian supported solutions. (A) SEM image of the postero-dorsal surface of P. metropolitanus sp. nov. (B) Magnified image of dorsal cuticle apodemes corresponding to the white dashed box in A. (C) Magnified image of the left apodemes in B. White-filled indented arrowheads indicate the apodemes. Scale bars = μm., Published as part of Sugiura, Kenta, Matsumoto, Midori & Kunieda, Takekazu, 2022, Description of a model tardigrade Paramacrobiotus metropolitanus sp. nov (Eutardigrada) from Japan with a summary of its life history, reproduction and genomics, pp. 92-112 in Zootaxa 5134 (1) on pages 96-102, DOI: 10.11646/zootaxa.5134.1.4, http://zenodo.org/record/6531340, {"references":["Sugiura, K., Minato, H., Suzuki, A. C., Arakawa, K., Kunieda, T. & Matsumoto, M. (2019) Comparison of sexual reproductive behaviors in two species of Macrobiotidae (Tardigrada: Eutardigrada). Zoological Science, 36 (2), 120 - 127. https: // doi. org / 10.2108 / zs 180103","Hara, Y., Shibahara, R., Kondo, K., Abe, W. & Kunieda T. (2021) Parallel evolution of trehalose production machinery in anhydrobiotic animals via recurrent gene loss and horizontal transfer. Open Biology, 11, 7. https: // doi. org / 10.1098 / rsob. 200413","Sugiura, K. & Matsumoto, M. (2021) Spermatozoa morphology changes during reproduction and first observation of acrosomal contact in two dioecious species of Macrobiotidae (Tardigrada: Eutardigrada). Zygote, 29, 42 - 48. https: // doi. org / 10.1017 / S 0967199420000490","Sugiura, K., Shiba, K., Inaba, K. & Matsumoto, M. (2022) Morphological differences in tardigrade spermatozoa induce variation in gamete motility. BMC Zoology, 7, 8. https: // doi. org / 10.1186 / s 40850 - 022 - 00109 - w","Pilato, G., Binda, M. G., Napolitano, A. & Moncada E. (2001) Notes on South American tardigrades with the description of two new species: Pseudechiniscus spinerectus and Macrobiotus danielae. Tropical Zoology, 14, 2, 223 - 231. https: // doi. org / 10.1080 / 03946975.2001.10531154","Kaczmarek. L., Michalczyk, L. & Diduszko, D. (2005) Some tardigrades from Siberia (Russia, Baikal region) with a description of Macrobiotus garynahi sp. nov. (Eutardigrada: Macrobiotidae: richtersi group). Zootaxa, 1053, 35 - 45. http: // doi. org / 10.11646 / zootaxa. 1053.1.3","Pilato, G., Claxton, S. & Binda, M. G. (1989) Tardigrades from Australia. III. Echiniscus marcusi and Macrobiotus peteri, new species of tardigrades from New South Wales. Animalia, 16, 43 - 48.","Pilato, G. & Binda M. G. (1990) Notizie sui tardigradi muscicoli di Bali (Indonesia). Animalia, 17, 209 - 218.","Mackness, B. S. (2003) A review of nematophagy by terrestrial tardigrades with new observations on some Australian species. Proceedings of the Royal Society of Victoria, 115, 1 - 5.","Claps, M. C. & Rossi, G. C. (1997) Tardigrados de Uruguay, com descripcion de dos nuevas especies (Echiniscidae, Macrobiotidae). Iheringia Serie Zoologia, 83, 17 - 22.","Guidetti, R., Cesari, M., Bertolani, R., Altiero, T. & Rebecchi, L. (2019) High diversity in species, reproductive modes and distribution within the Paramacrobiotus richtersi complex (Eutardigrada, Macrobiotidae). Zoological letters, 5 (1), 1 - 28. https: // doi. org / 10.1186 / s 40851 - 018 - 0113 - z"]}

SummaryMating behaviours for two species of dioecious eutardigrades: a strain of Paramacrobiotus sp. and Macrobiotus shonaicus (Stec et al., 2018) have been recorded previously, and observations have indicated that spermatozoa of both species are first released into the environment, then swim through the cloaca of the females and into the spermatheca. The fusion of gamete nuclei has not yet occurred in a laid egg. Therefore, it has been suggested that fertilization is completed externally as the egg is released into the environment before the nuclei of the gametes fuse. In the present study, the spermatozoa of both Paramacrobiotus sp. and M. shonaicus spermatozoa underwent morphological changes during reproduction. In morphometrical analyses of testicular spermatozoa, the tail, mid-piece, nucleus, and acrosome were significantly longer in Paramacrobiotus sp. compared with M. shonaicus. The nuclei of both the testicular and spermathecal spermatozoa were equally coiled, but the latter had shorter tails in both species. These spermatozoa were present on the surface of the egg chorion after oviposition. The tip of the acrosomes lay buried in the chorion, suggesting that penetration had occurred. We also proposed that the reduced tail is a conserved trait, at least in Macrobiotidae.

Phylum Tardigrada, which has been quite a little studied both around the world and Turkey, belongs to members of a metazoan. In this study eight moss and lichen samples were collected and investigated from a variety of elevations of the Verçenik Mountain in Rize (Turkey). In total 62 specimens, 11 exivium and 10 eggs of tardigrades were found. Five species were identified, and they belong to genera: Acanthechiniscus, Pseudechiniscus, Macrobiotus, Paramacrobiotus and, Ramazzottius. Three of them are new records for Turkish fauna i.e. Acanthechiniscus victor (Ehrenberg, 1853), Pseudechiniscus ramazzottii ramazzottii Maucci, 1952 and Macrobiotus spectabilis Thulin, 1928. Including these three species, the tardigrade fauna of Turkey increased to 54. In this paper, we present the previous study a full list of the known high mountain and alpine tardigrade species with their localities. All specimens are deposited in the Aquatic Animals Research Laboratory at Ankara University.

Incomplete descriptions of nominal taxa are one of the most significant obstacles in modern taxonomy, including the taxonomy of Tardigrada. Another major problem in tardigrade systematics is the lack of tests for the reliability of genetic markers in species delineation. Here, we employ an integrative taxonomy approach to redescribe the nominal taxon for the P. areolatus complex, Paramacrobiotus areolatus. Moreover, we obtained multilocus DNA sequences for another 16 populations representing 9–12 Paramacrobiotus species collected from Europe, North America, Africa and Australia, enabling us to reconstruct the most extensive phylogeny of the genus to date. The identification of a pair of potentially cryptic dioecious P. areolatus complex species with divergent genetic distances in ITS2 (1.4%) and COI (13.8%) provided an opportunity to test the biological species concept for the first time in the history of tardigrade taxonomy. Intra- and interpopulation crosses did not differ in reproductive success in terms of F1 offspring. However, because of the low F1 family sizes, we were unfortunately unable to test F1 hybrid fertility. Although our results are only partially conclusive, they offer a baseline not only for further taxonomic and phylogenetic research on the areolatus complex, but also for studies on species delineation in tardigrades in general.

Based on the original species descriptions, a review of the genus Paramacrobiotus was conducted. We divided the genus into two subgenera, Microplacoidus subgen. nov. and Paramacrobiotus subgen. nov., based on the presence or absence of a microplacoid, and characterized species within the genus based on seven different types of eggs. In a moss sample collected in Ecuador, Paramacrobiotus (Paramacrobiotus) spinosus sp. nov., was found. The new species differs from all species of the subgenus Parama crobiotus by the presence of richtersi type eggs and from other species by morphometric characters. Additionally, in the Ecuadorian material we found P. (Microplacoidus) magdalenae comb. nov., which is the first record of this species in Ecuador, and we provide the full set of measurements for this species, not included in the original description. An additional new record is P. (M.) alekseevi comb. nov. found in Vietnam for the first time. After examining microscope slides from the Iharos' collection deposited in the Hungarian Natural History Museum, we prepared re-descriptions of P. (P.) csotiensis comb nov., P. (M.) submorulatus comb. nov. and P. (M.) wauensis comb. nov. Based on the morphological and morphometric characters of adults and eggs, we developed a diagnostic key to the genus Paramacrobiotus.

Naturally occurring fluorescence has been observed in multiple species ranging from bacteria to birds. In macroscopic animals such as birds and fishes, fluorescence provides a visual communication signal. However, the functional significance of this phenomenon is not known in most cases. Though photoprotection is attributed to fluorescence under ultraviolet (UV) light in some organisms, it lacks direct experimental evidence. Here, we have identified a new species of eutardigrade belonging to the genus Paramacrobiotus, which exhibits fluorescence under UV light. Using a natural variant of the same species that lacks fluorescence, we show that the fluorescence confers tolerance to lethal UV radiation. Remarkably, we could transfer this property to UV-sensitive Hypsibius exemplaris, another eutardigrade, and also to C. elegans, a nematode. Using high performance liquid chromatography (HPLC) we isolated the fluorescent compound from Paramacrobiotus sp. This compound has excitation maxima (λex) at 370 nm and emission maxima (λem) at 420-430 nm. We propose that Paramacrobiotus sp. uses a fluorescent shield that absorbs harmful UV radiation, and emits harmless blue light, thereby protecting itself from the lethal effects of UV radiation.Summary statementTardigrades are well known for their tolerance to extreme environmental conditions. In this study, we have identified a new tardigrade species that employs a fluorescent shield to protect itself from the germicidal ultra violet radiation.

In this paper, Macrobiotus naskreckii sp. nov., a new species of the hufelandi group from the Gorongosa National Park in Mozambique, is described. The analysis have revealed that M. naskreckii sp. nov. is most similar to M. kristenseni, M. ramoli and M. serratus. The new species differs from M. kristenseni mainly through a different shape of egg processes and clearly dentate lunules IV; from M. ramoli mainly by smaller eggs and a smaller number of egg processes on the circumference; from M. serratus mainly by the absence of the reticulation between the egg processes. This is a third record of tardigrades from Mozambique and apart of the new species reported here, three other taxa from Gorongosa National Park are provided: Minibiotus cf. intermedius, Minibiotus sp. and Paramacrobiotus cf. richtersi.

Naturally occurring fluorescence has been observed in multiple species ranging from bacteria to birds. In macroscopic animals such as birds, fluorescence provides a visual communication signal. However, the functional significance of this phenomenon is unknown in most cases. Though photoprotection is attributed to fluorescence under ultraviolet (UV) light in some organisms, it lacks direct experimental evidence. Here, we demonstrate naturally occurring fluorescence under UV light in a eutardigrade belonging to the genus Paramacrobiotus . Using a natural variant that lacks fluorescence, we show that the fluorescence confers tolerance to lethal UV radiation. Remarkably, the fluorescent extract from Paramacrobiotus sp. could protect the UV-sensitive tardigrade Hypsibius exemplaris and nematode Caenorhabditis elegans from germicidal UV radiation. We propose that Paramacrobiotus sp. possess a protective fluorescent shield that absorbs harmful UV radiation and emits harmless blue light.

Tardigrada (water bears) are microscopic invertebrates inhabiting aquatic (freshwater and marine) and terrestrial habitats. They are thriving in almost all Earth ecosystems from deepest oceans to highest mountains, from tropics to polar regions. Water bears are probably most famous for their cryptobiotic abilities, which allow them to survive a broad spectrum of extreme environmental conditions. The Special Issue on tardigrades was launched to popularize research on these fascinating microinvertebrates. The published papers were focused on (a) marine and terrestrial tardigrades diversity, (b) interpopulation variability of Antarctic eutardigrade Paramacrobiotus fairbanksi, (c) encystment in freshwater eutardigrade Thulinius ruffoi and (d) use of a metabarcoding approach to community structures studies in microenvironments.

The recent re-description of Paramacrobiotus Guidetti, Schill, Bertolani, Dandekar and Wolf, 2009 has inadvertently led to the description of an objective synonym within its subgenera nominal taxa. To resolve this issue, we have re-described both subgenera, and proposed a new substitute name for one subgenus, in line with the International Code of Zoological Nomenclature. Additionally we have confirmed the placement of two recently published Paramacrobiotus species, not included in the last revision, within the respective subgenera established herein.

The water bears (Tardigrada) from Colombia are rather poorly known; thus, the aim of this research was to study the composition of tardigrade fauna inhabiting epiphytic and rock mosses in a high Andean forest in Guatavita. A 25-m2 plot was delimited and 6 randomly chosen trees with epiphytic mosses were sampled. Twelve samples of epiphytic mosses and 6 of rock mosses were collected and examined. A total of 277 specimens and 6 eggs of 8 tardigrade taxa were found in these samples. All taxa are new records for Guatavita and 5-Hypsibius dujardini (Doyère, 1840); Milnesium katarzynae Kaczmarek et al., 2004; Milnesium krzysztofi Kaczmarek & Michalczyk, 2007; Milnesium granulatum Ramazzotti 1962; and Paramacrobiotus danielae (Pilato et al., 2001)-are also new for Colombia.

Paramacrobiotus sagani sp. nov. (Figs. 2���3, Tables 2���3) Material examined: Holotype mounted in Hoyer���s medium, 11 paratypes and 10 eggs in Hoyer���s medium, and 15 paratypes in PVA, from locality Bonda, lower basin of the ManZanares River. One additional specimen and one additional egg from the locality Puerto Mosquito, lower basin of the Gaira River, were also observed (see: Table 1). In general, the structure of the microhabitat from which all the specimens and eggs were collected were either a mixed assemblage or single species of bryophytes and/or lichens growing on tree trunks from the following families: Bryophytes (Amblystegiaceae, Dicranaceae, Lejeuneaceae, Fissidentaceae, Frullaniaceae, Meteoriaceae, Racopilaceae, and Stereophyllaceae), and lichens (Strigulaceae, Acarosporaceae, and Thelotremataceae). Holotype found in single species of Stereophyllaceae growing on tree. Type repositories: The holotype, paratypes and eggs are deposited in the Centro de Colecciones Biol��gicas de la Universidad del Magdalena (CBUMAG), Santa Marta, Colombia. Holotype CBUMAG:TAR:00004-4, 26 paratypes and 10 eggs (Table 1). Species diagnosis: Body milky white in living specimens. Eye-spots present, difficult to see after mounting. Cuticle with fine granulation (Fig. 2E���F), also present on the ventral surface, but with sparser granules. Granulation very evident in the smallest specimens, becoming more faint with maturity, and practically invisible in specimens larger than ca. 450 ��m. Bucco-pharyngeal apparatus (Fig. 2B), including buccal armature, of Macrobiotus type (Pilato & Binda 2010), with buccal armature and placoid configuration as in P. richtersi: three slender rod-shaped macroplacoids, not forming an arc (Maucci 1973), microplacoid distant from the third macroplacoid (Pilato 1975); buccal armature as described by Pilato (1972). Stylet supports inserted at 77.1���81.5% of the buccal tube length. Claws of Macrobiotus type, slender, with long basal portion, well-developed accessory points on primary branches, and smooth lunules at the base of all legs (Fig. 2C���D). Cuticular bars on legs I���III present but poorly developed (Fig. 2C, arrow). Eggs white, spherical, areolated, with 10���13 processes round circumference, and 19���24 in the hemisphere (Fig. 3A���B). Trunco-conical processes, usually with short cylindrical indented apices, which are often enlarged at the very top (Fig. 3A���C). Processes showing in light microscopy the typical mesh, which also partially continues on the ridges delimiting the areolae (Fig. 3C). Areolae, 9���12 per process, clearly sculptured (Fig. 3D). Description of the holotype: Body length 446 ��m (Fig. 2A), milky white in vivo. Eye-spots present and still visible after mounting (N.B. in some other specimens they have disappeared). Cuticle with fine granulation, also present ventrally but with sparser granules, more evident on the legs (Fig. 2E���F), especially the hind legs. Buccopharyngeal apparatus of Macrobiotus type (i.e. with ten peribuccal lamellae, rigid buccal tube with ventral lamina which lacks ventral hook ��� Pilato & Binda 2010), with buccal armature which comprises three bands of teeth (Fig. 2B). The first band, close to the oral aperture, is composed of numerous, small, granular teeth disposed irregularly in many rows. The second band, right above the third, is composed of larger triangular teeth arranged in a single row. The third band is formed by three dorsal and three ventral ridges. The medio-ventral ridge is divided in two teeth. Buccal tube 55.4 ��m long and 14.1 ��m wide (pt = 25.5). The stylet support insertion point is placed at 78.9% of the buccal tube length. Apophyses and three rod-shaped macroplacoids present in the pharynx; microplacoid present, small and placed at a distance from the third macroplacoid. First macroplacoid, with a central constriction, 9.5 ��m long (pt = 17.0); second macroplacoid, 7.6 ��m (pt = 13.7); third macroplacoid, 11.4 ��m (pt = 20.5); microplacoid, 4.1 ��m long (pt = 7.4); entire placoid row, 39.9 ��m long (pt = 72.0); macroplacoid row, 29.3 ��m (pt = 52.8). Claws of Macrobiotus type, slender, with a long basal portion (Fig. 2C���D). Well-developed accessory points on primary branches. First pair of legs, primary branches of external claw, 11.1 ��m long (pt = 20.1), internal claws incorrectly aligned for measurement. Second pair of legs, external claw primary branches, 11.4 ��m long (pt = 20.6); primary branches of internal claw, 11.6 ��m long (pt = 20.9). Third pair of legs, external claw primary branches, 11.5 ��m long (pt = 20.7); internal claw primary branches, 11.5 ��m long (pt = 20. 7). Anterior claw primary branches of the hind legs, 13.0 ��m long (pt = 23.4); posterior claw primary branches, 13.7 ��m long (pt = 24.7). Smooth lunules present on all claw bases. Cuticular bars typical of the richtersi group (one under each claw parallel to the lunule) present on legs I���III, but poorly developed (Fig. 2C, arrow). Eggs white, spherical, areolated, with 10���13 processes round the circumference, and 19���24 in the hemisphere (Fig. 3A���B). Egg diameter 73.7���87.7 ��m excluding the processes, 92.5���109.7 ��m including. Trunco-conical processes usually with cylindrical, indented apices, variable in length but never very long, and often enlarged at the very top (processes 9.4���13.2 ��m in height, and 14.6���22.4 ��m in the basal diameter) (Fig. 3A���C); the cylindrical apices appearance and length seem to depend on the degree of distension of the processes. Processes, and also the proximal portions of the ridges delimiting the areolae showing in light microscopy the typical mesh, variable in shape, some elongated and sinuous, giving an irregular reticulated appearance (Fig. 3C). A slight, transverse annulation is visible on most processes, but this might depend on the possibility that the egg had been very recently laid. Areolae (9���12 per process) with thickened, sculptured central portion which appears dark, and with small pores appearing as bright dots (Fig. 3D, arrows). Quantitative characters of the eggs measured are given in Table 3. Sample Locality Geographic coordinates Altitude Tardigrade species Specimens Slide code CBUMAG: TAR (m a.s.l.) 1 MAN 11��14'03''N 79��06'54''W 70 Paramacrobiotus sagani sp. nov. -type 1 0 0 0 0 1 1 0 0 202 Sample Locality Geographic coordinates Altitude Tardigrade species Specimens Slide code CBUMAG: TAR (m. a.s.l.) 21 MAN 11��13'40.9"N 74��06'41.1"W 82 Paramacrobiotus sagani sp. nov. - type 1 0 0 229 1 0 0 232 1 0 0 234 1 0 0 235 22 MAN 11��13'41.1"N 74��06'41.1"W 82 Paramacrobiotus sagani sp. nov. - type 1 0 0 244 1 0 0 245 2 0 0 246 1 0 0 248 Doryphoribius rosanae sp. nov. - type 1 0 0 246 1 0 0 248 1 0 0 249 23 MAN 11��13'41.1"N 74��06'41.1"W 82 Doryphoribius rosanae sp. nov. - type 1 0 0 250 Paramacrobiotus sagani sp. nov. - type 1 0 0 251 24 GAI 11��10'21.8"N 74��10'33.1"W 49 Doryphoribius rosanae sp. nov. -add. mat. 1 0 0 278 25 GAI 11��10'15.6"N 74��10'27.2"W 60 Doryphoribius rosanae sp. nov. -add. mat. 9 0 0 282 0 0 283 0 0 285 0 0 286 0 0 288 0 0 289 0 0 291 0 0 293 Egg Paramacrobiotus sagani sp. nov. -add. mat. 1 0 0 294 CHARACTER N RANGE MEAN SD Holotype ��m pt ��m pt ��m pt ��m pt Body length 27 189 ��� 603 ��� 402 81 446 Buccopharyngeal tube Taxonomic remarks: The paratypes show the same characters of the holotype (Table 2), apart from the following details: the cuticle granulation is very evident in the smallest specimens and gradually becomes fainter with maturity, and is practically invisible, except on the legs, in the larger specimens over 450 ��m in length. Also, the medio-ventral ridge of the buccal armature, can be divided in two to five teeth (most frequently three); only in one case were these smaller and more numerous (7���8) teeth. Our data for the pt index of the buccal tube width in proportion to the body length seems to indicate allometric growing: with the pt index varying from about 19 in the smallest specimens (about 200 ��m) to almost 30 in the largest specimens (about 600 ��m) (Fig. 4). Etymology: This species is named after the astrophysicist and science communicator Carl Sagan, philosopher, and one of the greatest minds of the twentieth century. Differential diagnosis: The presence of microplacoids in the pharynx, and some details of the eggs (especially type of areolation and type of mesh on the processes), indicates Paramacrobiotus sagani sp. nov. belongs to the richtersi group, and the presence of a characteristic sculpture in the areolae places it in the subgroup with vanescens - type areolation (central portion of the areolae thickened and sculptured ���MichalcZyk & KacZmarek 2006). Other species in this subgroup are: P. alekseevi (Tumanov, 2005), P. corgatensis (Pilato, Binda & Lisi, 2002), P. danielisae (Pilato, Binda & Lisi, 2006), P. gerlachae (Pilato, Binda & Lisi, 2004), P. halei (Bartels, Pilato, Lisi & Nelson, 2009), P. lorenae (Biserov, 1996), P. magdalenae (MichalcZyk & KacZmarek, 2006), P. pius Lisi, Binda & Pilato, 2016, P. priviterae (Binda, Pilato, Moncada & Napolitano, 2001), P. sklodowskae (MichalcZyk, KacZmarek & Węglarska, 2006) and P. vanescens (Pilato, Binda & CatanZaro, 1991). Paramacrobiotus sagani sp. nov. differs from P. alekseevi by having cuticular granulation (only on the legs in P. alekseevi at all stages of maturity), generally wider buccal tube (pt 15.0��� 20.7 in P. alekseevi, 19.0��� 29.7 in P. sagani sp. nov.). Claws with more evident accessory points and smooth lunules on all legs (with indented margin on legs IV in P. alekseevi). Larger eggs (59.2���82.8 ��m excluding the processes in P. alekseevi, 73.7���87.7 ��m in P. sagani sp. nov.), with shorter processes (11.8���21.8 ��m in P. alekseevi, 9.4���13.2 ��m in P. sagani sp. nov.), with processes having cylindrical and indented apices (vesicular, cap-like apices in P. alekseevi), and areolae with evidently sculptured central portion (only faintly sculptured in P. alekseevi). The cuticular granulation of P. sagani sp. nov. is most similar to P. corgatensis, from which the new species differs by having smooth lunules on all legs (with teeth on legs IV in P. corgatensis). Eggs with shorter processes (about 20���25 ��m in P. corgatensis, 9.4���13.2 ��m in P. sagani sp. nov.), and with slightly different average number of processes on the egg circumference (8���11 in P. corgatensis, 10���13 in P. sagani sp. nov.), with cylindrical apices (tapering apices in P. corgatensis). Paramacrobiotus sagani sp. nov. differs from P. danielisae by having eye-spots, fine cuticular granulation (irregular, polygonal, flat tubercles in P. danielisae). Eggs with 10���13 trunco-conical processes on circumference (9���10 conical processes in P. danielisae), and with different apices (more or less tapering or stout, not regularly cylindrical and indented in P. danielisae). Paramacrobiotus sagani sp. nov. differs from P. gerlachae by having eye-spots, granulation on the cuticle (only on the hind legs in P. gerlachae at all stages of maturity). Wider buccal tube (pt 16.4���19.4 in P. gerlachae, 19.0��� 29.7 in P. sagani sp. nov.). Eggs with cylindrical, indented process apices (smooth, rounded or truncated apices in P. gerlachae) and thickened, clearly sculptured areolae (very poorly visible sculpture in P. gerlachae). Paramacrobiotus sagani sp. nov. differs from P. halei by having eye-spots, granular sculpture on the cuticle (tubercles in P. halei). Smaller eggs (91.0���96.0 ��m in P. halei, 73.7���87.7 ��m in P. sagani sp. nov. excluding processes), and with different processes apices (smooth, truncated in P. halei). Paramacrobiotus sagani sp. nov. differs from P. lorenae by having eye-spots, less caudal stylet support insertion point (pt 84.5 in the holotype in P. lorenae, 78.9 in the holotype in P. sagani sp. nov.), and eggs with differently shaped processes (conical processes with very narrow, elongated apices in P. lorenae). Paramacrobiotus sagani sp. nov. differs from P. magdalenae by having more rounded eye-spots (elongated eye-spots in P. magdalenae). Cuticular granulation (only on the hind legs in P. magdalenae at all stages of maturity). Generally, wider buccal tube (pt 16.1���22.2 in P. magdalenae, 19.0��� 29.7 in P. sagani sp. nov.). Presence of cuticular bars on legs I���III in P. sagani sp. nov. Larger eggs (70.0���74.0 ��m in P. magdalenae, 73.7���87.7 ��m in P. sagani sp. nov. excluding processes), differently shaped processes and with different type of apices (conical processes, usually with elongated apices, in P. magdalenae). Paramacrobiotus sagani sp. nov. differs from P. pius by having eye-spots, granulated cuticle (only on legs in P. pius at all stages of maturity), longer macroplacoid row (pt 41.4���43.1 in P. pius, 45.0��� 60.5 in P. sagani sp. nov.). Shorter claws (e.g. pt of total length of claws II���III internal and external collectively, 25.4���29.2 in P. pius, 18.7��� 24.5 in the P. sagani). Egg processes with cylindrical apices which are lacking in P. pius, its egg processes being simply truncated cones. Paramacrobiotus sagani sp. nov. differs from P. priviterae by having cuticular granulation (only on legs in P. priviterae also in small specimens), typical hufelandi - type claws with slender body and shorter distal portion of the branches and accessory points, while P. priviterae has less typically shaped claws for a Paramacrobiotus (see Binda et al. 2001, page 238, Fig. 3C���D), with stout main claw body, but with long, well tapering, distal portions of branches and long accessory points. Smaller eggs (90���107 ��m in P. priviterae, 73.7���87.7 ��m in P. sagani sp. nov. excluding processes) with 10���13 processes on circumference (14���17 processes for P. priviterae) and different apices of the processes (stout and thorn-shaped in P. priviterae). Paramacrobiotus sagani sp. nov. differs from P. sklodowskae by having a cuticular granulation (only on legs in P. sklodowskae at all stages of maturity). Generally, wider buccal tube (pt 17.0��� 20.5 in P. sklodowskae, 19.0��� 29.7 in P. sagani sp. nov.) with less caudal stylet support insertion point (pt 81.8���85.2 in P. sklodowskae, 77.1���81.5 in P. sagani sp. nov.). Slender claws (stouter in P. sklodowskae), presence of cuticular bars on legs I���III. Smaller eggs (88.4���96.9 ��m in P. sklodowskae, 73.7���87.7 ��m in P. sagani sp. nov. excluding processes), with different process apices, which are rounded in P. sklodowskae. The egg processes of P. sagani sp. nov. are most similar to P. vanescens (Pilato, Binda & CatanZaro, 1991; amended by Binda & Pilato 2001 and Pilato et al. 2001), from which the new species differs by having eye-spots and an obvious microplacoid (described as, ���little, faint, sometimes almost invisible��� in P. vanescens). Eggs with shorter processes (16���17 ��m in P. vanescens, 9.4���13.2 ��m in P. sagani sp. nov.), with slightly more cylindrical, elongated apices (just a short, indented crown on apices of P. vanescens). All those differences were confirmed by personal observation of type material of P. vanescens, including eggs, deposited in the Pilato and Binda collection., Published as part of Daza, Anisbeth, Caicedo, Mart��n, Lisi, Oscar & Quiroga, Sigmer, 2017, New records of tardigrades from Colombia with the description of Paramacrobiotus sagani sp. nov. and Doryphoribius rosanae sp. nov., pp. 29-50 in Zootaxa 4362 (1) on pages 32-39, DOI: 10.11646/zootaxa.4362.1.2, http://zenodo.org/record/1076122, {"references":["Maucci, W. (1973) Macrobiotus spallanzanii sp. nov. and redescription of Macrobiotus tenius Binda and Pilato (Tardigrada, Macrobiotidae). Bollettino di Zoologia, 40 (3 - 4), 261 - 267. https: // doi. org / 10.1080 / 11250007309429241","Pilato, G. (1975) On the taxonomic criteria of the Eutardigrada. Memorie dell'Istituto Italiano di Idrobiologia \" Dott. Marco De Marchi \", 32, 277 - 303.","Pilato, G. (1972) Structure, Intraspecific Variability and Systematic Value of the Buccal Armature of Eutardigrades. Zeitschrift fur zoologische Systematik und Evolutionsforschung, 1, 65 - 78. https: // doi. org / 10.1111 / j. 1439 - 0469.1972. tb 00785. x","Tumanov, D. V. (2005) Notes on the Tardigrada of Thailand, with a description of Macrobiotus alekseevi sp. nov. (Eutardigrada, Macrobiotidae). Zootaxa, 999 (1), 1 - 16. https: // doi. org / 10.11646 / zootaxa. 999.1.1","Pilato, G., Binda, M. G. & Lisi, O. (2002) Notes on tardigrades of the Seychelles with the description of two new species. Bolletino della Accademia Giornia di Scienze Naturali Catania, 35 (361), 503 - 517.","Pilato, G., Binda, M. G. & Lisi, O. (2004) Notes on tardigrades of the Seychelles with the description of three new species. Italian Journal of Zoology, 71, 171 - 178. https: // doi. org / 10.1080 / 11250000409356569","Bartels, P. J., Pilato, G., Lisi, O. & Nelson, D. R. (2009) Macrobiotus (Eutardigrada, Macrobiotidae) from the Great Smoky Mountains National Park, Tennessee / North Carolina, USA (North America): two new species and six new records. Zootaxa, 2022, 45 - 57.","Biserov, V. I. (1996) Macrobiotus lorenae sp. n., a new species of Tardigrada (Eutardigrada Macrobiotidae) from the Russian Far East. Arthropoda Selecta, 5 (1 / 2), 145 - 149.","Lisi, O., Binda, M. G. & Pilato, G. (2016) Eremobiotus ginevrae sp. nov. and Paramacrobiotus pius sp. nov., two new species of Eutardigrada. Zootaxa, 4103 (4), 344 - 360. http: // doi. org / 10.11646 / zootaxa. 4103.4.3","Michalczyk, L., Kaczmarek, L. & Weglarska, B. (2006) Macrobiotus sklodowskae sp. nov. (Tardigrada: Eutardigrada: Macrobiotidae, richtersi group) from Cyprus. Zootaxa, 1371, 45 - 56.","Pilato, G., Binda, M. G. & Catanzaro, R. (1991) Remarks of some tardigrades of the African fauna with the description of three new species of Macrobiotus Schultze 1834. Tropical Zoology, 4, 167 - 718. https: // doi. org / 10.1080 / 03946975.1991.10539487"]}

160. Paramacrobiotus sklodowskae (Michalczyk, Kaczmarek & Węglarska, 2006) [T] PaRamacRobiotus sklodowskae (Michalczyk et al., 2006) (Gąsiorek et al. 2017) Terra typica: Cyprus (Europe) Tunisia: • 36°43'20''N, 08°40'58''E; 668 m asl: Jendouba Governorate, Beni M'tir, forest, moss and moss and lichen on soil (2 samples). Gąsiorek et al. (2017) • 36°22'55''N, 10°07'03''E; 357 m asl: Zaghouan Governorate, Zaghouan, forest, lichen on soil. Gąsiorek et al. (2017) Record numbers. Tunisia: 2; total: 2. Remarks. Reported from Cyprus and Tunisia, indicates a Mediterranean distribution (Gąsiorek et al. 2017)., 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 57, DOI: 10.11646/zootaxa.4284.1.1, http://zenodo.org/record/1010400, {"references":["Michalczyk, L., Kaczmarek, L. & Weglarska, B. (2006) MacRobiotus sklodowskae sp. nov. (Tardigrada: Eutardigrada: Macrobiotidae, RichteRsi group) from Cyprus. Zootaxa, 1371, 45 - 56.","Gasiorek, P., Morek, W., Stec, D., Marnissi, J. & Michalczyk, L. (2017) The tardigrade fauna of Tunisia, with an integrative description of BRyodelphax maculatus sp. nov. (Tardigrada: Heterotardigrada: Echiniscoidea). African Zoology. http: // dx. doi. org / 10.1080 / 15627020.2017.1297688"]}

159. Paramacrobiotus richtersi (Murray, 1911) sensu lato [T] MacRobiotus RichteRsii Murray (Murray 1913, Pardi 1941) MacRobiotus schultzei Greeff 1866 (Rahm 1936) MacRobiotus RichteRsii J. Murray, 1911 (Rodriguez-Roda 1947) MacRobiotus schultzei Greeff 1866 (Rahm 1936, da Cunha & do Nascimento 1964) MacRobiotus RichteRsi (Iharos 1969a) MacRobiotus RichteRsi J. Murr., (1911) (Binda 1971) MacRobiotus RichteRsi J. Murr., 1911 (Pilato 1972) MacRobiotus RichteRsi Murray, 1911 (Pilato & Pennisi 1976, Binda 1984, Binda & Pilato 1987, Van Rompu et al. 1995, Meyer & Hinton 2009) MacRobiotus RichteRsi J. Murr. (Iharos 1978) M. RichteRsi (Ramløv & Kristensen 1985) M. RichteRsi J. Murray, 1911 (Séméria 1986) MacRobiotus cf. RichteRsi Murray, 1911 (Middleton 2003, Kaczmarek & Michalczyk 2004b) PaRamacRobiotus cf. RichteRsi (Murray, 1911)? (Gąsiorek et al. 2017) Terra typica: Ireland (Europe) Algeria: • 36°49′N, 05°47′E; 0 m asl: Jijel Province, Jijel, Les Aftis beach (Ziama Monsouria). Binda & Pilato (1987) • 36°46′N, 05°06′E; 400 m asl: Béjaïa Province, Pic des Singes. Binda & Pilato (1987) • 36°40′N, 05°09′E; 50 m asl: Béjaïa Province, Tichi [Tichy]. Binda & Pilato (1987) • 36°38′N, 05°45′E; 1,200 m asl: Jijel Province, M’Sid-Ech-Cheta (Texenna Pass). Binda & Pilato (1987) • 36°28′N, 04°14′E; 1,700 m asl: Tizi Ouzou Province, Massif of Djurdjura (Assouille depression). Binda & Pilato (1987) • 34°40′N, 06°11′E; 0 m asl: Biskra Province, 30 km from Biskra, El Oued [Zeribet el Oued]. Binda & Pilato (1987) Angola: • 07°47′S, 15°01′E; 650 m asl: Uíge Province, near Carmona [Uíge], Estate at Pumba Loge, mosses and lichens. da Cunha & do Nascimento (1964) • 14°44′S, 15°01′E: Undefined locality, Huíla Province, Impaca, mosses and lichens. da Cunha & do Nascimento (1964) Cameroon: • 04°57′N, 09°56′E; 800 m asl: Littoral Region, Kongsamba [Nkongsamba], moss on tree. Iharos (1969a) • 04°13′N, 09°10′E; 4,000 m asl: Southwest Region, Mount Cameroon National Park, Mount Cameroon, near fumaroles (80°C), moss on soil. Iharos (1969a) • 04°12′N, 09°12′E; 2,900–3,000 m asl: Southwest Region, Mount Cameroon National Park, Mount Cameroon, moss on trees and lava blocks. Iharos (1969a) • 04°11′N, 09°12′E; 1,900–1,950 m asl: Southwest Region, Mount Cameroon National Park, Mount Cameroon, primeval forest, above bothy No. 1, moss on trees and lava. Iharos (1969a) • 04°10′N, 09°13′E; 1,300–1,500 m asl: Southwest Region, Mount Cameroon National Park, Mount Cameroon, rainforest, moss on tree. Iharos (1969a) • 04°10′N, 09°13′E; 1,660–1,800 m asl: Southwest Region, Mount Cameroon National Park, Mount Cameroon, rainforest, moss on tree. Iharos (1969a) Equatorial Guinea: • 03°39′N, 08°49′E; 1,500 m asl: Fernando Póo [Bioko Island], Bioko Norte Province, Pico Basilé, among mosses and liverworts. Rodriguez-Roda (1947) Ethiopia: • 04°03′N, 38°19′E; 1,700 m asl: Oromia Region, Mēga, former English residence, moss (PleuRochaete sp.) on tree (JunipeRus pRoceRa), moss in stream, moss on (JunipeRus pRoceRa) and moss (NeckeRa sp.) from tree (JunipeRus pRoceRa) (4 samples). Pardi (1941) Gabon: • 00°18′N, 09°30′E; 10–40 m asl: Estuaire Province, near Libreville, Owendo, moss on tree. Iharos (1969a) • Israel and Palestinian National Authority: • 32°53′N, 35°34′E; - 200 m asl: N shore of Sea of Galilee, Capernaum, moss (Hypnum sp.). Rahm (1936) Kenya: • 00°01′S, 37°54′E: Undefined locality, British East Africa [Kenya], moss. Murray (1913) 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) Libya: • 32°29′N, 20°50′E; 350 m asl: Marj District, Barce [Marj], moss. Pilato & Pennisi (1976) Morocco: • 33°31′N, 05°07′W; 1,650 m asl: Meknès-Tafilalet Region, Ras al Ma [Res el Ma], mosses on trees (Cedrus) and rocks. Séméria (1986) • 32°56′N, 05°40′W; 1,700 m asl [850 m asl]: Meknès-Tafilalet Region, Khenifra [Khénifra], moss. Binda (1971), Pilato (1972) • 32°25′N, 05°13′W; 1,750 m asl: Meknès-Tafilalet Region, S of Azrou, cedar forest, moss on basalt boulders in the forest glade. Ramløv & Kristensen (1985) • 31°08′N, 07°55′W; 1,700 m asl: Tadla Tadla-Azilal Region, Imlil, walnut forest, moss on limestone cliffs. Ramløv & Kristensen (1985) Republic of South Africa: • 24°38′S, 30°51′E; 1,800 m asl: Undefined locality, Gauteng or Mpumalanga Province, Dragon Mountains, near the canyon of Blade River, moss on soil in the grass. Kaczmarek & Michalczyk (2004b) • 25°45′S, 28°11′E; 1,300 m asl: Gauteng Province, Pretoria, moss. Murray (1913) • 29°44′S, 22°44′E: Undefined locality, Cape Colony [Cape Province]. 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) • 34°01′S, 23°55′E; 200 m asl: Eastern Cape Province, Tizitzikama / Tzitzikama [Tsitsikamma National Park], moss. Binda (1984) Tunisia: • 36°57′N, 08°45′E; 0 m asl: Jendouba Governorate, surroundings of Tabarka. Binda & Pilato (1987) • 36°53'57''N, 10°18'19''E; 67 m asl: Tunis Governorate, Gammaarth, rural, lichen on tree and moss on soil (2 samples). Gąsiorek et al. (2017) • 36°46′N, 08°41′E; 700 m asl: Jendouba Governorate, surroundings of Ain Draham, forest of QueRcus saubeR and Q. libanotis, moss on soil and vegetable debris mixed with soil (2 samples). Iharos (1978) • 36°43'19''N, 08°43'30''E; 474 m asl: Jendouba Governorate, Beni M'tir, forest, moss on soil. Gąsiorek et al. (2017) • 36°07′N, 08°30′E; 350 m asl: Kef Governorate, along the road between El Kef and Sakiet, 23 km W El Kef, Qued el Kohl, scrub, sparse vegetation, with stony ground and thin layer of surface soil, soil with vegetable debris or litter under trees (Pinus halepensis and JunipeRus phoenicus). 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 and moss on soil mixed with fine-grained soil (2 samples). Iharos (1978) • 36°22'55''N, 10°07'03''E; 357 m asl: Zaghouan Governorate, Zaghouan, forest, mosses on soil (4 samples). Gąsiorek et al. (2017) Uganda: • 01°22′N, 32°17′E: Undefined locality. Murray (1913) Zimbabwe: • 17°46′S, 31°03′E; 1 500 m asl: Harare City, Tobacco Research Board, nematology seed bed site, under grass, dry soil. Van Rompu et al. (1995) • 18°20′S, 26°28′E; 700 m asl: Matabeleland North Province, 370 km from Bulawayo, beyond Hwange village, along the road to Livingstone, mopane forest, dry soil. Van Rompu et al. (1995) • 18°44′S, 26°57′E; 1,100 m asl: Matabeleland North Province, Hwange National Park, near Main Camp, under grass and shrubs, dry sandy soil. Van Rompu et al. (1995) • 18°56′S, 27°46′E; 1,000 m asl: Matabeleland North Province, 182 km from Bulawayo alongside the road to Hwange, Lupane, around a tree, dry sandy soil with some organic debris. Van Rompu et al. (1995) • 20°06′S, 31°00′E; 1,050 m asl: Masvingo Province, Popoteke Gorge [Popoteki Gorge], along the river, under reeds, humid soil. Van Rompu et al. (1995) • 20°26′S, 28°31′E; 1,350 m asl: Matabeleland South Province, Matobo National Park, Matopos, Moth Shrine, under grass, dry soil. Van Rompu et al. (1995) • 20°29′S, 28°31′E; 1,350 m asl: Matabeleland South Province, Matobo National Park, Matopos, close to the parking near White Rhino Cave, dry river bed, under grass and mopane tree, dry soil. Van Rompu et al. (1995) • 20°31′S, 29°21′E; 1,100 m asl: Matabeleland South Province, right side of the road from Masvingo to Bulawayo, 175 km from Masvingo, under shrubs and mopane tree, dry soil. Van Rompu et al. (1995) • 20°32′S, 28°28′E; 1,350 m asl: Matabeleland South Province, Matobo National Park, Matopos, Nswatugi Cave, 10 m from the parking, under grass and various trees, dry soil. Van Rompu et al. (1995) • 20°33′S, 28°31′E; 1,400 m asl: Matabeleland South Province, Matobo National Park, Matopos, left from the path to Pomongwe Cave, 10 m before the cave, under various trees, dry soil. Van Rompu et al. (1995) • 21°27′S, 32°04′E; 200 m asl: Masvingo Province, Gonaredzhou National Park [South Gonarezhou National Park], Mwatambo viewpoint, under grass and shrubs, dry soil. Van Rompu et al. (1995) Record numbers. Algeria: 6, Angola: 2, Cameroon: 6, Equatorial Guinea: 1, Ethiopia: 1, Gabon: 1, Israel and Palestinian National Authority: 1, Kenya: 1, Lesotho: 2, Libya: 1, Morocco: 4, Republic of South Africa: 5, Tunisia: 7, Uganda: 1, Zimbabwe: 11; total: 50. Remarks. Paramacrobiotus richtersi was, until recently, believed to be cosmopolitan (McInnes 1994). New data has shown that this is the nominal species for a species complex – the ‘ richtersi group’ – requiring careful examination of both adults and eggs. Most African reports pre-date the taxonomic re-assessment and several lack eggs, so proper identification cannot be confirmed (e.g. Van Rompu et al. (1995), whose drawings could be interpreted as an adult of the M. harmsworthi group). Paramacrobiotus richtersi was originally described from Europe (Murray, 1911), suggesting ‘some confusion’ and that all African records should be treated with caution until the original material is re-examined or the species is otherwise verified from the region.

157. Paramacrobiotus kenianus Schill, F��rster, Dandekar & Wolf, 2010 [T] MacRobiotus ���RichteRsi gRoups��� 1 and 2 (Hengherr et al. 2008, 2009a) MacRobiotus ���RichteRsi gRoups��� 2 and 1 (Hengherr et al. 2009b) MacRobiotus ���RichteRsi gRoups��� 2 and 3 (Guidetti et al. 2009) PaRamacRobiotus kenianus sp. nov. (Schill et al. 2010) Terra typica: Kenya (Africa) Kenya: ��� 00��18���S; 36��05���E; 1,800 m asl: Type Locality: Nakuru County, Nakuru, moss. Hengherr et al. (2008, 2009a, b), Guidetti et al. (2009), Schill et al. (2010) ��� 00��43���S; 36��26���E; 1,900 m asl: Nakuru County, Naivasha, moss. Hengherr et al. (2008, 2009a, b), Guidetti et al. (2009), Schill et al. (2010) Record numbers. Kenya: 2; total: 2. Remarks. This species, identified only via DNA barcoding and biophysical data, is morphologically identical to P. richtersi (Guidetti et al. 2009, Schill et al. 2010). The International Commission of the Zoological Nomenclature code states, ���An author, when drawing up the description of a new nominal taxon, should include comparisons with appropriate related taxa in order to assist later identification of the taxon. Namebearing type material should be illustrated (or a reference given to such illustration).��� (http://www.nhm.ac.uk/ hosted-sites/iczn/code/index.jsp?nfv=true&booksection=appendixB). This taxon, in the absence of such illustration, remains a cryptic species identifiable only via DNA and biophysical data. Currently endemic to Kenya., 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 55, DOI: 10.11646/zootaxa.4284.1.1, http://zenodo.org/record/1010400, {"references":["Schill, R. O., Forster, F., Dandekar, T. & Wolf, N. (2010) Using compensatory base change analysis of internal transcribed spacer 2 secondary structures to identify three new species in PaRamacRobiotus (Tardigrada). ORganisms DiveRsity & Evolution, 10 (4), 287 - 296. https: // doi. org / 10.1007 / s 13127 - 010 - 0025 - z","Hengherr, S., Heyer, A. G., Kohler, H. R. & Schill, R. O. (2008) Trehalose and anhydrobiosis in tardigrades-evidence for divergence in responses to dehydration. FEBS JouRnal, 275, 281 - 288. https: // doi. org / 10.1111 / j. 1742 - 4658.2007.06198. x","Hengherr, S., Worland, M. R., Reuner, A., Brummer, F. & Schill, R. O. (2009 a) High-temperature tolerance in anhydrobiotic tardigrades is limited by glass transition. Physiological and Biochemical Zoology, 82, 749 - 755. https: // doi. org / 10.1086 / 605954","Hengherr, S., Worland, M. R., Reuner, A., Brummer, F. & Schill, R. O. (2009 b) Freeze tolerance, supercooling points and ice formation: comparative studies on the subzero temperature survival of limno-terrestrial tardigrades. JouRnal of ExpeRimental Biology, 212, 802. https: // doi. org / 10.1242 / jeb. 025973","Guidetti, R., Schill, R. O., Bertolani, R., Dandekar, T. & Wolf, M. (2009) New molecular data for tardigrade phylogeny, with the erection of PaRamacRobiotus gen. nov. JouRnal of Zoological Systematics and EvolutionaRy ReseaRch, 47, 315 - 321. https: // doi. org / 10.1111 / j. 1439 - 0469.2009.00526. x"]}

156. Paramacrobiotus areolatus (Murray, 1907c) sensu lato [T] MacRobiotus aReolatus Murray, 1907 (Binda 1984, Binda & Pilato 1987) PaRamacRobiotus cf. aReolatus (Murray, 1907)? (Gąsiorek et al. 2017) Terra typica: Svalbard (Spitsbergen) Algeria: • 36°40′N, 04°36′E; 1,400 m asl: Tizi Ouzou Province, Akfadou Forest. Binda & Pilato (1987) Mozambique: • 23°52′S, 35°23′E; 0 m asl: Inhambene Province, Inhambene, moss. Binda (1984) Tunisia: • 37°10'46''N, 10°15'11''E; 120 m asl: Bizerte Governorate, Raf Raf, forest, moss on soil. Gąsiorek et al. (2017) • 36°53'57''N, 10°18'19''E; 67 m asl: Tunis Governorate, Gammarth, rural, moss on soil. Gąsiorek et al. (2017) • 36°46'30''N, 08°44'23''E; 636 m asl: Jendouba Governorate, Beni M'tir, forest, moss on soil. Gąsiorek et al. (2017) • 36°47'51''N, 08°52'24''E; 590 m asl: Jendouba Governorate, Beni M'tir, forest, moss on soil. Gąsiorek et al. (2017) • 36°45'59''N, 08°47'52''E; 561 m asl: Jendouba Governorate, Beni M'tir, forest, moss on soil. Gąsiorek et al. (2017) • 36°44'20''N, 08°43'43''E; 511 m asl: Jendouba Governorate, Beni M'tir, forest, moss on soil. Gąsiorek et al. (2017) • 36°44'13''N, 08°40'59''E; 765 m asl: Jendouba Governorate, Beni M'tir, forest, moss on soil. Gąsiorek et al. (2017) • 36°43'45''N, 08°44'11''E; 373 m asl: Jendouba Governorate, Beni M'tir, forest, moss on soil. Gąsiorek et al. (2017) • 36°43'43''N, 08°40'50''E; 742 m asl: Jendouba Governorate, Beni M'tir, forest, moss on soil. Gąsiorek et al. (2017) • 36°43'20''N, 08°40'58''E; 668 m asl: Jendouba Governorate, Beni M'tir, forest, moss on soil. Gąsiorek et al. (2017) • 36°43'19''N, 08°43'30''E; 474 m asl: Jendouba Governorate, Beni M'tir, forest, moss on soil. Gąsiorek et al. (2017) • 36°42'48''N, 08°43'30''E; 371 m asl: Jendouba Governorate, Beni M'tir, forest, moss on soil. Gąsiorek et al. (2017) • 36°42'10''N, 08°40'46''E; 462 m asl: Jendouba Governorate, Beni M'tir, forest, moss on soil. Gąsiorek et al. (2017) Record numbers. Algeria: 1, Mozambique: 1, Tunisia: 13; total: 15. Remarks. The genus Paramacrobiotus (until recently a species-group within Macrobiotus) has been divided into richtersi and areolatus (with and without macroplacoid, respectively) species groups. Paramacrobiotus areolatus, once considered cosmopolitan (McInnes 1994), is now recognised as the nominal species for a group of very similar taxa that require careful taxonomic examination of adults and egg morphology for correct identification. A few new (including African) species of the areolatus group (e.g. Michalczyk & Kaczmarek 2006b, Degma et al. 2008, Kaczmarek et al. 2014b). Both Richters (1908) and Binda (1984) predate the taxonomic changes and failed to find eggs, essential for correct species identification. This, in conjunction with P. areolatus being originally described from the Arctic, implies all African records should be treated with caution until relevant examples are re-examined or new African material is confirmed.

158. Paramacrobiotus priviterae (Binda, Pilato, Moncada & Napolitano, 2001) [T] MacRobiotus pRiviteRae n. sp. (Binda et al. 2001) Terra typica: Guinea (Africa) Guinea: • 11°06′N, 13°49′W; 200 m asl: Type Locality: Boké Region, Sangarédi, moss (IsopteRigium teneRum). Binda et al. (2001) Record numbers. Guinea: 1; total: 1. Remarks. This species belongs to richtersi group and is currently endemic to Guinea., 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 55, DOI: 10.11646/zootaxa.4284.1.1, http://zenodo.org/record/1010400, {"references":["Binda, M. G., Pilato, G., Moncada, E. & Napolitano, A. (2001) Some tardigrades from Central Africa with the description of two new species: MacRobiotus Ragonesei and M. pRiviteRae (Eutardigrada: Macrobiotidae). TRopical Zoology, 14, 233 - 242. https: // doi. org / 10.1080 / 03946975.2001.10531155"]}

159. Paramacrobiotus richtersi (Murray, 1911) sensu lato [T] MacRobiotus RichteRsii Murray (Murray 1913, Pardi 1941) MacRobiotus schultzei Greeff 1866 (Rahm 1936) MacRobiotus RichteRsii J. Murray, 1911 (Rodriguez-Roda 1947) MacRobiotus schultzei Greeff 1866 (Rahm 1936, da Cunha & do Nascimento 1964) MacRobiotus RichteRsi (Iharos 1969a) MacRobiotus RichteRsi J. Murr., (1911) (Binda 1971) MacRobiotus RichteRsi J. Murr., 1911 (Pilato 1972) MacRobiotus RichteRsi Murray, 1911 (Pilato & Pennisi 1976, Binda 1984, Binda & Pilato 1987, Van Rompu et al. 1995, Meyer & Hinton 2009) MacRobiotus RichteRsi J. Murr. (Iharos 1978) M. RichteRsi (Raml��v & Kristensen 1985) M. RichteRsi J. Murray, 1911 (S��m��ria 1986) MacRobiotus cf. RichteRsi Murray, 1911 (Middleton 2003, Kaczmarek & Michalczyk 2004b) PaRamacRobiotus cf. RichteRsi (Murray, 1911)? (Gąsiorek et al. 2017) Terra typica: Ireland (Europe) Algeria: ��� 36��49���N, 05��47���E; 0 m asl: Jijel Province, Jijel, Les Aftis beach (Ziama Monsouria). Binda & Pilato (1987) ��� 36��46���N, 05��06���E; 400 m asl: B��ja��a Province, Pic des Singes. Binda & Pilato (1987) ��� 36��40���N, 05��09���E; 50 m asl: B��ja��a Province, Tichi [Tichy]. Binda & Pilato (1987) ��� 36��38���N, 05��45���E; 1,200 m asl: Jijel Province, M���Sid-Ech-Cheta (Texenna Pass). Binda & Pilato (1987) ��� 36��28���N, 04��14���E; 1,700 m asl: Tizi Ouzou Province, Massif of Djurdjura (Assouille depression). Binda & Pilato (1987) ��� 34��40���N, 06��11���E; 0 m asl: Biskra Province, 30 km from Biskra, El Oued [Zeribet el Oued]. Binda & Pilato (1987) Angola: ��� 07��47���S, 15��01���E; 650 m asl: U��ge Province, near Carmona [U��ge], Estate at Pumba Loge, mosses and lichens. da Cunha & do Nascimento (1964) ��� 14��44���S, 15��01���E: Undefined locality, Hu��la Province, Impaca, mosses and lichens. da Cunha & do Nascimento (1964) Cameroon: ��� 04��57���N, 09��56���E; 800 m asl: Littoral Region, Kongsamba [Nkongsamba], moss on tree. Iharos (1969a) ��� 04��13���N, 09��10���E; 4,000 m asl: Southwest Region, Mount Cameroon National Park, Mount Cameroon, near fumaroles (80��C), moss on soil. Iharos (1969a) ��� 04��12���N, 09��12���E; 2,900���3,000 m asl: Southwest Region, Mount Cameroon National Park, Mount Cameroon, moss on trees and lava blocks. Iharos (1969a) ��� 04��11���N, 09��12���E; 1,900���1,950 m asl: Southwest Region, Mount Cameroon National Park, Mount Cameroon, primeval forest, above bothy No. 1, moss on trees and lava. Iharos (1969a) ��� 04��10���N, 09��13���E; 1,300���1,500 m asl: Southwest Region, Mount Cameroon National Park, Mount Cameroon, rainforest, moss on tree. Iharos (1969a) ��� 04��10���N, 09��13���E; 1,660���1,800 m asl: Southwest Region, Mount Cameroon National Park, Mount Cameroon, rainforest, moss on tree. Iharos (1969a) Equatorial Guinea: ��� 03��39���N, 08��49���E; 1,500 m asl: Fernando P��o [Bioko Island], Bioko Norte Province, Pico Basil��, among mosses and liverworts. Rodriguez-Roda (1947) Ethiopia: ��� 04��03���N, 38��19���E; 1,700 m asl: Oromia Region, Mēga, former English residence, moss (PleuRochaete sp.) on tree (JunipeRus pRoceRa), moss in stream, moss on (JunipeRus pRoceRa) and moss (NeckeRa sp.) from tree (JunipeRus pRoceRa) (4 samples). Pardi (1941) Gabon: ��� 00��18���N, 09��30���E; 10���40 m asl: Estuaire Province, near Libreville, Owendo, moss on tree. Iharos (1969a) ��� Israel and Palestinian National Authority: ��� 32��53���N, 35��34���E; - 200 m asl: N shore of Sea of Galilee, Capernaum, moss (Hypnum sp.). Rahm (1936) Kenya: ��� 00��01���S, 37��54���E: Undefined locality, British East Africa [Kenya], moss. Murray (1913) 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) Libya: ��� 32��29���N, 20��50���E; 350 m asl: Marj District, Barce [Marj], moss. Pilato & Pennisi (1976) Morocco: ��� 33��31���N, 05��07���W; 1,650 m asl: Mekn��s-Tafilalet Region, Ras al Ma [Res el Ma], mosses on trees (Cedrus) and rocks. S��m��ria (1986) ��� 32��56���N, 05��40���W; 1,700 m asl [850 m asl]: Mekn��s-Tafilalet Region, Khenifra [Kh��nifra], moss. Binda (1971), Pilato (1972) ��� 32��25���N, 05��13���W; 1,750 m asl: Mekn��s-Tafilalet Region, S of Azrou, cedar forest, moss on basalt boulders in the forest glade. Raml��v & Kristensen (1985) ��� 31��08���N, 07��55���W; 1,700 m asl: Tadla Tadla-Azilal Region, Imlil, walnut forest, moss on limestone cliffs. Raml��v & Kristensen (1985) Republic of South Africa: ��� 24��38���S, 30��51���E; 1,800 m asl: Undefined locality, Gauteng or Mpumalanga Province, Dragon Mountains, near the canyon of Blade River, moss on soil in the grass. Kaczmarek & Michalczyk (2004b) ��� 25��45���S, 28��11���E; 1,300 m asl: Gauteng Province, Pretoria, moss. Murray (1913) ��� 29��44���S, 22��44���E: Undefined locality, Cape Colony [Cape Province]. 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) ��� 34��01���S, 23��55���E; 200 m asl: Eastern Cape Province, Tizitzikama / Tzitzikama [Tsitsikamma National Park], moss. Binda (1984) Tunisia: ��� 36��57���N, 08��45���E; 0 m asl: Jendouba Governorate, surroundings of Tabarka. Binda & Pilato (1987) ��� 36��53'57''N, 10��18'19''E; 67 m asl: Tunis Governorate, Gammaarth, rural, lichen on tree and moss on soil (2 samples). Gąsiorek et al. (2017) ��� 36��46���N, 08��41���E; 700 m asl: Jendouba Governorate, surroundings of Ain Draham, forest of QueRcus saubeR and Q. libanotis, moss on soil and vegetable debris mixed with soil (2 samples). Iharos (1978) ��� 36��43'19''N, 08��43'30''E; 474 m asl: Jendouba Governorate, Beni M'tir, forest, moss on soil. Gąsiorek et al. (2017) ��� 36��07���N, 08��30���E; 350 m asl: Kef Governorate, along the road between El Kef and Sakiet, 23 km W El Kef, Qued el Kohl, scrub, sparse vegetation, with stony ground and thin layer of surface soil, soil with vegetable debris or litter under trees (Pinus halepensis and JunipeRus phoenicus). 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 and moss on soil mixed with fine-grained soil (2 samples). Iharos (1978) ��� 36��22'55''N, 10��07'03''E; 357 m asl: Zaghouan Governorate, Zaghouan, forest, mosses on soil (4 samples). Gąsiorek et al. (2017) Uganda: ��� 01��22���N, 32��17���E: Undefined locality. Murray (1913) Zimbabwe: ��� 17��46���S, 31��03���E; 1 500 m asl: Harare City, Tobacco Research Board, nematology seed bed site, under grass, dry soil. Van Rompu et al. (1995) ��� 18��20���S, 26��28���E; 700 m asl: Matabeleland North Province, 370 km from Bulawayo, beyond Hwange village, along the road to Livingstone, mopane forest, dry soil. Van Rompu et al. (1995) ��� 18��44���S, 26��57���E; 1,100 m asl: Matabeleland North Province, Hwange National Park, near Main Camp, under grass and shrubs, dry sandy soil. Van Rompu et al. (1995) ��� 18��56���S, 27��46���E; 1,000 m asl: Matabeleland North Province, 182 km from Bulawayo alongside the road to Hwange, Lupane, around a tree, dry sandy soil with some organic debris. Van Rompu et al. (1995) ��� 20��06���S, 31��00���E; 1,050 m asl: Masvingo Province, Popoteke Gorge [Popoteki Gorge], along the river, under reeds, humid soil. Van Rompu et al. (1995) ��� 20��26���S, 28��31���E; 1,350 m asl: Matabeleland South Province, Matobo National Park, Matopos, Moth Shrine, under grass, dry soil. Van Rompu et al. (1995) ��� 20��29���S, 28��31���E; 1,350 m asl: Matabeleland South Province, Matobo National Park, Matopos, close to the parking near White Rhino Cave, dry river bed, under grass and mopane tree, dry soil. Van Rompu et al. (1995) ��� 20��31���S, 29��21���E; 1,100 m asl: Matabeleland South Province, right side of the road from Masvingo to Bulawayo, 175 km from Masvingo, under shrubs and mopane tree, dry soil. Van Rompu et al. (1995) ��� 20��32���S, 28��28���E; 1,350 m asl: Matabeleland South Province, Matobo National Park, Matopos, Nswatugi Cave, 10 m from the parking, under grass and various trees, dry soil. Van Rompu et al. (1995) ��� 20��33���S, 28��31���E; 1,400 m asl: Matabeleland South Province, Matobo National Park, Matopos, left from the path to Pomongwe Cave, 10 m before the cave, under various trees, dry soil. Van Rompu et al. (1995) ��� 21��27���S, 32��04���E; 200 m asl: Masvingo Province, Gonaredzhou National Park [South Gonarezhou National Park], Mwatambo viewpoint, under grass and shrubs, dry soil. Van Rompu et al. (1995) Record numbers. Algeria: 6, Angola: 2, Cameroon: 6, Equatorial Guinea: 1, Ethiopia: 1, Gabon: 1, Israel and Palestinian National Authority: 1, Kenya: 1, Lesotho: 2, Libya: 1, Morocco: 4, Republic of South Africa: 5, Tunisia: 7, Uganda: 1, Zimbabwe: 11; total: 50. Remarks. Paramacrobiotus richtersi was, until recently, believed to be cosmopolitan (McInnes 1994). New data has shown that this is the nominal species for a species complex ��� the ��� richtersi group��� ��� requiring careful examination of both adults and eggs. Most African reports pre-date the taxonomic re-assessment and several lack eggs, so proper identification cannot be confirmed (e.g. Van Rompu et al. (1995), whose drawings could be interpreted as an adult of the M. harmsworthi group). Paramacrobiotus richtersi was originally described from Europe (Murray, 1911), suggesting ���some confusion��� and that all African records should be treated with caution until the original material is re-examined or the species is otherwise verified from the region., 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 55-57, DOI: 10.11646/zootaxa.4284.1.1, http://zenodo.org/record/1010400, {"references":["Murray, J. (1911) Arctiscoida. PRoceedings of the Royal IRish Academy, 31, 1 - 16.","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","Pardi, L. (1941) Tardigrada. Missione Biologica Sagan-Omo, Reale Accademia d'Italia, Roma Zoologia, 6, 221 - 232.","Rahm, G. (1936) Tardigraden aus Palastina. ZoologischeR AnzeigeR, 115, 65 - 76.","Rodriguez-Roda, J. (1947) Algunos Tardigrados de Fernando Poo. Publicaciones del Instituto de Biologia Aplicada, 4, 149 - 159.","Da Cunha, A. X. da & Nascimento Ribeiro, F. (1964) Tardigrados de Angola. GaRcia de ORta, Lisboa, 12, 397 - 406.","Iharos, G. (1969 a) Tardigraden aus Mittelwestafrika. Opuscula Zoologica (Budapest), 9, 115 - 120.","Pilato, G. & Pennisi, G. (1976) Prime notizie sui Tardigradi della Cirenaica. Animalia, 3, 243 - 258.","Van Rompu, E. A., De Smet, W. H. & Coomans, A. (1995) Some terrestrial tardigrades from Zimbabwe. Biologisch JaaRbock Dodonea, 62, 48 - 55.","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","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.","Semeria, Y. (1986) Notes sur les Tardigrades du Maroc. Biocosme mesogeen, Nice, 3, 117 - 119.","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","Kaczmarek, L. & Michalczyk, L. (2004 b) Notes on some tardigrades from South Africa, with the description of Diphascon (Diphascon) zaniewi sp. nov. (Eutardigrada: Hypsibiidae). Zootaxa, 576 (1), 1 - 6. https: // doi. org / 10.11646 / zootaxa. 576.1.1","Gasiorek, P., Morek, W., Stec, D., Marnissi, J. & Michalczyk, L. (2017) The tardigrade fauna of Tunisia, with an integrative description of BRyodelphax maculatus sp. nov. (Tardigrada: Heterotardigrada: Echiniscoidea). African Zoology. http: // dx. doi. org / 10.1080 / 15627020.2017.1297688","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"]}

161. Paramacrobiotus vanescens (Pilato, Binda & Catanzaro, 1991) [T] MacRobiotus vanescens n. sp. (Pilato et al. 1991) MacRobiotus vanescens Pilato, Binda & Catanzaro 1991 (Binda & Pilato 1995a, Pilato et al. 2003) Terra typica: Tanzania (Africa) Democratic Republic of Congo: • 00°04′S, 29°18′E; 1,950 m asl: North Kivu Province, outskirts of Lukanga, liverwort (TaRgionia hypophylla). Pilato et al. (2003) Guinea: • 09°31′N, 13°42′W; 0 m asl: Conakry Region, Conkary, moss (BaRbula sp.). Pilato et al. (2003) Republic of Zambia: • 17 ° 55 ′ S, 25 ° 52 ′ E; 890 m asl: Southern Province, near Livingstone, in the Victoria Falls gorge, mosses and leaf litter on soil. Zawierucha et al. (2012) Tanzania: • 03°09′S, 35°29′E: Type Locality: Undefined locality, Arusha Region, Ngorongoro Volcano, mosses. Pilato et al. (1991) • 03°12′S, 37°31′E; 2,200–2,400 m asl: Kilimanjaro Region, near Marangu [gate], outskirts of the entrance of Kilimanjaro National Park, mosses. Binda & Pilato (1995a) Record numbers. Democratic Republic of Congo: 1, Guinea: 1, Republic of Zambia: 1, Tanzania: 2; total: 5. Remarks. This species belongs to richtersi group and is currently reported from a central band across 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 57-58, DOI: 10.11646/zootaxa.4284.1.1, http://zenodo.org/record/1010400, {"references":["Pilato, G., Binda, M. G. & Catanzaro, R. (1991) Remarks on some tardigrades of the African fauna with the description of three new species of MacRobiotus Schultze 1834. TRopical Zoology, 4, 167 - 178. https: // doi. org / 10.1080 / 03946975.1991.10539487","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","Zawierucha, K., Michalczyk, L. & Kaczmarek, L. (2012) The first record of Tardigrada from the Republic of Zambia, with a description of DoRyphoRibius niedbalai sp. nov. (Eutardigrada: Isohypsibiidae, the evelinae group). AfRican Zoology, 47 (2), 275 - 284. https: // doi. org / 10.3377 / 004.047.0224"]}

espanolDebido a la gran cantidad de especies y a las dificultades en su identificacion morfologica, muchos estudios taxonomicos actuales de tardigrados se basan en el uso de tecnicas geneticas y moleculares. En Colombia son necesarios estudios enfocados en esta disciplina que permitan complementar la informacion morfologica; por consiguiente se analizaron siete secuencias del gen COI en el genero de tardigrado Paramacrobiotus de la cuenca baja del rio Manzanares, en Santa Marta, y se compararon con las secuencias del mismo genero presentes en la base de datos de GenBank a traves de un analisis de divergencia especifica y analisis de vecino mas cercano (NJ). Las secuencias obtenidas en el estudio mostraron una clara diferenciacion de las secuencias de GenBank y una baja distancia intraespecifica, obteniendose el codigo de barras genetico correspondiente a una sola especie. Las diferencias de los resultados con el analisis morfologico reflejan la eficacia de tecnicas moleculares como los codigos de barras y evidencian la importancia de la integracion de tecnicas moleculares a los estudios taxonomicos de los tardigrados. EnglishBecause of the great number of species and troubles with their morphological identification, many current taxonomic studies in tardigrades are based on the use of genetic and molecular techniques. In Colombia, studies focused in this discipline are needed to be able to complement the morphological information; consequently, seven COI gene sequences were analyzed in the genus of tardigrade Paramacrobiotus at the lower basin of the Manzanares River, in Santa Marta, and were compared with the sequences of the same genus present in the GenBank database through an analysis of specific divergence and neighbor-joining (NJ). The sequences obtained in the study showed a clear differentiation of the GenBank sequences, while at the same time showed a low intraspecific distance, obtaining a DNA barcoding for a single species. The differences obtained in comparison with the morphological analyses reflect the effectiveness of molecular techniques such as barcoding, emphasizing the importance of integration of molecular techniques to taxonomic studies on tardigrades.

Reproductive strategy is an important aspect of biological diversity. In tardigrades, several reproductive modes, including sexual reproduction, are known. However, tardigrade mating behavior has been observed only rarely in most species, and in some cases, especially in the freely ovipositing eutardigrades, remains entirely unknown. In the present study, we cultured two sexually reproducing tardigrade species that lay eggs freely, Paramacrobiotus sp. TYO strain and Macrobiotus shonaicus, to investigate and compare their courtship, mating, and chromosome morphology. Mating behavior was observed and recorded in both species. The entire mating sequence, including courtship, was categorized into five discrete steps common to two species, as follows: [1] Tracking: the male tracks and orientates toward the female; [2] Touching: the male makes contact with the cloaca of the female; [3] Standstill: the female ceases movement until male ejaculation is complete; [4] Ejaculation: the male curls its caudal end and ejaculates into the cloaca from close range; [5] Contraction: the female contracts its ventral side after ejaculation to capture spermatozoa deposited in the external environment in close proximity to the cloaca. Some notable differences between the two species were observed in the steps 3–4. First, oviposition was observed at 40 min in Paramacrobiotus sp. TYO strain, and a few days after mating in M. shonaicus, respectively. Comparisons of chromosome morphology before and after mating indicated that oocytes are arrested at metaphase I in both species. Spermatozoa attach to the interior of the chorion of laid eggs.

Paramacrobiotus pius sp. nov. (Fig. 5) Type locality. Gangi (Palermo, Sicily), 37 �� 48 ���00��� N 14 �� 12 ��� 00���E. Former monastery of Gangivecchio (Gangi, Palermo) at 850 m a.s.l.: moss sample collected on rock by Pilato & Binda in August 2002. Material examined. Holotype (slide no. 5562), 7 paratypes (slides no. 5562 ��� 3; 5704, 5706��� 7), and 10 eggs (slides nos. 5562���5563; 5705���5707); some with fully developed embryos. Paramacrobiotus pius sp . nov.No. Slide5562 paratype 5706 newborn 5704 paratypenewborn 5704 paratype5562 holotype5704 paratypeBody length???270?Buccal tube length30.130.231.131.534.3External buccal tube width5.96.15.86.17.3 pt 19.6 20.218.6 19.4 21.3 pt stylet supports insertion point 77.7 77.976.9 77.2 77.9First macroplacoid3.94.84.44.4? pt 13.0 15.914.1 14.0 ?Second macroplacoid3.13.43.43.6? pt 10.3 11.310.9 11.4 ?Third macroplacoid4.44.34.54.5? pt 14.6 14.214.5 14.3 ?Microplacoid2.42.32.42.5? pt 8.0 7.67.7 7.9 ?Placoid row16.116.017.417.7? pt 53.5 53.055.9 56.2 ?Macroplacoid row12.512.513.413.2? pt 41.5 41.443.1 41.9 ?External claw I8.2?8.78.6? pt 27.2 ?28.0 27.3 ?Internal claw I7.7?8.58.18.7 pt 25.6 ?27.3 25.7 25.4External claw II8.3?8.99.2? pt 27.6 ?28.6 29.2 ?Internal claw II7.7??8.2? pt 25.6 ?? 26.0 ?External claw III8.3?8.99.2? pt 27.6 ?28.6 29.2 ?Internal claw III7.7??8.2? pt 25.6 ?? 26.0 ?Posterior claw IV???9.7? pt ? ?? 30.8 ?Anterior claw IV???8.99.7 pt???28.328.3 Specific diagnosis. Colourless; eye spots absent (also in living specimens), cuticle smooth. Bucco-pharyngeal apparatus of the Macrobiotus type similar to that of Paramacrobiotus richtersi; three macroplacoids and a microplacoid (at a distance from the third macroplacoid) present. Well developed claws of the hufelandi type with accessory points on the main branches and lunules present. Eggs, free laid, with reticulated trunco-conical processes with short terminal thorns; egg shell areolated with clearly sculptured areolae. Description of the holotype. Body length 270 ��m long, colourless; cuticle smooth without pores (only very small dots are present on the legs); eye spots absent; bucco-pharyngeal apparatus of Macrobiotus type (buccal tube rigid with ventral lamina) (Fig. 5 A), mouth terminal with 10 peribuccal lamellae; buccal armature composed of an anterior band of small teeth, a posterior ring of small triangular teeth and, more caudally, three dorsal and three ventral transverse ridges. Rigid buccal tube 31.5 ��m long and 6.1 ��m wide externally (pt = 19.4) (Table 3); stylet supports inserted on the buccal tube at 77.2 % of its length (pt = 77.2). Pharyngeal bulb with apophyses, three macroplacoids, of almost the same length, and the microplacoid set at a distance from the third macroplacoid, which is a character of the Paramacrobiotus richtersi group (Fig. 5 A); first macroplacoid 4.4 ��m long (pt = 14.0); second 3.6 ��m (pt = 11.4); third 4.5 ��m (pt = 14.3); small microplacoid 2.5 ��m long (pt = 7.9); the entire placoid row 17.7 ��m long (pt = 56.2); the macroplacoid row length 13.2 ��m (pt = 41.9). The first macroplacoid does not appear to have a central constriction. Claws of the Macrobiotus hufelandi type, well developed, slender, with a long basal portion (Fig. 5 B,C). External and internal claws of the first pair of legs 8.6 ��m long (pt = 27.3) and 8.1 ��m long (pt = 25.7) respectively; those of the second and third pairs of legs 9.2 ��m long (pt = 29.2) and 8.2 ��m (pt =26.0) long respectively (Table 3); posterior and anterior claws of the hind legs 9.7 ��m long (pt = 30.8) and 8.9 ��m (pt = 28.3), respectively. Main branches of all claws with thin accessory points (Fig. 5 B, arrow). Smooth lunules present, well developed on the hind legs (Fig. 5 C, arrows); a cuticular bar is present near the base of the claws on the first three pairs of legs. Remarks. The paratypes are similar to the holotype in both morphological and metric characters; the measurements of some structures of the holotype and four paratypes are given in Table 3. Eggs: The eggs, free laid, are areolated and have trunco-conical processes (10 around the circumference, 21-22 per hemisphere) (Fig. 5 D). The diameter is 82-88 ��m excluding the processes, 105-111 ��m including these structures, which are 11.8-12.3 ��m high with a basal diameter ca. 20 ��m or slightly wider (Table 4). Short spines are more or less visible on the apices of the processes (Fig. 5 D). The processes have a reticular ornamentation (Fig. 5 E) with an elongated mesh (with slightly sinuous margins) near the base, and an isodiametric mesh near the apices; the areolae are sculptured with obvious small tubercles (Fig. 5 F). . Etymology. The name pius (meaning: holy, devoutly religious) refers to the fact that the population was found in a moss sample collected in the neighbourhood of the ancient former monastery of Gangivecchio (Gangi, Palermo). Differential diagnosis. As is known, the species of the Paramacrobiotus richtersi group are all very similar and it is often very difficult to identify them without eggs. Paramacrobiotus pius sp. nov. has a smooth cuticle (only very small dots are present on the legs) and therefore differs from Paramacrobiotus corgatensis (which has very fine dots over dorsal and ventral cuticle) and Paramacrobiotus danielisae (which has a sculptured cuticle, with small polygons); stylet supports are inserted on the buccal tube in a slightly more cephalic position (pt = 76.9-77.7 in Paramacrobiotus pius sp. nov., 80.2-80.7 in Paramacrobiotus corgatensis, 80.2-81.2 in Paramacrobiotus danielisae); higher values of the pt index relative to the claw length (Table 5), and in having conical and shorter egg processes (Figs. 5 D and 6 A and 6 B). It also differs from Paramacrobiotus corgatensis in the lack of eye spots. The new species differs from Paramacrobiotus halei by having a smooth cuticle, stylet supports inserted on the buccal tube in a slightly more cephalic position (pt = 76.9-77.7 in Paramacrobiotus pius sp. nov., 81.5 in Paramacrobiotus halei) and higher values of the pt index relative to the claw lengths (Table 5). The new species differs from Paramacrobiotus savai, Paramacrobiotus danielae and Paramacrobiotus peteri by having higher values of the pt index relative to the claw lengths (Tables 5 and 6), and by having sculptured egg areolae. In addition, it differs from Paramacrobiotus savai and Paramacrobiotus danielae by lacking eye spots, from Paramacrobiotus danielae by having a smooth cuticle and stylet supports inserted on the buccal tube in a slightly more cephalic position (pt = 76.9-77.7 in Paramacrobiotus pius sp. nov., 79.9-80.5 in Paramacrobiotus danielae); it differs from Paramacrobiotus peteri also in the shape of the egg processes (Figs. 5 D and 6 C). The lack of eye spots distinguishes Paramacrobiotus pius sp. nov. from Paramacrobiotus priviterae and Paramacrobiotus sklodowskae; in addition, it differs from the former species by the claw shape (Figs. 5 B,C and 6 D), and from Paramacrobiotus sklodowskae by a lower value of the pt index relative to the stylet supports insertion point (76.9-77.7 in Paramacrobiotus pius sp. nov., 81.8-85.2 in Paramacrobiotus sklodovskae). The new species differs from Paramacrobiotus magdalenae by lacking eye spots, by having higher values of the pt index relative to the claw lengths (Table 6), and by characters of the eggs (shape of processes and shorter processes). * According to Michalczyk & Kaczmarek (2006). Slide deposited in the Natural History Collections, Faculty of Biology, A. Mickiewicz University, Poznan, Poland. ** Slide deposited in Te Papa Tongarewa, National Museum of New Zealand, Wellington. ***According to Kaczmarek, Michalczyk and Diduszko, 2005. Paramacrobiotus pius sp. nov. differs from Paramacrobiotus alekseevi by higher values of the pt index relative to the microplacoid length (7.7 ���8.0 in Paramacrobiotus pius sp. nov., 4.0��� 6.2 in Paramacrobiotus alekseevi), different shape of the egg processes, and egg areolae with more obvious ornamentation. The new species differs from Paramacrobiotus hapukuensis by lower values of the pt index relative to macroplacoid lengths but higher values relative to the microplacoid length (Table 6); slightly higher values of that index relative to the claw lengths (Table 6); egg processes shorter and different in shape, and sculptured egg areolae (Figs. 5 D and 7 A). Paramacrobiotus pius sp. nov. is very similar to Paramacrobiotus gerlachae but differs by slightly higher values of the pt index relative to the external claws and a more marked difference in length between external and internal claws on each leg (Table 6); and overall, in some characters of the egg (small apical spines present, egg areolae with obvious ornamentation, which in comparison is very faint in Paramacrobiotus gerlachae where the central portion of the areolae is thicker than the periphery) (Figs. 5 F and 7 B). The new species differs from Paramacrobiotus vanescens by having smooth cuticle with small dots only on the legs (the original description of Paramacrobiotus vanescens did not report that the whole cuticle was finely dotted (Pilato, Binda & Catanzaro, 1991); however, this characteristic was subsequently stressed by Pilato et al. (2001). In addition, the new species differs from Paramacrobiotus vanescens by having lower values of the pt index relative to the microplacoid (7.7 ���8.0 in Paramacrobiotus pius sp. nov., and ca. 4 in Paramacrobiotus vanescens: Figs. 5 A and 7 C), and in details of the eggs (less indented apices of processes, and areolae with more obvious sculpture: Figs. 5 D-F and 7 D). Paramacrobiotus pius sp. nov. differs from Paramacrobiotus garynahi by lacking cuticular pores; by having slightly higher values of the pt index relative to the claw lengths (Table 6); a different shape of the egg processes, and egg areolae with a more obvious sculpture. The new species differs from Paramacrobiotus richtersi overall by having slightly lower values of the index pt relative to the stylet supports insertion point (76.9-77.7 in Paramacrobiotus pius sp. nov., 79.2-81.8 in Paramacrobiotus richtersi), and in some characters of the eggs: the egg processes of the new species have small terminal spines (Fig. 5 D) whereas in Paramacrobiotus richtersi they have blunt apices; and the entire surface of the areolae has an obvious reticular sculpture (Fig. 5 F). Paramacrobiotus pius sp. nov. differs from Paramacrobiotus lorenae by a smooth cuticle, lower value of the pt index relative to the stylet supports insertion point (76.9-77.7 in Paramacrobiotus pius sp. nov. and ca. 81 in Paramacrobiotus lorenae), and in some characters of the eggs (processes with neither elongate nor flexible apices, more evident areolae sculpture)., Published as part of Lisi, Oscar, Binda, Maria Grazia & Pilato, Giovanni, 2016, Eremobiotus ginevrae sp. nov. and Paramacrobiotus pius sp. nov., two new species of Eutardigrada, pp. 344-360 in Zootaxa 4103 (4) on pages 351-359, DOI: 10.11646/zootaxa.4103.4.3, http://zenodo.org/record/267185, {"references":["Kaczmarek, L., Michalczyk, L. & Diduszko, D. (2005) Some tardigrades from Siberia with a description of Macrobiotus garynhai sp. nov. (Eutardigrada: Macrobiotidae, richtersi group). Zootaxa, 1053, 35 - 45.","Pilato, G., Binda, M. G. & Catanzaro R. (1991) Remarks on some tardigrades of the African fauna with the description of three new species of Macrobiotus Schultze, 1834. Tropical Zoology, 4, 167 - 178."]}

Up to few years ago, the phylogenies of tardigrade taxa have been investigated using morphological data, but relationships within and between many taxa are still unresolved. Our aim has been to verify those relationships adding molecular analysis to morphological analysis, using nearly complete 18S ribosomal DNA gene sequences (five new) of 19 species, as well as cytochrome oxidase subunit 1 (COI) mitochondrial DNA gene sequences (15 new) from 20 species, from a total of seven families. The 18S rDNA tree was calculated by minimum evolution, maximum parsimony (MP) and maximum likelihood (ML) analyses. DNA sequences coding for COI were translated to amino acid sequences and a tree was also calculated by neighbour-joining, MP and ML analyses. For both trees (18S rDNA and COI) posterior probabilities were calculated by MrBayes. Prominent findings are as follows: the molecular data on Echiniscidae (Heterotardigrada) are in line with the phylogenetic relationships identifiable by morphological analysis. Among Eutardigrada, orders Apochela and Parachela are confirmed as sister groups. Ramazzottius (Hypsibiidae) results more related to Macrobiotidae than to the genera here considered of Hypsibiidae. Macrobiotidae and Macrobiotus result not monophyletic and confirm morphological data on the presence of at least two large groups within Macrobiotus. Using 18S rDNA and COI mtDNA genes, a new phylogenetic line has been identified within Macrobiotus, corresponding to the ‘richtersi-areolatus group’. Moreover, cryptic species have been identified within the Macrobiotus‘richtersi group’ and within Richtersius. Some evolutionary lines of tardigrades are confirmed, but others suggest taxonomic revision. In particular, the new genus Paramacrobiotus gen. n. has been identified, corresponding to the phylogenetic line represented by the ‘richtersi-areolatus group’. Zusammenfassung Die Anzahl der Arten im Phylum Tardigrada ist in den letzten 25 Jahren von 500 Arten auf inzwischen fast 1000 Arten angestiegen. Zurzeit besteht die Gruppe aus zwei Klassen (Heterotardigrada und Eutardigrada), vier Ordnungen, 21 Familien, und 104 Gattungen. Trotz der Haufigkeit der Tardigraden wurde ihnen seit ihrer Entdeckung im Jahr 1773 nur wenig Aufmerksamkeit geschenkt. Bis vor wenigen Jahren wurden ausschlieslich morphologische Merkmale verwendet, um die Phylogenie der Tardigrada zu untersuchen. Dennoch sind die Verhaltnisse zwischen und innerhalb vieler Arten noch nicht eindeutig geklart. Das Ziel der vorliegenden Arbeit war es, die bereits bekannten, morphologischen Verhaltnisse mit molekularen Ergebnissen zu belegen. Hierzu wurden nur vollstandige Sequenzen der ribosomalen 18S rDNA von 19 Arten verwendet. Funf neue Sequenzen wurden dabei hinzugefugt. Weiterhin wurden von 15 Arten neue mitochrondriale COI Sequenzen verwendet, die mit funf bekannten COI Sequenzen zu insgesamt sieben Familien gehoren. Der 18S rDNA-Baum wurde durch ME, maximum parsimony (MP) and ML Analysen berechnet. Die fur COI kodierenden Sequenzen wurden in Aminosauren ubersetzt und der Baum mit NJ, MP and ML Analysen berechnet. Fur beide Baume (18 rDNA und COI) wurden die Wahrscheinlichkeiten durch MrBayes ermittelt. Dabei ergab sich, dass molekulare Daten mit den morphologischen Untersuchungen bei den Echiniscidae (Heterotardigrada) ubereinstimmen. Bei Eutardigrada wurden die Ordnungen Apochela und Parachela als Schwestergruppen bestatigt. Ramazzottius (Hypsibiidae) gehort zu der Familie Macrobiotidae und weniger zu Hypsibiidae, zu der die Gattung gegenwartig gestellt wird. Die molekularen und morphologischen Daten deuten darauf hin, dass es mindestens zwei groser Gruppen innerhalb von Macrobiotus gibt. Durch die 18 rDNA und COI mtDNA Sequenzen konnte eine neue phylogenetische Linie innerhalb von Macrobiotus, der ‘richtersi-areolatus Gruppe’ zugehorig, identifiziert werden. Weiterhin sind kryptische Arten innerhalb der Macrobiotus richtersi Gruppe’ und innerhalb von Richtersius gefunden worden. Die vorliegende Arbeit verifiziert die in vorangegangene Untersuchungen erarbeitete Phylogenie von Tardigraden. Es konnten einige Entwicklungslinien innerhalb den Tardigraden bestatigt werden, andere deuten zukunftige, taxonomische Revisionen an. So wurde die neue Gattung Paramacrobiotus eingefuhrt, entsprechend der phylogenetischen Linie, die bisher durch die ‘richtersi-areolatus Gruppe’ vertreten war.

207. Paramacrobiotus danielae (Pilato, Binda, Napolitano & Moncada, 2001) [T] Macrobiotus danielae n. sp. (Pilato et al. 2001) Macrobiotus danielae Pilato, Binda, Napolitano & Moncada, 2001 (Pilato et al. 2002, 2003a, 2004) Terra typica: Peru (South America) Ecuador: ��� 00�� 54 ���N, 78 �� 33 ���W; 800 m asl: Esmeraldas Province, Alto Tambo, liverwort (Plagiochila punctata). Pilato et al. (2004) ��� 00�� 11 ���S, 78 �� 36 ���W; 4,500 m asl: Pichincha Province, Pichincha Volcano, soil sample. Pilato et al. (2004) ��� 00�� 22 ���S, 78 �� 55 ���W; 1,100 m asl: Cotopaxi Province, Otonga (San Pedro de las Pampas) [Otonga Nature Reserve], liverwort (Plagiochila punctata and Bazzania bearsonii). Pilato et al. (2002) ��� 00�� 22 ���S, 78 ��09���W; 4,000 m asl: Napo Province, Papallacta Pass, moss (Leptodontium viticulosoides). Pilato et al. (2003 a) ��� 00�� 41 ���S, 77 �� 18 ���W; 400 m asl: Type Locality: Orellana Province, Loreto, moss. Pilato et al. (2001) ��� 00�� 58 ���S, 77 �� 45 ���W; 550 m asl: Napo Province, near Rio Hollin, liverwort (Porella sp.). Pilato et al. (2003 a) Peru: ��� 12 �� 35 ���S, 69 �� 11 ���W; 200 m asl: Region Madre de Dios, Puerto Maldonado, outskirts of Rio Madre de Dios, moss (Pseudocrossidium replicatum). Pilato et al. (2001) Record numbers: Ecuador: 6, Peru: 1; total: 7. Remarks: Currently endemic to South America, this species belongs to the Paramacrobiotus richtersi group (see: P. areolatus above and P. richtersi below)., Published as part of Kaczmarek, ��ukasz, Michalczyk, ��ukasz & Mcinnes, Sandra J., 2015, Annotated zoogeography of non-marine Tardigrada. Part II: South America, pp. 1-107 in Zootaxa 3923 (1) on page 85, DOI: 10.11646/zootaxa.3923.1.1, http://zenodo.org/record/241936, {"references":["Pilato, G., Binda, M. G., Napolitano, A. & Moncada, E. (2001) Notes on South American tardigrades with the description of two new species: Pseudechiniscus spinerectus and Macrobiotus danielae. Tropical Zoology, 14, 223 - 231. http: // dx. doi. org / 10.1080 / 03946975.2001.10531154","Pilato, G., Binda, M. G., Napolitano, A. & Moncada, E. (2002) Tardigrades from Ecuador, with the description of two new species: Mixibius ornatus n. sp. and Diphascon (Adropion) onorei n. sp. (Eutardigrada, Hypsibiidae). Studies on Neotropical Fauna and Environment, 37, 175 - 179. http: // dx. doi. org / 10.1076 / snfe. 37.2.175.8584","Pilato, G., Binda, M. G. & Lisi, O. (2003 a) Remarks on some species of tardigrades from South America with description of Minibiotus sidereus n. sp. Zootaxa, 195, 1 - 8.","Pilato, G., Binda, M. G., Napolitano, A. & Moncada, E. (2004) Remarks on some species of tardigrades from South America with description of two new species. Journal of Natural History, 38, 1081 - 1086. http: // dx. doi. org / 10.1080 / 0022293031000071541"]}

211. Paramacrobiotus rioplatensis (Claps & Rossi, 1997) [T] Macrobiotus rioplatensis sp. n. (Claps & Rossi 1997) Terra typica: Uruguay (South America) Uruguay: • 34 ° 25 ′S, 57 ° 44 ′W; 50 m asl: Type Locality: Colonia Department, Riachuelo, 223 km W of Montevideo, lichens on tree on the banks of the Rio de la Plata. Claps & Rossi (1997) Record numbers: Uruguay: 1; total: 1. Remarks: Currently endemic to Colombia, this species belongs to the Paramacrobiotus areolatus group (see: P. areolatus and P. richtersi above).

210. Paramacrobiotus richtersi (Murray, 1911) sensu lato [T] M. richtersii J. Murr. (Marcus 1939) Macrobiotus richtersii J. Murr., 1908 (de Barros 1942 b) M. richtersi J. Murray, 1911 (du Bois-Reymond Marcus 1944) Macrobiotus richtersii J. Murr. (Iharos 1963) Macrobiotus richtersi Murr. (Iharos 1969) Macrobiotus richtersi Murray, 1911 (Rossi & Claps 1980) M. richtersi Murray, 1911 (Claps & Rossi 1981, 1984, 1988) Macrobiotus richtersi J. Murray, 1911 (Iharos 1982) Macrobiotus richtersi Murray, 1911 (Maucci 1988, Rossi & Claps 1991, Jerez & Narváez 2001) M. richtersi J. Murr., 1911 (Rossi & Claps 1989) M. richtersi Murray (Claps & Rossi 1997) Macrobiotus harmsworthi / richtersi (Montoya et al. 2010) Terra typica: Ireland (Europe) Argentina: • 23 °08′S, 64 ° 19 ′W; 350 m asl: Salta Province, Orán [San Ramón de la Nueva Orán], mosses on rock. Claps & Rossi (1984) • 23 ° 36 ′S, 65 °04′W; 2,000 m asl: Jujuy Province, Departamento Santa Bárbara, lichens on vina lito. Claps & Rossi (1984) • 24 °07′S, 65 ° 24 ′W; 1,450 m asl: Jujuy Province, road to Yala, mosses from the edge of irrigation ditch and on tree (2 samples). Claps & Rossi (1984) • 24 ° 18 ′S, 64 ° 54 ′W; 700 m asl: Jujuy Province, road to Higueritas, mosses on soil. Claps & Rossi (1984) • 25 ° 43 ′S, 54 ° 25 ′W; 250 m asl: Misiones Province, Iguazú National Park, moss on rock and mosses on trees in the forest (3 samples). Claps & Rossi (1988) • 25 ° 58 ′S, 54 ° 34 ′W; 200 m asl: Misiones Province, Wanda, mosses on rocks. Claps & Rossi (1988) • 26 °08′S, 54 ° 35 ′W; 150 m asl: Misiones Province, Arroyo Torocuá, 12 km S Puerto Esperanza, mosses on rocks (2 samples). Claps & Rossi (1988) • 26 ° 47 ′S, 65 ° 20 ′W; 750 m asl: Tucumán Province, Horco Molle, mosses on soil and tree (2 samples). Claps & Rossi (1984) • 26 ° 51 ′S, 65 ° 26 ′W; 700 m asl: Tucumán Province, road from San Miguel de Tucumán to Tafi del Valle, mosses and lichens on rocks (2 samples). Claps & Rossi (1984) • 26 ° 56 ′S, 55 °04′W; 200 m asl: Misiones Province, Capioví, mosses on trees in the forest (2 samples). Claps & Rossi (1988) • 27 ° 27 ′S, 58 ° 39 ′W; 50 m asl: Corrientes Province, old road to Santa Ana, Selaginella sp. in lagoon and moss on tree (2 samples). Claps & Rossi (1988) • 35 ° 27 ′S, 58 ° 48 ′W; 0 m asl: Buenos Aires Province, Laguna San Miguel del Monte, on plant (Ceratophyllum sp.) Rossi & Claps (1991). • 36 ° 22 ′S, 56 ° 43 ′W; 0 m asl: Buenos Aires Province, camping San Clemente del Tuy, lichens on tree (Eucalyptus) in shade. Rossi & Claps (1980) • 36 ° 24 ′S, 56 ° 57 ′W; 0 m asl: Buenos Aires Province, Port General Lavalle, lichens on tree (Tamarisk) in full sun. Rossi & Claps (1980) • 40 ° 50 ′S, 71 ° 38 ′W; 900 m asl: Neuquén Province, Peninsula de Quetrihue, semi-wet, mosses on rocks on the sunny slope. Claps & Rossi (1981) • 41 °00′S, 71 ° 30 ′W; 800 m asl: Neuquén Province, Isla Victoria, mosses on tree. Rossi & Claps (1989) • 41 ° 13 ′S, 71 ° 30 ′W; 2,250 m asl: Rio Negro Province, Nahuel Huapi National Park, Cerro Catedral, semi-wet, lichens and mosses on trees and roots (Nothofagus sp.) in the forest, very wet, mosses on soil and dead wood and mosses on leaf litter in the forest (6 samples). Claps & Rossi (1981) • 41 ° 58 ′S, 71 ° 29 ′W; 1,200 m asl: Rio Negro Province, El Bolsón, Piltriquitron Mt., mosses on dry soil under Mulinum spinosum and between Berberis shrubs. Iharos (1963) • 41 ° 58 ′S, 71 ° 28 ′W; 1,460 m asl: Rio Negro Province, El Bolsón, Piltriquitron Mt., mosses on rocks and barks of trees near edge within Nothofagus pumilio forest. Iharos (1963) • 41 ° 59 ′S, 71 ° 34 ′W; 300 m asl: Rio Negro Province, El Bolsón, Valley of Rio Azul, mosses on tree (Myrceugenia exupca) in the Myrceugenia exupca - Nothofagus dombeyi marsh forest. Iharos (1963) • 41 ° 59 ′S, 71 ° 31 ′W; 360 m asl: Rio Negro Province, El Bolsón, foot of Piltriquitron Mt., mosses on rocks in shaded Libocedrus chilensis -Lomatia obliqua forest. Iharos (1963) • 41 ° 59 ′S, 71 ° 31 ′W; 370 m asl: Rio Negro Province, El Bolsón, foot of Piltriquitron Mt., slope above Pampa Azcona, mosses on soil of shrubby area. Iharos (1963) • 42 ° 13 ′S, 71 ° 42 ′W; 220 m asl: Chubut Province, El Turbio, Puelo Lake, mosses on wet soil in primeval Nothofagus dombeyi - Libocedrus chilensis forest near the lake. Iharos (1963) • 54 ° 39 ′S, 68 ° 30 ′W; 800 m asl: Tierra del Fuego Province, National Park Ushuaia [Tierra del Fuego National Park], soil and litter (2 samples). Iharos (1982) Bolivia: • 10 ° 49 ′S, 65 ° 22 ′W; 150 m asl: Beni Department, Guayaramerín, Estancia Esperanza, gallerian forest along Mamore river, leaf litter, leaf litter near water, decayed leaves on banana plantation, dry leaf litter, leaf litter from cacao plantation and soil from Leguminosae plantation (7 samples). Iharos (1969) • 10 ° 56 ′S, 65 ° 28 ′W; 150 m asl: Beni Department, Guayaramerín, 15 km on W on the city, leaf litter in evergreen forest, near the road to Riberalta, leaf litter (2 samples). Iharos (1969) • 15 ° 31 ′S, 67 ° 33 ′W; 800 m asl: La Paz Department, between Alcoche and Puerto Linares, 20 km on Alcoche, mosses in shaded place. Iharos (1969) • 16 °08′S, 67 ° 43 ′W; 1,600 m asl: La Paz Department, Coroico, 5 km on N of the city, wet leaf litter from citrus and coffee plantation. Iharos (1969) • 16 ° 11 ′S, 69 °05′W; 4,100 m asl: La Paz Department, Copacabana [on the coast of Titicaca Lake], leaf litter from Stipa vegetation. Iharos (1969) • 16 ° 16 ′S, 67 ° 53 ′W; 3,100 m asl: La Paz Department, 5 km on N of Unduavi, leaf litter and soil from Vaccinum plantation. Iharos (1969) • 16 ° 19 ′S, 67 ° 54 ′W; 4,000 m asl: La Paz Department, near Unduavi, near the river, very wet leaf litter under shrubs. Iharos (1969) Brazil: • 03° 17 ′S, 59 ° 53 ′W; 0 m asl: Amazonas State, Manaus, 20 km on S of the city, leaf litter from evergreen forest. Iharos (1969) • 21 ° 49 ′S, 49 °05′W: Undefined localities in São Paulo State, few localities. Marcus (1939), de Barros (1942 b) • 22 ° 44 ′S, 45 ° 35 ′W; 1,650 m asl: São Paulo State, Campos do Jordão, mosses or aquatic plants. du Bois-Reymond Marcus (1944) • 22 ° 45 ′S, 47 ° 24 ′W; 550 m asl: São Paulo State, Município de Sta. Bárbara [Município Santa Bárbara d ′Oeste], mosses or aquatic plants. du Bois-Reymond Marcus (1944) • 22 ° 54 ′S, 47 °04′W; 700 m asl: São Paulo State, Município Campinas, in bromeliads. du Bois-Reymond Marcus (1944) • 25 ° 26 ′S, 49 ° 16 ′W; 900 m asl: Paraná State, Curitiba, mosses or aquatic plants. du Bois-Reymond Marcus (1944) Chile: • 18 ° 11 ′S, 69 ° 13 ′W; 4,800 m asl [4,550 m asl]: Region XV Arica y Parinacota, Laguna La Cotacotani, leaf litter. Iharos (1969) • 18 ° 34 ′S, 70 °05′W; 500 m asl: Region XV Arica y Parinacota, Azapa Chico, 25 km. on East of Arica, near the river Rio Lauca, wet soil on bamboo plantation. Iharos (1969) • 20 ° 12 ′S, 69 ° 17 ′W: Undefined locality in Region I Tarapac, Bofedal de Bajuco, soil under red cactus. Iharos (1969) • 30 ° 17 ′S, 71 ° 16 ′W; 400 m asl: Region IV Coquimbo, Las Cardas, leaf litter under shrubs. Iharos (1969) • 30 ° 40 ′S, 71 ° 41 ′W; 900 m asl [650 m asl]: Region IV Coquimbo, Fray Jorge [Bosque de Fray Jorge National Park], foggy forest, leaf litter. Iharos (1969) • 31 ° 55 ′S, 71 ° 31 ′W; 0 m asl: Region IV Coquimbo, Los Vilos, jungle soil and leaf litter. Iharos (1969) • 32 ° 58 ′S, 71 °01′W; 1,900 m asl: Region V Valparaíso (Región de Valparaíso), Cordiliera de la Costa, Cerro El Roble, 84 km NW of Santiago de Chile, leaf litter under snow, under the bushes. Iharos (1969) • 32 ° 58 ′S, 71 °01′W; 1,700 m asl: Region V Valparaíso (Región de Valparaíso), Cordiliera de la Costa, Cerro El Roble, 84 km NW of Santiago de Chile, leaf litter between dry bushes. Iharos (1969) • 33 °04′S, 71 °00′W; 1,000 m asl: Region RM Metropolitana (Región Metropolitana de Santiago), Tiltil, Cuesta La Dormida, jungle leaf litter. Iharos (1969) • 33 ° 21 ′S, 70 ° 19 ′W; 2,300 m asl: Region RM Metropolitana (Región Metropolitana de Santiago), Farellones, 30 km on East of Santiago de Chile, detritus and soil under the bushes, wet mosses from rocky cave and leaf litter (3 samples). Iharos (1969) • 33 ° 26 ′S, 70 ° 38 ′W; 800 m asl: Region RM Metropolitana (Región Metropolitana de Santiago), Santiago de Chile, San Cristóbal Mt., mosses on rocks. Iharos (1969) • 33 ° 30 ′S, 70 ° 55 ′W; 850 m asl: Region RM Metropolitana (Región Metropolitana de Santiago), Quebrada de La Plata, shrubs upland, moss and lichen on rocks (2 samples). Iharos (1969) • 37 ° 13 ′S, 72 ° 23 ′W; 100 m asl: Region VIII Biobío, Salto del Laja, near waterfalls, mosses on rocky wall. Iharos (1969) • 50 ° 59 ′S, 73 °00′W; 1,200 m asl: Region XII Magallanes (Región de Magallanes y de la Antártica Chilena), Torres del Paine National Park, mosses on trees and rocks, nearly all in half sunlight. Maucci (1988) • 51 ° 34 ′S, 72 ° 36 ′W; 200 m asl: Region XII Magallanes (Región de Magallanes y de la Antártica Chilena), Ultima Esperanza, near Cueva del Milodón, mosses on rocks, in sunlight. Maucci (1988) Colombia: • 07°07′N, 73 °02′W; 1,795–1,970 m asl: Santander Department, El Diviso Natural Reserve, Pine forest (Pinus patula), mosses (Hypopterygium tamariscinum, Fabronia ciliaris and Sematophyllum insularum). Jerez & Narváez (2001) Uruguay: • 30 ° 15 ′S, 57 ° 36 ′W; 50 m asl: Artigas Department, Bella Unión, mosses or lichens on rocks, trees or posts or aquatic macrophytes. Claps & Rossi (1997) • 31 °03′S, 57 ° 38 ′W; 50 m asl: Salto Department, Arroyo Palomas, mosses or lichens on rocks, trees or posts or aquatic macrophytes. Claps & Rossi (1997) • 31 ° 20 ′S, 57 ° 51 ′W; 50 m asl: Salto Department, Arroyo San Antônio de Chico, mosses or lichens on rocks, trees or posts or aquatic macrophytes. Claps & Rossi (1997) Venezuela: • 08° 46 ′N, 70 ° 49 ′W; 4,300 m asl: Region Los Andes, Mérida, Mucuñuque Mt., moss. Montoya et al. (2010) • 08° 45 ′N, 70 ° 49 ′W; 4,450 m asl: Region Los Andes, Mérida, Mucuñuque Mt., surface samples of peat bog. Montoya et al. (2010) Record numbers: Argentina: 24, Bolivia: 7, Brazil: 6, Chile: 15, Colombia: 1, Uruguay: 3, Venezuela: 2; total: 58. Remarks: Paramacrobiotus richtersi was, until recently, believed to be cosmopolitan (McInnes 1994 a). New taxonomic data has shown that this is the nominal species for a species complex – the ‘ richtersi group’ – requiring careful examination of both adults and eggs. Several newly described species of this group have been added to the faunas of South and Central America (e.g. Michalczyk & Kaczmarek 2006 b, Michalczyk et al. 2006, Pilato et al. 2006). In the light of these reports, and given P. richtersi was originally described from Europe, we suggest all species records from South America should be treated with caution until the original slides are re-examined or the species is otherwise verified from the region.

205. Paramacrobiotus areolatus (Murray, 1907 a) sensu lato [T] Makrobiotus aff. areolatus Murray (Richters 1911 a) Macrobiotus areolatus Murray (Murray & Wailes 1913) M. areolatus Murr. (Iharos 1969) Macrobiotus areolatus Murray, 1907 (Rossi & Claps 1980, Claps & Rossi 1984, Maucci 1988, Jerez & Narváez 2001, Nickel et al. 2001, Moly de Peluffo et al. 2006, Peluffo et al. 2007) M. areolatus Murray, 1907 (Claps & Rossi 1988) Macrobiotus areolatus J. Murr., 1907 (Rossi & Claps 1989) M. areolatus Murray (Claps & Rossi 1997) Terra typica: Svalbard (Spitsbergen) (Norway, Arctic Ocean) Argentina: • 24 ° 44 ′S, 65 ° 29 ′W; 1,350 m asl: Salta Province, San Lorenzo, lichens on wood on soil in the mountains. Claps & Rossi (1984) • 24 ° 45 ′S, 65 ° 28 ′W; 1,300 m asl: Salta Province, road from Salta to San Lorenzo, lichens on trees (2 samples). Claps & Rossi (1984) • 25 ° 38 ′S, 65 ° 39 ′W; 1,250 m asl: Salta Province, road from Cafayate to Salta, mosses on tree and mosses on tree in the mountains (2 samples). Claps & Rossi (1984) • 27 ° 57 ′S, 58 ° 48 ′W; 50 m asl: Corrientes Province, Empedrado, Paraje Sombrerito, mosses on fallen tree. Claps & Rossi (1988) • 35 ° 40 ′S, 63 ° 44 ′W; 143 m asl: La Pampa Province, General Pico, mosses and lichens on trees. Moly de Peluffo et al. (2006) • 36 ° 22 ′S, 56 ° 43 ′W; 0 m asl: Buenos Aires Province, San Clemente del Tuy, lichens on tree (Eucalyptus) in full sun, lichens on tree (Eucalyptus) in shade, lichens on tree (Tamarisk) in near the forest and lichens on tree (Tamarisk) in full sun (4 samples). Rossi & Claps (1980) • 36 ° 24 ′S, 56 ° 57 ′W; 0 m asl: Buenos Aires Province, Port General Lavalle, lichens on tree (Tamarisk) in full sun and lichens on tree (Eucalyptus) (3 samples). Rossi & Claps (1980) • 36 ° 39 ′S, 64 ° 17 ′W; 177 m asl: La Pampa Province, Santa Rosa [Santa Rosa de Toay], mosses and lichens on trees (Robinia pseudoacacia, few species of Fraxinus or Prosopis caldenia). Peluffo et al. (2007) • 40 ° 49 ′S, 63 °00′W; 0 m asl: Rio Negro Province, Viedma, mosses on soil. Rossi & Claps (1989) • 50 °06′S, 73 ° 18 ′W; 200 m asl: Santa Cruz Province, Los Glaciares National Park, shores of Argentino Lake, near the Onelli glacier, Nothofagus forests, in the shade, mosses and lichens on trees and rocks. Maucci (1988) Bolivia: • 14 ° 49 ′S, 69 °04′W; 11–12,000 ft asl / 3,350–3,650 m asl: La Paz Department, Pelechuco. Murray & Wailes (1913) Chile: • 33 °04′S, 71 °00′W; 1,000 m asl: Region RM Metropolitana (Región Metropolitana de Santiago), Tiltil, Cuesta La Dormida, soil on dry forest (2 samples). Iharos (1969) • 51 ° 44 ′S, 72 ° 30 ′W; 0 m asl: Region XII Magallanes (Región de Magallanes y de la Antártica Chilena), Puerto Natales, mosses on trees and rocks. Maucci (1988) Colombia: • 07°07′N, 73 °02′W; 970 – 1,850 m asl: Santander Department, El Diviso Natural Reserve, Cypress forest (Cupresus lusitanica) and trail, moss (Lyperodon tomentosus). Jerez & Narváez (2001) Paraguay: • 22 ° 39 ′S, 57 ° 46 ′W; 50 m asl: Concepción Department, Puerto Max, between Concepción and Rio Ata, lichens, mosses and hepatics (Forsstroemia cuspidata and Frullania sp.). Richters (1911 a) • 25 ° 20 ′S, 57 ° 13 ′W; 200 m asl: Cordillera Department, Cordillera de Altos, Tucangua, mosses and lichens on palm tree (Acrocomia sclerocarpa). Richters (1911 a) Peru: • 13 °04′ 48 ′′S, 72 ° 18 ′00′′W; 2,350 m asl [13 ° 10 ′S, 72 ° 33 ′W; 2,500 m asl]: Cusco Region, Machu Picchu, mosses or lichens. Nickel et al. (2001) • 13 ° 12 ′S, 69 ° 38 ′W; 1–3,000 ft asl / 300–900 m asl: Madre de Dios Region, Tambopata Valley. Murray & Wailes (1913) • 13 °08′ 24 ′′S, 72 ° 10 ′ 12 ′′W; 2,300 m asl [13 ° 15 ′S, 72 ° 16 ′W; 2,850 m asl]: Cusco Region, Ollantaytambo, mosses or lichens. Nickel et al. (2001) Uruguay: • 30 ° 15 ′S, 57 ° 36 ′W; 50 m asl: Artigas Department, Bella Unión, mosses or lichens on rocks, trees or posts or aquatic macrophytes. Claps & Rossi (1997) • 30 ° 57 ′S, 57 ° 31 ′W; 50 m asl: Salto Department, Termas del Arapey, mosses or lichens on rocks, trees or posts or aquatic macrophytes. Claps & Rossi (1997) • 31 °03′S, 57 ° 38 ′W; 50 m asl: Salto Department, Arroyo Palomas, mosses or lichens on rocks, trees or posts or aquatic macrophytes. Claps & Rossi (1997) • 32 ° 31 ′S, 55 ° 46 ′W: Undefined locality, Cañada Cousillas, mosses or lichens on rocks, trees or posts or aquatic macrophytes. Claps & Rossi (1997) • 34 ° 10 ′S, 55 ° 49 ′W; 50 m asl: Florida Department, Arroyo San Juan, mosses or lichens on rocks, trees or posts or aquatic macrophytes. Claps & Rossi (1997) • 34 ° 25 ′S, 57 ° 44 ′W; 50 m asl: Colonia Department, Riachuelo, mosses or lichens on rocks, trees or posts or aquatic macrophytes. Claps & Rossi (1997) Record numbers: Argentina: 10, Bolivia: 1, Chile: 2, Colombia: 1, Paraguay: 2, Peru: 3, Uruguay: 6; total: 25. Remarks: The genus Paramacrobiotus (until recently a species group within Macrobiotus) is currently divided into two groups: richtersi and areolatus (with and without macroplacoid, respectively). P. areolatus, once considered cosmopolitan (McInnes 1994 a), is now recognised as the nominal species for a group of very similar taxa that require careful taxonomic examination of adults and egg morphology for correct identification. Recently, new species belonging to the areolatus group have been described from Central and South America (e.g. Michalczyk & Kaczmarek 2006 b, Degma et al. 2008, Kaczmarek et al. 2014 b). In the light of these reports and because P. areolatus was originally described from the Arctic, we feel all records of this species from South America should be treated with caution until relevant examples are re-examined or the presence in the region is otherwise verified.

209. Paramacrobiotus intii Kaczmarek, Cytan, Zawierucha, Diduszko & Michalczyk 2014 [T] Paramacrobiotus intii sp. nov. (Kaczmarek et al. 2014 b) Terra typica: Peru (South America) Peru: ��� 13 �� 25 ���S, 71 �� 51 ���W; 3,000 m asl: Type Locality: Cusco Region, Pisac near Cusco, mixed mosses and lichens on rock. Kaczmarek et al. (2014 b) Record numbers: Peru: 1; total: 1. Remarks: Currently endemic to Peru, this species belongs to the Paramacrobiotus areolatus group (see: P. areolatus above and P. richtersi below)., Published as part of Kaczmarek, ��ukasz, Michalczyk, ��ukasz & Mcinnes, Sandra J., 2015, Annotated zoogeography of non-marine Tardigrada. Part II: South America, pp. 1-107 in Zootaxa 3923 (1) on page 85, DOI: 10.11646/zootaxa.3923.1.1, http://zenodo.org/record/241936, {"references":["Kaczmarek, L., Cytan, J., Zawierucha, K., Diduszko, D. & Michalczyk, L. (2014 b) Tardigrades from Peru (South America), with descriptions of three new species of Parachela. Zootaxa, 3790 (2), 357 - 379. http: // dx. doi. org / 10.11646 / zootaxa. 3790.2.5"]}

207. Paramacrobiotus danielae (Pilato, Binda, Napolitano & Moncada, 2001) [T] Macrobiotus danielae n. sp. (Pilato et al. 2001) Macrobiotus danielae Pilato, Binda, Napolitano & Moncada, 2001 (Pilato et al. 2002, 2003a, 2004) Terra typica: Peru (South America) Ecuador: • 00° 54 ′N, 78 ° 33 ′W; 800 m asl: Esmeraldas Province, Alto Tambo, liverwort (Plagiochila punctata). Pilato et al. (2004) • 00° 11 ′S, 78 ° 36 ′W; 4,500 m asl: Pichincha Province, Pichincha Volcano, soil sample. Pilato et al. (2004) • 00° 22 ′S, 78 ° 55 ′W; 1,100 m asl: Cotopaxi Province, Otonga (San Pedro de las Pampas) [Otonga Nature Reserve], liverwort (Plagiochila punctata and Bazzania bearsonii). Pilato et al. (2002) • 00° 22 ′S, 78 °09′W; 4,000 m asl: Napo Province, Papallacta Pass, moss (Leptodontium viticulosoides). Pilato et al. (2003 a) • 00° 41 ′S, 77 ° 18 ′W; 400 m asl: Type Locality: Orellana Province, Loreto, moss. Pilato et al. (2001) • 00° 58 ′S, 77 ° 45 ′W; 550 m asl: Napo Province, near Rio Hollin, liverwort (Porella sp.). Pilato et al. (2003 a) Peru: • 12 ° 35 ′S, 69 ° 11 ′W; 200 m asl: Region Madre de Dios, Puerto Maldonado, outskirts of Rio Madre de Dios, moss (Pseudocrossidium replicatum). Pilato et al. (2001) Record numbers: Ecuador: 6, Peru: 1; total: 7. Remarks: Currently endemic to South America, this species belongs to the Paramacrobiotus richtersi group (see: P. areolatus above and P. richtersi below).

211. Paramacrobiotus rioplatensis (Claps & Rossi, 1997) [T] Macrobiotus rioplatensis sp. n. (Claps & Rossi 1997) Terra typica: Uruguay (South America) Uruguay: ��� 34 �� 25 ���S, 57 �� 44 ���W; 50 m asl: Type Locality: Colonia Department, Riachuelo, 223 km W of Montevideo, lichens on tree on the banks of the Rio de la Plata. Claps & Rossi (1997) Record numbers: Uruguay: 1; total: 1. Remarks: Currently endemic to Colombia, this species belongs to the Paramacrobiotus areolatus group (see: P. areolatus and P. richtersi above)., Published as part of Kaczmarek, ��ukasz, Michalczyk, ��ukasz & Mcinnes, Sandra J., 2015, Annotated zoogeography of non-marine Tardigrada. Part II: South America, pp. 1-107 in Zootaxa 3923 (1) on page 88, DOI: 10.11646/zootaxa.3923.1.1, http://zenodo.org/record/241936, {"references":["Claps, M. C. & Rossi, G. C. (1997) Tardigrados de Uruguay, com descripcion de dos nuevas especies (Echiniscidae, Macrobiotidae). Iheringia, Serie Zoologia, 83, 17 - 22."]}

205. Paramacrobiotus areolatus (Murray, 1907 a) sensu lato [T] Makrobiotus aff. areolatus Murray (Richters 1911 a) Macrobiotus areolatus Murray (Murray & Wailes 1913) M. areolatus Murr. (Iharos 1969) Macrobiotus areolatus Murray, 1907 (Rossi & Claps 1980, Claps & Rossi 1984, Maucci 1988, Jerez & Narv��ez 2001, Nickel et al. 2001, Moly de Peluffo et al. 2006, Peluffo et al. 2007) M. areolatus Murray, 1907 (Claps & Rossi 1988) Macrobiotus areolatus J. Murr., 1907 (Rossi & Claps 1989) M. areolatus Murray (Claps & Rossi 1997) Terra typica: Svalbard (Spitsbergen) (Norway, Arctic Ocean) Argentina: ��� 24 �� 44 ���S, 65 �� 29 ���W; 1,350 m asl: Salta Province, San Lorenzo, lichens on wood on soil in the mountains. Claps & Rossi (1984) ��� 24 �� 45 ���S, 65 �� 28 ���W; 1,300 m asl: Salta Province, road from Salta to San Lorenzo, lichens on trees (2 samples). Claps & Rossi (1984) ��� 25 �� 38 ���S, 65 �� 39 ���W; 1,250 m asl: Salta Province, road from Cafayate to Salta, mosses on tree and mosses on tree in the mountains (2 samples). Claps & Rossi (1984) ��� 27 �� 57 ���S, 58 �� 48 ���W; 50 m asl: Corrientes Province, Empedrado, Paraje Sombrerito, mosses on fallen tree. Claps & Rossi (1988) ��� 35 �� 40 ���S, 63 �� 44 ���W; 143 m asl: La Pampa Province, General Pico, mosses and lichens on trees. Moly de Peluffo et al. (2006) ��� 36 �� 22 ���S, 56 �� 43 ���W; 0 m asl: Buenos Aires Province, San Clemente del Tuy, lichens on tree (Eucalyptus) in full sun, lichens on tree (Eucalyptus) in shade, lichens on tree (Tamarisk) in near the forest and lichens on tree (Tamarisk) in full sun (4 samples). Rossi & Claps (1980) ��� 36 �� 24 ���S, 56 �� 57 ���W; 0 m asl: Buenos Aires Province, Port General Lavalle, lichens on tree (Tamarisk) in full sun and lichens on tree (Eucalyptus) (3 samples). Rossi & Claps (1980) ��� 36 �� 39 ���S, 64 �� 17 ���W; 177 m asl: La Pampa Province, Santa Rosa [Santa Rosa de Toay], mosses and lichens on trees (Robinia pseudoacacia, few species of Fraxinus or Prosopis caldenia). Peluffo et al. (2007) ��� 40 �� 49 ���S, 63 ��00���W; 0 m asl: Rio Negro Province, Viedma, mosses on soil. Rossi & Claps (1989) ��� 50 ��06���S, 73 �� 18 ���W; 200 m asl: Santa Cruz Province, Los Glaciares National Park, shores of Argentino Lake, near the Onelli glacier, Nothofagus forests, in the shade, mosses and lichens on trees and rocks. Maucci (1988) Bolivia: ��� 14 �� 49 ���S, 69 ��04���W; 11���12,000 ft asl / 3,350���3,650 m asl: La Paz Department, Pelechuco. Murray & Wailes (1913) Chile: ��� 33 ��04���S, 71 ��00���W; 1,000 m asl: Region RM Metropolitana (Regi��n Metropolitana de Santiago), Tiltil, Cuesta La Dormida, soil on dry forest (2 samples). Iharos (1969) ��� 51 �� 44 ���S, 72 �� 30 ���W; 0 m asl: Region XII Magallanes (Regi��n de Magallanes y de la Ant��rtica Chilena), Puerto Natales, mosses on trees and rocks. Maucci (1988) Colombia: ��� 07��07���N, 73 ��02���W; 970 ��� 1,850 m asl: Santander Department, El Diviso Natural Reserve, Cypress forest (Cupresus lusitanica) and trail, moss (Lyperodon tomentosus). Jerez & Narv��ez (2001) Paraguay: ��� 22 �� 39 ���S, 57 �� 46 ���W; 50 m asl: Concepci��n Department, Puerto Max, between Concepci��n and Rio Ata, lichens, mosses and hepatics (Forsstroemia cuspidata and Frullania sp.). Richters (1911 a) ��� 25 �� 20 ���S, 57 �� 13 ���W; 200 m asl: Cordillera Department, Cordillera de Altos, Tucangua, mosses and lichens on palm tree (Acrocomia sclerocarpa). Richters (1911 a) Peru: ��� 13 ��04��� 48 ������S, 72 �� 18 ���00������W; 2,350 m asl [13 �� 10 ���S, 72 �� 33 ���W; 2,500 m asl]: Cusco Region, Machu Picchu, mosses or lichens. Nickel et al. (2001) ��� 13 �� 12 ���S, 69 �� 38 ���W; 1���3,000 ft asl / 300���900 m asl: Madre de Dios Region, Tambopata Valley. Murray & Wailes (1913) ��� 13 ��08��� 24 ������S, 72 �� 10 ��� 12 ������W; 2,300 m asl [13 �� 15 ���S, 72 �� 16 ���W; 2,850 m asl]: Cusco Region, Ollantaytambo, mosses or lichens. Nickel et al. (2001) Uruguay: ��� 30 �� 15 ���S, 57 �� 36 ���W; 50 m asl: Artigas Department, Bella Uni��n, mosses or lichens on rocks, trees or posts or aquatic macrophytes. Claps & Rossi (1997) ��� 30 �� 57 ���S, 57 �� 31 ���W; 50 m asl: Salto Department, Termas del Arapey, mosses or lichens on rocks, trees or posts or aquatic macrophytes. Claps & Rossi (1997) ��� 31 ��03���S, 57 �� 38 ���W; 50 m asl: Salto Department, Arroyo Palomas, mosses or lichens on rocks, trees or posts or aquatic macrophytes. Claps & Rossi (1997) ��� 32 �� 31 ���S, 55 �� 46 ���W: Undefined locality, Ca��ada Cousillas, mosses or lichens on rocks, trees or posts or aquatic macrophytes. Claps & Rossi (1997) ��� 34 �� 10 ���S, 55 �� 49 ���W; 50 m asl: Florida Department, Arroyo San Juan, mosses or lichens on rocks, trees or posts or aquatic macrophytes. Claps & Rossi (1997) ��� 34 �� 25 ���S, 57 �� 44 ���W; 50 m asl: Colonia Department, Riachuelo, mosses or lichens on rocks, trees or posts or aquatic macrophytes. Claps & Rossi (1997) Record numbers: Argentina: 10, Bolivia: 1, Chile: 2, Colombia: 1, Paraguay: 2, Peru: 3, Uruguay: 6; total: 25. Remarks: The genus Paramacrobiotus (until recently a species group within Macrobiotus) is currently divided into two groups: richtersi and areolatus (with and without macroplacoid, respectively). P. areolatus, once considered cosmopolitan (McInnes 1994 a), is now recognised as the nominal species for a group of very similar taxa that require careful taxonomic examination of adults and egg morphology for correct identification. Recently, new species belonging to the areolatus group have been described from Central and South America (e.g. Michalczyk & Kaczmarek 2006 b, Degma et al. 2008, Kaczmarek et al. 2014 b). In the light of these reports and because P. areolatus was originally described from the Arctic, we feel all records of this species from South America should be treated with caution until relevant examples are re-examined or the presence in the region is otherwise verified., Published as part of Kaczmarek, ��ukasz, Michalczyk, ��ukasz & Mcinnes, Sandra J., 2015, Annotated zoogeography of non-marine Tardigrada. Part II: South America, pp. 1-107 in Zootaxa 3923 (1) on pages 83-84, DOI: 10.11646/zootaxa.3923.1.1, http://zenodo.org/record/241936, {"references":["Murray, J. (1907 a) Arctic Tardigrada, collected by Wm. S. Bruce. Transactions of the Royal Society of Edinburgh, 45, 669 - 681. http: // dx. doi. org / 10.1017 / s 0080456800011789","Richters, F. (1911 a) Sudamerikanische Tardigraden. Zoologisher Anzeiger, 38, 273 - 277.","Murray, J. & Wailes, G. H. (1913) Notes on the Natural History of Bolivia and Peru. The Scottish oceanographic laboratory Edinburgh, 1 - 45.","Iharos, G. (1969) The scientific results of the Hungarian soil zoological expeditions to South America: 15. Tardigraden aus den Sammlungen der ersten und zweiten Expedition. Opuscula Zoologica, Budapest, 9, 279 - 289.","Rossi, G. C. and Claps, M. C. (1980) Contribucion al conocimiento de los tardigrados de Argentina. I. Revista de la Sociedad Entomologica Argentina, 39, 243 - 250","Claps, M. C. & Rossi, G. C. (1984) Contribucion al conocimiento de los tardigrados de Argentina. IV. Acta Zoologica Lilloana, 38, 45 - 50.","Maucci, W. (1988) Tardigrada from Patagonia (Southern South America) with description of three new species. Revista Chilena de Entomologia, 16, 5 - 13.","Jerez Jaimes, J. H. & Narvaez Parra, E. X. (2001) Tardigrados (Animalia, tardigrada) de la Reserva El Diviso - Santander, Colombia. Biota Colombiana, 2, 145 - 151.","Nickel, K., Miller, W. R. & Marley, N. (2001) Tardigrades of South America: Machu Picchu and Ollantayambo, Peru. Zoologischer Anzeiger, 240, 505 - 509. http: // dx. doi. org / 10.1078 / 0044 - 5231 - 00060","Peluffo, J. R., Rocha, A. M. & Moly de Peluffo, M. C. (2007) Species diversity and morphometrics of tardigrades in a mediumsized city in the Neotropical Region: Santa Rosa (La Pampa, Argentina). Animal Biodiversity and Conservation, 30, 43 - 51.","Claps, M. C. & Rossi, G. C. (1988) Contribucion al conocimiento de los tardigrados de Argentina. VI. Iheringia, 67, 3 - 11.","Rossi, G. C. & Claps, M. C. (1989) Contribucion al conocimiento de los Tardigrados de la Argentina V. Revista de la Sociedad Entomologica Argentina, 47, 133 - 142.","Claps, M. C. & Rossi, G. C. (1997) Tardigrados de Uruguay, com descripcion de dos nuevas especies (Echiniscidae, Macrobiotidae). Iheringia, Serie Zoologia, 83, 17 - 22.","McInnes, S. J. (1994 a) Zoogeographic distribution of terrestrial / freshwater tardigrades from current literature. Journal of Natural History, 28, 257 - 352. http: // dx. doi. org / 10.1080 / 00222939400770131","Degma, P., Michalczyk, L. & Kaczmarek, L. (2008) Macrobiotus derkai, a new species of Tardigrada (Eutardigrada, Macrobiotidae, huzori group) from the Colombian Andes (South America). Zootaxa, 1731, 1 - 23.","Kaczmarek, L., Cytan, J., Zawierucha, K., Diduszko, D. & Michalczyk, L. (2014 b) Tardigrades from Peru (South America), with descriptions of three new species of Parachela. Zootaxa, 3790 (2), 357 - 379. http: // dx. doi. org / 10.11646 / zootaxa. 3790.2.5"]}

210. Paramacrobiotus richtersi (Murray, 1911) sensu lato [T] M. richtersii J. Murr. (Marcus 1939) Macrobiotus richtersii J. Murr., 1908 (de Barros 1942 b) M. richtersi J. Murray, 1911 (du Bois-Reymond Marcus 1944) Macrobiotus richtersii J. Murr. (Iharos 1963) Macrobiotus richtersi Murr. (Iharos 1969) Macrobiotus richtersi Murray, 1911 (Rossi & Claps 1980) M. richtersi Murray, 1911 (Claps & Rossi 1981, 1984, 1988) Macrobiotus richtersi J. Murray, 1911 (Iharos 1982) Macrobiotus richtersi Murray, 1911 (Maucci 1988, Rossi & Claps 1991, Jerez & Narv��ez 2001) M. richtersi J. Murr., 1911 (Rossi & Claps 1989) M. richtersi Murray (Claps & Rossi 1997) Macrobiotus harmsworthi / richtersi (Montoya et al. 2010) Terra typica: Ireland (Europe) Argentina: ��� 23 ��08���S, 64 �� 19 ���W; 350 m asl: Salta Province, Or��n [San Ram��n de la Nueva Or��n], mosses on rock. Claps & Rossi (1984) ��� 23 �� 36 ���S, 65 ��04���W; 2,000 m asl: Jujuy Province, Departamento Santa B��rbara, lichens on vina lito. Claps & Rossi (1984) ��� 24 ��07���S, 65 �� 24 ���W; 1,450 m asl: Jujuy Province, road to Yala, mosses from the edge of irrigation ditch and on tree (2 samples). Claps & Rossi (1984) ��� 24 �� 18 ���S, 64 �� 54 ���W; 700 m asl: Jujuy Province, road to Higueritas, mosses on soil. Claps & Rossi (1984) ��� 25 �� 43 ���S, 54 �� 25 ���W; 250 m asl: Misiones Province, Iguaz�� National Park, moss on rock and mosses on trees in the forest (3 samples). Claps & Rossi (1988) ��� 25 �� 58 ���S, 54 �� 34 ���W; 200 m asl: Misiones Province, Wanda, mosses on rocks. Claps & Rossi (1988) ��� 26 ��08���S, 54 �� 35 ���W; 150 m asl: Misiones Province, Arroyo Torocu��, 12 km S Puerto Esperanza, mosses on rocks (2 samples). Claps & Rossi (1988) ��� 26 �� 47 ���S, 65 �� 20 ���W; 750 m asl: Tucum��n Province, Horco Molle, mosses on soil and tree (2 samples). Claps & Rossi (1984) ��� 26 �� 51 ���S, 65 �� 26 ���W; 700 m asl: Tucum��n Province, road from San Miguel de Tucum��n to Tafi del Valle, mosses and lichens on rocks (2 samples). Claps & Rossi (1984) ��� 26 �� 56 ���S, 55 ��04���W; 200 m asl: Misiones Province, Capiov��, mosses on trees in the forest (2 samples). Claps & Rossi (1988) ��� 27 �� 27 ���S, 58 �� 39 ���W; 50 m asl: Corrientes Province, old road to Santa Ana, Selaginella sp. in lagoon and moss on tree (2 samples). Claps & Rossi (1988) ��� 35 �� 27 ���S, 58 �� 48 ���W; 0 m asl: Buenos Aires Province, Laguna San Miguel del Monte, on plant (Ceratophyllum sp.) Rossi & Claps (1991). ��� 36 �� 22 ���S, 56 �� 43 ���W; 0 m asl: Buenos Aires Province, camping San Clemente del Tuy, lichens on tree (Eucalyptus) in shade. Rossi & Claps (1980) ��� 36 �� 24 ���S, 56 �� 57 ���W; 0 m asl: Buenos Aires Province, Port General Lavalle, lichens on tree (Tamarisk) in full sun. Rossi & Claps (1980) ��� 40 �� 50 ���S, 71 �� 38 ���W; 900 m asl: Neuqu��n Province, Peninsula de Quetrihue, semi-wet, mosses on rocks on the sunny slope. Claps & Rossi (1981) ��� 41 ��00���S, 71 �� 30 ���W; 800 m asl: Neuqu��n Province, Isla Victoria, mosses on tree. Rossi & Claps (1989) ��� 41 �� 13 ���S, 71 �� 30 ���W; 2,250 m asl: Rio Negro Province, Nahuel Huapi National Park, Cerro Catedral, semi-wet, lichens and mosses on trees and roots (Nothofagus sp.) in the forest, very wet, mosses on soil and dead wood and mosses on leaf litter in the forest (6 samples). Claps & Rossi (1981) ��� 41 �� 58 ���S, 71 �� 29 ���W; 1,200 m asl: Rio Negro Province, El Bols��n, Piltriquitron Mt., mosses on dry soil under Mulinum spinosum and between Berberis shrubs. Iharos (1963) ��� 41 �� 58 ���S, 71 �� 28 ���W; 1,460 m asl: Rio Negro Province, El Bols��n, Piltriquitron Mt., mosses on rocks and barks of trees near edge within Nothofagus pumilio forest. Iharos (1963) ��� 41 �� 59 ���S, 71 �� 34 ���W; 300 m asl: Rio Negro Province, El Bols��n, Valley of Rio Azul, mosses on tree (Myrceugenia exupca) in the Myrceugenia exupca - Nothofagus dombeyi marsh forest. Iharos (1963) ��� 41 �� 59 ���S, 71 �� 31 ���W; 360 m asl: Rio Negro Province, El Bols��n, foot of Piltriquitron Mt., mosses on rocks in shaded Libocedrus chilensis -Lomatia obliqua forest. Iharos (1963) ��� 41 �� 59 ���S, 71 �� 31 ���W; 370 m asl: Rio Negro Province, El Bols��n, foot of Piltriquitron Mt., slope above Pampa Azcona, mosses on soil of shrubby area. Iharos (1963) ��� 42 �� 13 ���S, 71 �� 42 ���W; 220 m asl: Chubut Province, El Turbio, Puelo Lake, mosses on wet soil in primeval Nothofagus dombeyi - Libocedrus chilensis forest near the lake. Iharos (1963) ��� 54 �� 39 ���S, 68 �� 30 ���W; 800 m asl: Tierra del Fuego Province, National Park Ushuaia [Tierra del Fuego National Park], soil and litter (2 samples). Iharos (1982) Bolivia: ��� 10 �� 49 ���S, 65 �� 22 ���W; 150 m asl: Beni Department, Guayaramer��n, Estancia Esperanza, gallerian forest along Mamore river, leaf litter, leaf litter near water, decayed leaves on banana plantation, dry leaf litter, leaf litter from cacao plantation and soil from Leguminosae plantation (7 samples). Iharos (1969) ��� 10 �� 56 ���S, 65 �� 28 ���W; 150 m asl: Beni Department, Guayaramer��n, 15 km on W on the city, leaf litter in evergreen forest, near the road to Riberalta, leaf litter (2 samples). Iharos (1969) ��� 15 �� 31 ���S, 67 �� 33 ���W; 800 m asl: La Paz Department, between Alcoche and Puerto Linares, 20 km on Alcoche, mosses in shaded place. Iharos (1969) ��� 16 ��08���S, 67 �� 43 ���W; 1,600 m asl: La Paz Department, Coroico, 5 km on N of the city, wet leaf litter from citrus and coffee plantation. Iharos (1969) ��� 16 �� 11 ���S, 69 ��05���W; 4,100 m asl: La Paz Department, Copacabana [on the coast of Titicaca Lake], leaf litter from Stipa vegetation. Iharos (1969) ��� 16 �� 16 ���S, 67 �� 53 ���W; 3,100 m asl: La Paz Department, 5 km on N of Unduavi, leaf litter and soil from Vaccinum plantation. Iharos (1969) ��� 16 �� 19 ���S, 67 �� 54 ���W; 4,000 m asl: La Paz Department, near Unduavi, near the river, very wet leaf litter under shrubs. Iharos (1969) Brazil: ��� 03�� 17 ���S, 59 �� 53 ���W; 0 m asl: Amazonas State, Manaus, 20 km on S of the city, leaf litter from evergreen forest. Iharos (1969) ��� 21 �� 49 ���S, 49 ��05���W: Undefined localities in S��o Paulo State, few localities. Marcus (1939), de Barros (1942 b) ��� 22 �� 44 ���S, 45 �� 35 ���W; 1,650 m asl: S��o Paulo State, Campos do Jord��o, mosses or aquatic plants. du Bois-Reymond Marcus (1944) ��� 22 �� 45 ���S, 47 �� 24 ���W; 550 m asl: S��o Paulo State, Munic��pio de Sta. B��rbara [Munic��pio Santa B��rbara d ���Oeste], mosses or aquatic plants. du Bois-Reymond Marcus (1944) ��� 22 �� 54 ���S, 47 ��04���W; 700 m asl: S��o Paulo State, Munic��pio Campinas, in bromeliads. du Bois-Reymond Marcus (1944) ��� 25 �� 26 ���S, 49 �� 16 ���W; 900 m asl: Paran�� State, Curitiba, mosses or aquatic plants. du Bois-Reymond Marcus (1944) Chile: ��� 18 �� 11 ���S, 69 �� 13 ���W; 4,800 m asl [4,550 m asl]: Region XV Arica y Parinacota, Laguna La Cotacotani, leaf litter. Iharos (1969) ��� 18 �� 34 ���S, 70 ��05���W; 500 m asl: Region XV Arica y Parinacota, Azapa Chico, 25 km. on East of Arica, near the river Rio Lauca, wet soil on bamboo plantation. Iharos (1969) ��� 20 �� 12 ���S, 69 �� 17 ���W: Undefined locality in Region I Tarapac, Bofedal de Bajuco, soil under red cactus. Iharos (1969) ��� 30 �� 17 ���S, 71 �� 16 ���W; 400 m asl: Region IV Coquimbo, Las Cardas, leaf litter under shrubs. Iharos (1969) ��� 30 �� 40 ���S, 71 �� 41 ���W; 900 m asl [650 m asl]: Region IV Coquimbo, Fray Jorge [Bosque de Fray Jorge National Park], foggy forest, leaf litter. Iharos (1969) ��� 31 �� 55 ���S, 71 �� 31 ���W; 0 m asl: Region IV Coquimbo, Los Vilos, jungle soil and leaf litter. Iharos (1969) ��� 32 �� 58 ���S, 71 ��01���W; 1,900 m asl: Region V Valpara��so (Regi��n de Valpara��so), Cordiliera de la Costa, Cerro El Roble, 84 km NW of Santiago de Chile, leaf litter under snow, under the bushes. Iharos (1969) ��� 32 �� 58 ���S, 71 ��01���W; 1,700 m asl: Region V Valpara��so (Regi��n de Valpara��so), Cordiliera de la Costa, Cerro El Roble, 84 km NW of Santiago de Chile, leaf litter between dry bushes. Iharos (1969) ��� 33 ��04���S, 71 ��00���W; 1,000 m asl: Region RM Metropolitana (Regi��n Metropolitana de Santiago), Tiltil, Cuesta La Dormida, jungle leaf litter. Iharos (1969) ��� 33 �� 21 ���S, 70 �� 19 ���W; 2,300 m asl: Region RM Metropolitana (Regi��n Metropolitana de Santiago), Farellones, 30 km on East of Santiago de Chile, detritus and soil under the bushes, wet mosses from rocky cave and leaf litter (3 samples). Iharos (1969) ��� 33 �� 26 ���S, 70 �� 38 ���W; 800 m asl: Region RM Metropolitana (Regi��n Metropolitana de Santiago), Santiago de Chile, San Crist��bal Mt., mosses on rocks. Iharos (1969) ��� 33 �� 30 ���S, 70 �� 55 ���W; 850 m asl: Region RM Metropolitana (Regi��n Metropolitana de Santiago), Quebrada de La Plata, shrubs upland, moss and lichen on rocks (2 samples). Iharos (1969) ��� 37 �� 13 ���S, 72 �� 23 ���W; 100 m asl: Region VIII Biob��o, Salto del Laja, near waterfalls, mosses on rocky wall. Iharos (1969) ��� 50 �� 59 ���S, 73 ��00���W; 1,200 m asl: Region XII Magallanes (Regi��n de Magallanes y de la Ant��rtica Chilena), Torres del Paine National Park, mosses on trees and rocks, nearly all in half sunlight. Maucci (1988) ��� 51 �� 34 ���S, 72 �� 36 ���W; 200 m asl: Region XII Magallanes (Regi��n de Magallanes y de la Ant��rtica Chilena), Ultima Esperanza, near Cueva del Milod��n, mosses on rocks, in sunlight. Maucci (1988) Colombia: ��� 07��07���N, 73 ��02���W; 1,795���1,970 m asl: Santander Department, El Diviso Natural Reserve, Pine forest (Pinus patula), mosses (Hypopterygium tamariscinum, Fabronia ciliaris and Sematophyllum insularum). Jerez & Narv��ez (2001) Uruguay: ��� 30 �� 15 ���S, 57 �� 36 ���W; 50 m asl: Artigas Department, Bella Uni��n, mosses or lichens on rocks, trees or posts or aquatic macrophytes. Claps & Rossi (1997) ��� 31 ��03���S, 57 �� 38 ���W; 50 m asl: Salto Department, Arroyo Palomas, mosses or lichens on rocks, trees or posts or aquatic macrophytes. Claps & Rossi (1997) ��� 31 �� 20 ���S, 57 �� 51 ���W; 50 m asl: Salto Department, Arroyo San Ant��nio de Chico, mosses or lichens on rocks, trees or posts or aquatic macrophytes. Claps & Rossi (1997) Venezuela: ��� 08�� 46 ���N, 70 �� 49 ���W; 4,300 m asl: Region Los Andes, M��rida, Mucu��uque Mt., moss. Montoya et al. (2010) ��� 08�� 45 ���N, 70 �� 49 ���W; 4,450 m asl: Region Los Andes, M��rida, Mucu��uque Mt., surface samples of peat bog. Montoya et al. (2010) Record numbers: Argentina: 24, Bolivia: 7, Brazil: 6, Chile: 15, Colombia: 1, Uruguay: 3, Venezuela: 2; total: 58. Remarks: Paramacrobiotus richtersi was, until recently, believed to be cosmopolitan (McInnes 1994 a). New taxonomic data has shown that this is the nominal species for a species complex ��� the ��� richtersi group��� ��� requiring careful examination of both adults and eggs. Several newly described species of this group have been added to the faunas of South and Central America (e.g. Michalczyk & Kaczmarek 2006 b, Michalczyk et al. 2006, Pilato et al. 2006). In the light of these reports, and given P. richtersi was originally described from Europe, we suggest all species records from South America should be treated with caution until the original slides are re-examined or the species is otherwise verified from the region., Published as part of Kaczmarek, ��ukasz, Michalczyk, ��ukasz & Mcinnes, Sandra J., 2015, Annotated zoogeography of non-marine Tardigrada. Part II: South America, pp. 1-107 in Zootaxa 3923 (1) on pages 85-87, DOI: 10.11646/zootaxa.3923.1.1, http://zenodo.org/record/241936, {"references":["Murray, J. (1911) Arctiscoida. Proceedings of the Royal Irish Academy, 31, 1 - 16.","Marcus, E. (1939) Tardigrada. The Percy Sladen Trust Exped. to Lake Titicaca in 1937. Transactions of the Linnean Society of London, Series 3, 1, 45 - 59.","De Barros, R. (1942 b) Tardigrados de Estado de Sao Paulo, Brasil. II. Genero Macrobiotus. Revista Brasileira de Biologia, 2, 373 - 386.","Bois-Reymond Marcus, E. du. (1944) Sobre tardigrados brasileiros. Comunicaciones Zoologicas del Museo de Historia Natural de Montevideo, 1 (13), 1 - 19.","Iharos, G. (1963) The zoological results of Gy. Topal's collections in South Argentina, 3. Tardigrada. Annales Historico Naturales Musei Nationalis Hungarici, 55, 293 - 299.","Iharos, G. (1969) The scientific results of the Hungarian soil zoological expeditions to South America: 15. Tardigraden aus den Sammlungen der ersten und zweiten Expedition. Opuscula Zoologica, Budapest, 9, 279 - 289.","Rossi, G. C. and Claps, M. C. (1980) Contribucion al conocimiento de los tardigrados de Argentina. I. Revista de la Sociedad Entomologica Argentina, 39, 243 - 250","Claps, M. C. & Rossi, G. C. (1981) Contribucion al conocimiento de los tardigrados de Argentina. II. Revista de la Sociedad Entomologica Argentina, 40 (1 - 4), 107 - 114.","Iharos, G. (1982) Tardigradologische Notizen, I. Miscellanea Zoologica Hungarica, 1, 85 - 90.","Maucci, W. (1988) Tardigrada from Patagonia (Southern South America) with description of three new species. Revista Chilena de Entomologia, 16, 5 - 13.","Rossi, G. C. & Claps, M. C. (1991) Tardigrados dulceacuicolas de la Argentina. Fauna de aqua dulce de la Republica Argentina, 19, 1 - 70.","Jerez Jaimes, J. H. & Narvaez Parra, E. X. (2001) Tardigrados (Animalia, tardigrada) de la Reserva El Diviso - Santander, Colombia. Biota Colombiana, 2, 145 - 151.","Rossi, G. C. & Claps, M. C. (1989) Contribucion al conocimiento de los Tardigrados de la Argentina V. Revista de la Sociedad Entomologica Argentina, 47, 133 - 142.","Claps, M. C. & Rossi, G. C. (1997) Tardigrados de Uruguay, com descripcion de dos nuevas especies (Echiniscidae, Macrobiotidae). Iheringia, Serie Zoologia, 83, 17 - 22.","Montoya, E., Rull, V. & van Geel, B. (2010) Non-pollen palynomorphs from surface sediments along an altitudinal transect of the Venezuelan Andes. Palaeogeography, Palaeoclimatology, Palaeoecology, 297, 169 - 183. http: // dx. doi. org / 10.1016 / j. palaeo. 2010.07.026","Claps, M. C. & Rossi, G. C. (1984) Contribucion al conocimiento de los tardigrados de Argentina. IV. Acta Zoologica Lilloana, 38, 45 - 50.","Claps, M. C. & Rossi, G. C. (1988) Contribucion al conocimiento de los tardigrados de Argentina. VI. Iheringia, 67, 3 - 11.","McInnes, S. J. (1994 a) Zoogeographic distribution of terrestrial / freshwater tardigrades from current literature. Journal of Natural History, 28, 257 - 352. http: // dx. doi. org / 10.1080 / 00222939400770131"]}

206. Paramacrobiotus centesimus (Pilato, 2000) [T] Macrobiotus centesimus sp. n. (Pilato 2000) Terra typica: Brazil and Ecuador (South America) Brazil: ��� 25 �� 41 ���S, 54 �� 26 ���W; 200 m asl: Type Locality: Paran�� State, Igua��u Falls, moss. Pilato (2000) Ecuador: ��� 0��09���S, 78 �� 32 ���W [00��09���S, 78 �� 32 ���W]; 3,800 m asl: Pichincha Province, Pichincha Volcano (eastern slope), soil sample. Pilato (2000) Record numbers: Brazil: 1, Ecuador: 1; total: 2. Remarks: Currently endemic to South America, this species belongs to the Paramacrobiotus richtersi group (see: P. areolatus above and P. richtersi below)., Published as part of Kaczmarek, ��ukasz, Michalczyk, ��ukasz & Mcinnes, Sandra J., 2015, Annotated zoogeography of non-marine Tardigrada. Part II: South America, pp. 1-107 in Zootaxa 3923 (1) on page 85, DOI: 10.11646/zootaxa.3923.1.1, http://zenodo.org/record/241936

206. Paramacrobiotus centesimus (Pilato, 2000) [T] Macrobiotus centesimus sp. n. (Pilato 2000) Terra typica: Brazil and Ecuador (South America) Brazil: • 25 ° 41 ′S, 54 ° 26 ′W; 200 m asl: Type Locality: Paraná State, Iguaçu Falls, moss. Pilato (2000) Ecuador: • 0°09′S, 78 ° 32 ′W [00°09′S, 78 ° 32 ′W]; 3,800 m asl: Pichincha Province, Pichincha Volcano (eastern slope), soil sample. Pilato (2000) Record numbers: Brazil: 1, Ecuador: 1; total: 2. Remarks: Currently endemic to South America, this species belongs to the Paramacrobiotus richtersi group (see: P. areolatus above and P. richtersi below).

208. Paramacrobiotus derkai (Degma, Michalczyk & Kaczmarek, 2008) [T] Macrobiotus derkai sp. nov. (Degma et al. 2008) Terra typica: Colombia (South America) Colombia: ��� 06�� 26 ���N, 72 �� 15 ���W; 4,300 m asl: Arauca Department, Cordillera Oriental, Sierra Nevada del Cocuy National Park, near Laguna de la Plaza (mountain lake), moss on rock. Degma et al. (2008) Record numbers: Peru: 1; total: 1. Remarks: Currently endemic to Colombia, this species belongs to the Paramacrobiotus richtersi group (see: P. areolatus above and P. richtersi below)., Published as part of Kaczmarek, ��ukasz, Michalczyk, ��ukasz & Mcinnes, Sandra J., 2015, Annotated zoogeography of non-marine Tardigrada. Part II: South America, pp. 1-107 in Zootaxa 3923 (1) on page 85, DOI: 10.11646/zootaxa.3923.1.1, http://zenodo.org/record/241936, {"references":["Degma, P., Michalczyk, L. & Kaczmarek, L. (2008) Macrobiotus derkai, a new species of Tardigrada (Eutardigrada, Macrobiotidae, huzori group) from the Colombian Andes (South America). Zootaxa, 1731, 1 - 23."]}

208. Paramacrobiotus derkai (Degma, Michalczyk & Kaczmarek, 2008) [T] Macrobiotus derkai sp. nov. (Degma et al. 2008) Terra typica: Colombia (South America) Colombia: • 06° 26 ′N, 72 ° 15 ′W; 4,300 m asl: Arauca Department, Cordillera Oriental, Sierra Nevada del Cocuy National Park, near Laguna de la Plaza (mountain lake), moss on rock. Degma et al. (2008) Record numbers: Peru: 1; total: 1. Remarks: Currently endemic to Colombia, this species belongs to the Paramacrobiotus richtersi group (see: P. areolatus above and P. richtersi below).

209. Paramacrobiotus intii Kaczmarek, Cytan, Zawierucha, Diduszko & Michalczyk 2014 [T] Paramacrobiotus intii sp. nov. (Kaczmarek et al. 2014 b) Terra typica: Peru (South America) Peru: • 13 ° 25 ′S, 71 ° 51 ′W; 3,000 m asl: Type Locality: Cusco Region, Pisac near Cusco, mixed mosses and lichens on rock. Kaczmarek et al. (2014 b) Record numbers: Peru: 1; total: 1. Remarks: Currently endemic to Peru, this species belongs to the Paramacrobiotus areolatus group (see: P. areolatus above and P. richtersi below).

By examining material collected in the Sierra Nevada de Santa Marta (Department of Magdalena, Colombia), the authors report a new record for the entire South America, Doryphoribius gibber Beasley & Pilato, 1987, and two species new to science, Paramacrobiotus sagani sp. nov., and Doryphoribius rosane sp. nov., are described. Paramacrobiotus sagani sp. nov. belongs to the richtersi group, vanescens subgroup (by having sculptured egg areolae) and is characterized by a peculiar cuticular granulation, trunco-conical egg processes with cylindrical indented apices, and other more detailed cha-racters both qualitative and metric; by the unique combination of characters, it differs from all the other known species of the genus. Doryphoribius rosanae sp. nov. is characterized by reticulated dorsal cuticle with gibbosities (formula IX:4-6-2-6-2-6-4-2-2), two macroplacoids in the pharynx without microplacoid or septulum, and large, stout claws without “free” accessory points but with lunules. It differs from all the other Doryphoribius species with gibbosities by having a unique formula, as well as other more detailed characters. Thanks to this contribution, the number of tardigrade species known for Colombia increases from 52 to 55.

Two new eutardigrade species are described: Eremobiotus ginevrae sp. nov. and Paramacrobiotus pius sp. nov. The first is similar to Eremobiotus alicatai (Binda, 1969) but differs in the claw shape and dimensions. It has been found in Sicily, Israel and Russia. The second species, belonging to the richtersi group, is currently found exclusively in Sicily. It has a smooth cuticle, three macroplacoids and a microplacoid, eggs with reticulated trunco-conical processes with small terminal thorns; the egg-shell is areolated and the areolae are clearly sculptured.

7. Paramacrobiotus csotiensis (Iharos, 1966) Margoitsy (2 moss samples: 7 specimens + 6 eggs). The species, already recorded from Belarus, was recorded from some European localities. 8. Paramacrobiotus klymenki sp. nov. Margoitsy (1 moss sample: 71 specimens + 45 eggs). Referred to as Macrobiotus sp. 4. gr. richtersi by Kiosya (2010)., Published as part of Pilato, Giovanni, Kiosya, Yevgen, Lisi, Oscar & Sabella, Giorgio, 2012, New records of Eutardigrada from Belarus with the description of three new species, pp. 39-60 in Zootaxa 3179 on page 42, DOI: 10.5281/zenodo.279945, {"references":["Iharos, G. (1966) Neue Tardigraden-Arten aus Ungarn. (Neuere Beitrage zur Kenntnis der Tardigraden-Fauna Ungarns. VI). Acta Zoologica Academiae Scientiarum Hungaricae, XII (1 - 2), 111 - 122.","Kiosya, Y. O. (2010) New data on Tardigrada of Belarus. Proceedings of the National Academy of Sciences of Belarus, Supplement \" Young scientists 2009 \", series Biology (4), 135 - 138. [In Russian]."]}

Paramacrobiotus richtersi (Murray, 1911) Material examined in this study: 8 specimens and 6 eggs in sample B. Remarks: Species known from many localities throughout the world (including Turkey), but as this name incorporates a species complex, the majority of records need to be verified (McInnes 1994, Guidetti et al. 2009). ......continued on the next page......continued on the next page, Published as part of Kaczmarek, ��ukasz, Jakubowska, Natalia & Michalczyk, ��ukasz, 2012, Current knowledge on Turkish tardigrades with a description of Milnesium beasleyi sp. nov. (Eutardigrada: Apochela: Milnesiidae, the granulatum group), pp. 49-64 in Zootaxa 3589 on pages 58-59, DOI: 10.5281/zenodo.213616, {"references":["Murray, J. (1911) Arctiscoida. Proceedings of the Royal Irish Academy, 31, 1 - 16.","McInnes, S. J. (1994) Zoogeographic distribution of terrestrial / freshwater tardigrades from current literature. Journal of Natural History, 28, 257 - 352.","Guidetti, R., Schill, R. O., Bertolani, R., Dandekar, T. & Wolf, M. (2009) New molecular data for tardigrade phylogeny, with the erection of Paramacrobiotus gen. nov. Journal of Zoological Systematics and Evolutionary Research, 47, 315 - 321."]}

Paramacrobiotus klymenki sp. nov. (Fig. 6) Type material. Vitebsk Oblast, Sennensky district, spruce forest, vicinity of Margoitsy village (54 ° 40 ' 19 "N, 29 ° 37 ' 27 "E); moss sample collected (June 2002) by G.O. Mazepa: holotype, 71 paratypes and 45 eggs. Type repository. Holotype (slide No. 5425), 23 paratypes and 14 eggs (slide No. 5425) are deposited in the collection of Binda & Pilato (Museum of the Department of Animal Biology “Marcello La Greca”, University of Catania, Italy). Other specimens and eggs are deposited in the collection of Kiosya (Kharkiv National University, Ukraine). Specific diagnosis. Colourless, cuticle smooth without pores; small dots present on all legs. Eye spots absent. Pharyngeal bulb with three macroplacoids; microplacoid absent; buccal tube not very wide (pt 14.7–16.6); stylet supports inserted on the buccal tube at 79–80 %. Claws, of hufelandi - type, slender, with accessory points on the main branches; lunules present. The eggs, freely laid, were spherical, with 10–11 conical processes on the circumference. The processes’ surface had a well visible reticular design consisting of elongate mesh with rectilinear margins. Egg shell areolated; the central area of each areola slightly thickened and not subdivided. Description of the holotype: Body length 340 µm; colourless, cuticle smooth without pores; small dots present on the legs. Eye spots absent. Bucco-pharyngeal apparatus of Macrobiotus - type (Fig. 6 A); mouth terminal with ten peribuccal lamellae; buccal armature with an anterior ring of small teeth, a posterior ring of triangular teeth and a system of three dorsal and three ventral transverse ridges. Buccal tube 42.1 µm long and 6.6 µm wide externally (pt = 15.7). Stylet supports inserted on the buccal tube at 79.8 % of its length (pt = 79.8). Pharyngeal bulb with apophyses and three macroplacoids; microplacoid absent (Fig. 6 A). Length of first macroplacoid 7.4 µm (pt = 17.6), second 5.2 µm (pt = 12.4), third 7.3 µm (pt = 17.3); entire macroplacoid row 21.2 µm long (pt = 50.4) Claws, of hufelandi - type, slender, with accessory points on the main branches (Fig. 6 B–D). External and internal claws of the first pair of legs 10.4 µm (pt = 24.7) and 10.2 µm (pt = 24.2) long respectively; those of the second and third pair of legs 11.7 µm (pt = 27.8) and 10.9 µm (pt = 25.9) long respectively; posterior and anterior claws of the hind legs 13.0 µm (pt = 30.9) and 11.8 µm (pt = 28.0) long respectively. Lunules present, small and smooth on the first three pairs of legs, larger and with indented margin on the hind legs. Two cuticular bars present below the claws on the first three pairs of legs (Fig. 6 B, arrow). Description of the eggs. The eggs, freely laid, were spherical, with 10–11 conical processes on the circumference and 22–26 on the hemisphere. The diameter was 69–71 µm excluding the processes, 101–109 µm including. The processes were conical, elongate (sometimes with slightly forked apices), 14.5–18.5 high with basal diameter of 16.4–18.2 µm. The processes surface had a well visible reticular design consisting of elongate mesh with rectilinear margins (Fig. 6 E). A short apical portion of each process was smooth (Fig. 6 E, arrows). Egg shell areolated and the central area of each areola slightly thickened. Remarks. The paratypes were similar to the holotype in both qualitative and metric characters (we referred to the pt index values when the specimens had different body length). The measurements of some specimens, the smallest and the largest ones included, are presented in Table 3. Etymology. The specific name is in honour to Prof. Vyacheslav Klymenko (Kharkiv National University, Ukraine), the scientific adviser of Ye. Kiosya. Differential diagnosis. Prior to the new species described above, six species of the genus Paramacrobiotus were described with a smooth cuticle, three macroplacoids, lacking the microplacoid and producing eggs with areolated shell and conical processes: P. areolatus, P. crenatus, P. walteri, P. centesimus, P. huziori and P. de r k a i. Comparing adults of, where possible, comparable size the new species can be from the other species (Table 3) as follows: Paramacrobiotus klymenki sp. nov. was very similar to P. a re o l a t u s but differed in lacking eye spots; in having the buccal tube narrower and slightly longer with respect to the body length (Table 3; Figs. 6 A and 7 A); and with slightly shorter claws (Table 3) (however, the pt values may appear similar as the buccal tube of P. a re o l a t u s was slightly shorter with respect to the body length). The new species differed from P. crenatus in having a narrower buccal tube; lunules on the hind legs less developed; smaller eggs (diameter 69–71 µm excluding the processes in P. klymenki sp. nov., 92–128 µm in P. c re - natus); smaller egg processes (14.5–18.5 µm high in P. klymenki sp. nov., 40–44 µm in P. crenatus, with basal diameter 16.4–18.2 µm in P. klymenki sp. nov. and 24–28 µm in P. crenatus). Paramacrobiotus klymenki sp. nov. differed from P. w al te r i in lacking eye spots; in having the stylet supports inserted on the buccal tube in a more caudal position (Table 3), shorter claws (Table 3), also different in shape (in P. walteri the difference in length between main and secondary branch was markedly higher, as seen in Figs. 6 B–D and 7 D). Paramacrobiotus klymenki sp. nov. differed from P. centesimus in having a narrower buccal tube (Table 3; Figs. 6 A and 8 A), lunules on the hind legs indented; longer egg processes (14.5–18.5 µm in P. klymenki sp. nov., 7–11 µm in P. centesimus), slightly different reticular design of the processes (the design was less dense, and the mesh larger and more elongate in P. klymenki sp. nov., while the design was more dense and the mesh almost isodiametric in P. centesimus (Figs. 6 E and 8 B). Paramacrobiotus klymenki sp. nov. differed from P. h u z i o r i in lacking eye spots, in having a narrower buccal tube, lunules on the hind legs indented, egg processes with less dense reticular design and more clearly elongate mesh with more rectilinear margins (Figs. 6 E and 8 C, D, arrow a); and egg shell areolae not subdivided (Figs. 6 E and 8 C, D, arrow b, and 8 C,D, arrows b). Lack of eye spots, indented lunules on hind legs, claws with less developed accessory points, egg shell areolae not subdivided, egg processes with basal diameter longer than the process height and reticular design less dense, distinguishes Paramacrobiotus klymenki sp. nov. from P. derkai.

Paramacrobiotus klymenki sp. nov. (Fig. 6) Type material. Vitebsk Oblast, Sennensky district, spruce forest, vicinity of Margoitsy village (54 �� 40 ' 19 "N, 29 �� 37 ' 27 "E); moss sample collected (June 2002) by G.O. Mazepa: holotype, 71 paratypes and 45 eggs. Type repository. Holotype (slide No. 5425), 23 paratypes and 14 eggs (slide No. 5425) are deposited in the collection of Binda & Pilato (Museum of the Department of Animal Biology ���Marcello La Greca���, University of Catania, Italy). Other specimens and eggs are deposited in the collection of Kiosya (Kharkiv National University, Ukraine). Specific diagnosis. Colourless, cuticle smooth without pores; small dots present on all legs. Eye spots absent. Pharyngeal bulb with three macroplacoids; microplacoid absent; buccal tube not very wide (pt 14.7���16.6); stylet supports inserted on the buccal tube at 79���80 %. Claws, of hufelandi - type, slender, with accessory points on the main branches; lunules present. The eggs, freely laid, were spherical, with 10���11 conical processes on the circumference. The processes��� surface had a well visible reticular design consisting of elongate mesh with rectilinear margins. Egg shell areolated; the central area of each areola slightly thickened and not subdivided. Description of the holotype: Body length 340 ��m; colourless, cuticle smooth without pores; small dots present on the legs. Eye spots absent. Bucco-pharyngeal apparatus of Macrobiotus - type (Fig. 6 A); mouth terminal with ten peribuccal lamellae; buccal armature with an anterior ring of small teeth, a posterior ring of triangular teeth and a system of three dorsal and three ventral transverse ridges. Buccal tube 42.1 ��m long and 6.6 ��m wide externally (pt = 15.7). Stylet supports inserted on the buccal tube at 79.8 % of its length (pt = 79.8). Pharyngeal bulb with apophyses and three macroplacoids; microplacoid absent (Fig. 6 A). Length of first macroplacoid 7.4 ��m (pt = 17.6), second 5.2 ��m (pt = 12.4), third 7.3 ��m (pt = 17.3); entire macroplacoid row 21.2 ��m long (pt = 50.4) Claws, of hufelandi - type, slender, with accessory points on the main branches (Fig. 6 B���D). External and internal claws of the first pair of legs 10.4 ��m (pt = 24.7) and 10.2 ��m (pt = 24.2) long respectively; those of the second and third pair of legs 11.7 ��m (pt = 27.8) and 10.9 ��m (pt = 25.9) long respectively; posterior and anterior claws of the hind legs 13.0 ��m (pt = 30.9) and 11.8 ��m (pt = 28.0) long respectively. Lunules present, small and smooth on the first three pairs of legs, larger and with indented margin on the hind legs. Two cuticular bars present below the claws on the first three pairs of legs (Fig. 6 B, arrow). Description of the eggs. The eggs, freely laid, were spherical, with 10���11 conical processes on the circumference and 22���26 on the hemisphere. The diameter was 69���71 ��m excluding the processes, 101���109 ��m including. The processes were conical, elongate (sometimes with slightly forked apices), 14.5���18.5 high with basal diameter of 16.4���18.2 ��m. The processes surface had a well visible reticular design consisting of elongate mesh with rectilinear margins (Fig. 6 E). A short apical portion of each process was smooth (Fig. 6 E, arrows). Egg shell areolated and the central area of each areola slightly thickened. Remarks. The paratypes were similar to the holotype in both qualitative and metric characters (we referred to the pt index values when the specimens had different body length). The measurements of some specimens, the smallest and the largest ones included, are presented in Table 3. Etymology. The specific name is in honour to Prof. Vyacheslav Klymenko (Kharkiv National University, Ukraine), the scientific adviser of Ye. Kiosya. Differential diagnosis. Prior to the new species described above, six species of the genus Paramacrobiotus were described with a smooth cuticle, three macroplacoids, lacking the microplacoid and producing eggs with areolated shell and conical processes: P. areolatus, P. crenatus, P. walteri, P. centesimus, P. huziori and P. de r k a i. Comparing adults of, where possible, comparable size the new species can be from the other species (Table 3) as follows: Paramacrobiotus klymenki sp. nov. was very similar to P. a re o l a t u s but differed in lacking eye spots; in having the buccal tube narrower and slightly longer with respect to the body length (Table 3; Figs. 6 A and 7 A); and with slightly shorter claws (Table 3) (however, the pt values may appear similar as the buccal tube of P. a re o l a t u s was slightly shorter with respect to the body length). The new species differed from P. crenatus in having a narrower buccal tube; lunules on the hind legs less developed; smaller eggs (diameter 69���71 ��m excluding the processes in P. klymenki sp. nov., 92���128 ��m in P. c re - natus); smaller egg processes (14.5���18.5 ��m high in P. klymenki sp. nov., 40���44 ��m in P. crenatus, with basal diameter 16.4���18.2 ��m in P. klymenki sp. nov. and 24���28 ��m in P. crenatus). Paramacrobiotus klymenki sp. nov. differed from P. w al te r i in lacking eye spots; in having the stylet supports inserted on the buccal tube in a more caudal position (Table 3), shorter claws (Table 3), also different in shape (in P. walteri the difference in length between main and secondary branch was markedly higher, as seen in Figs. 6 B���D and 7 D). Paramacrobiotus klymenki sp. nov. differed from P. centesimus in having a narrower buccal tube (Table 3; Figs. 6 A and 8 A), lunules on the hind legs indented; longer egg processes (14.5���18.5 ��m in P. klymenki sp. nov., 7���11 ��m in P. centesimus), slightly different reticular design of the processes (the design was less dense, and the mesh larger and more elongate in P. klymenki sp. nov., while the design was more dense and the mesh almost isodiametric in P. centesimus (Figs. 6 E and 8 B). Paramacrobiotus klymenki sp. nov. differed from P. h u z i o r i in lacking eye spots, in having a narrower buccal tube, lunules on the hind legs indented, egg processes with less dense reticular design and more clearly elongate mesh with more rectilinear margins (Figs. 6 E and 8 C, D, arrow a); and egg shell areolae not subdivided (Figs. 6 E and 8 C, D, arrow b, and 8 C,D, arrows b). Lack of eye spots, indented lunules on hind legs, claws with less developed accessory points, egg shell areolae not subdivided, egg processes with basal diameter longer than the process height and reticular design less dense, distinguishes Paramacrobiotus klymenki sp. nov. from P. derkai., Published as part of Pilato, Giovanni, Kiosya, Yevgen, Lisi, Oscar & Sabella, Giorgio, 2012, New records of Eutardigrada from Belarus with the description of three new species, pp. 39-60 in Zootaxa 3179 on pages 49-54, DOI: 10.5281/zenodo.279945

6. Paramacrobiotus richtersi (Murray, 1911) Margoitsy (1 moss sample: 9 specimens + 5 eggs); Minsk (1 moss sample: 5 specimens + 2 eggs); Zhodino (2 moss samples: 3 eggs). The species, already reported from Belarus, according to the current literature is considered cosmopolitan but many diagnoses should be confirmed., Published as part of Pilato, Giovanni, Kiosya, Yevgen, Lisi, Oscar & Sabella, Giorgio, 2012, New records of Eutardigrada from Belarus with the description of three new species, pp. 39-60 in Zootaxa 3179 on page 42, DOI: 10.5281/zenodo.279945, {"references":["Murray, J. (1911) Artiscoidea. Proceedings of the Royal Irish Academy, 31 Clare Island Survey, 37, 1 - 16."]}

Paramacrobiotus richtersi (Murray, 1911) Material examined in this study: 8 specimens and 6 eggs in sample B. Remarks: Species known from many localities throughout the world (including Turkey), but as this name incorporates a species complex, the majority of records need to be verified (McInnes 1994, Guidetti et al. 2009). ......continued on the next page......continued on the next page

Paramacrobiotus cf. lorenae (Biserov, 1996) Material examined. Two moss samples from Site 5 contained a total of 37 adults (three in the simplex stage) and 26 eggs. Remarks. All morphometric data of these specimens were consistent with Paramacrobiotus lorenae except the annulation of the egg processes was not seen. However, these specimens were different from other species in the P. richtersi group by one or more characteristics. P. lorenae was described from Popov Island, part of eastern Russia, and has so far not been reported from any other location. This is a new record for China., Published as part of Beasley, Clark W. & Miller, William R., 2012, Additional Tardigrada from Hubei Province, China, with the description of Doryphoribius barbarae sp. nov. (Eutardigrada: Parachela: Hypsibiidae), pp. 55-63 in Zootaxa 3170 on page 61, DOI: 10.5281/zenodo.212706, {"references":["Biserov, V. (1996) Macrobiotus lorenae sp. n., a new species of Tardigrada (Eutardigrada Macrobiotidae) from the Russian Far East. Arthropoda Selecta, 5, 1 - 2, 145 - 149."]}

In three mixed samples (mosses and leaf litter) collected in Zambia (southern Africa), forty-three specimens and six eggs of eutardigrades were found. Among them, 29 specimens belonged to a new species of the family Isohypsibiidae, Doryphoribius niedbalai n.sp. The new species belongs to the Doryphoribius evelinae group and it differs from other members of the group mainly by a different number and configuration of dorsal gibbosities, as well as by some morphometric characters. Apart from the new species, the examined material contained also two rare African eutardigrades: Milnesium tetralamellatum and Calcarobiotus (Calcarobiotus) parvicalcar. These species are recorded for the first time outside their type localities. Additionally, Paramacrobiotus vanescens is recorded for the fourth time from Africa. In this paper, together with the description of the new species, first ever photomicrographs of Milnesium tetralamellatum and Paramacrobiotus vanescens are also provided. This is the first account of the p...

Paramacrobiotus richtersi (Murray, 1911) Four eggs, samples 2 d (Table 1), leaf litter sample. Cosmopolitan species, also reported from the Dominican Republic, Puerto Rico and Saint Lucia., Published as part of Meyer, Harry A., 2011, Tardigrada of Grand Cayman, West Indies, with descriptions of two new species of eutardigrade, Doryphoribius tessellatus (Hypsibiidae) and Macrobiotus caymanensis (Macrobiotidae), pp. 28-40 in Zootaxa 2812 on page 38, DOI: 10.5281/zenodo.205995, {"references":["Murray, J. (1911) Arctiscoida. Proceedings of the Royal Irish Academy, 31, 1 - 16."]}