taxonID	type	description	language	source
7D3DFF4AFFE1FFA1582FFC7CCB0A2368.taxon	description	urn: lsid: zoobank. org: act: 0 F 5 EEA 46 - 2 A 3 D- 4 D 72 - B 776 - CB 812 C 013993 Figs 4 – 8	en	Hulterström, Jens, Guidetti, Roberto, Jönsson, K. Ingemar, Atherton, Sarah (2025): Integrative taxonomy supports the description of two new species of Macrobiotidae (Tardigrada, Eutardigrada) from Kristianstads Vattenrike Biosphere Reserve in Sweden. European Journal of Taxonomy 1030: 1-52, DOI: 10.5852/ejt.2025.1030.3135, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3135/13991
7D3DFF4AFFE1FFA1582FFC7CCB0A2368.taxon	diagnosis	Diagnosis Macrobiotus with three bands of teeth in oral cavity armature (OCA): first and second of granules, and third of a transverse crest non-uniform in shape on the dorsal and ventral side, with a dorsal side with one elongated tooth and ventral side with two lateral and two medial teeth. Two macroplacoids, the first with a slight central constriction, the second with a pre-terminal constriction, and a comma-shaped microplacoid. Claws of similar size on all legs, with large, smooth lunulae on legs I – III and serrated on legs IV. Eggs of hufelandi - type with inverse goblet-shaped processes with indented process disks without granulation. Egg chorion with a wrinkled or granulated surface, appears solid in LM but with a ring of small pores around each process visible in SEM.	en	Hulterström, Jens, Guidetti, Roberto, Jönsson, K. Ingemar, Atherton, Sarah (2025): Integrative taxonomy supports the description of two new species of Macrobiotidae (Tardigrada, Eutardigrada) from Kristianstads Vattenrike Biosphere Reserve in Sweden. European Journal of Taxonomy 1030: 1-52, DOI: 10.5852/ejt.2025.1030.3135, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3135/13991
7D3DFF4AFFE1FFA1582FFC7CCB0A2368.taxon	etymology	Etymology The new species is named after the area of the type locality within Kristianstads Vattenrike Biosphere Reserve, in dedication to its high tardigrade biodiversity.	en	Hulterström, Jens, Guidetti, Roberto, Jönsson, K. Ingemar, Atherton, Sarah (2025): Integrative taxonomy supports the description of two new species of Macrobiotidae (Tardigrada, Eutardigrada) from Kristianstads Vattenrike Biosphere Reserve in Sweden. European Journal of Taxonomy 1030: 1-52, DOI: 10.5852/ejt.2025.1030.3135, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3135/13991
7D3DFF4AFFE1FFA1582FFC7CCB0A2368.taxon	materials_examined	Material examined A total of 16 animals and 7 eggs observed, including: 10 animals and 6 eggs mounted in Hoyer’s fluid, 2 animals and 1 egg fixed for SEM, and 4 animals used for DNA extraction. Type material Holotype SWEDEN • Skåne, Sånnarna; 55 ° 55 ′ 41.6 ″ N, 14 ° 15 ′ 11.1 ″ E; 8 m a. s. l.; 22 Mar. 2021; S. Atherton, R. Guidetti and K. I. Jönsson leg.; moss on calcareous-rich sand and rock; SMNH, slide SMNH-Type- 10010. Paratypes SWEDEN • 9 specs, 5 eggs; same collection data as for holotype; GenBank nos: PX 093653 to PX 093655 (COI), PX 093644 (ITS), PX 093663, PX 093664 (18 S), PX 093649, PX 093650 (28 S); SMNH, slides SMNH-Type- 10011 to SMNH-Type- 10015, SEM stub SMNH-Type- 10016 • 2 specs, 2 eggs; same collection data as for holotype; MUSN, slides 24 - 081, 24 - 083.	en	Hulterström, Jens, Guidetti, Roberto, Jönsson, K. Ingemar, Atherton, Sarah (2025): Integrative taxonomy supports the description of two new species of Macrobiotidae (Tardigrada, Eutardigrada) from Kristianstads Vattenrike Biosphere Reserve in Sweden. European Journal of Taxonomy 1030: 1-52, DOI: 10.5852/ejt.2025.1030.3135, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3135/13991
7D3DFF4AFFE1FFA1582FFC7CCB0A2368.taxon	description	Description Morphometric measurements of animals and eggs are given in Tables 1 and 2, respectively (raw morphometric data in Supp. file 10). Adult body length 335 – 530 µm, yellowish with black eyespots visible in live animals (Fig. 4 A) and animals fixed in Hoyer’s. Cuticle with small (up to 1 µm in diameter), round pores, visible with DIC (Fig. 4 E) and clearly visible with SEM (Fig. 6 C), scattered throughout the body cuticle. Very fine granulation clearly visible only with SEM distributed over the entire body, less dense on the ventral side (Fig. 6 B – C). Medium granulation lateral and external to claws on legs I – III visible with DIC (Fig. 5 B) and SEM (Fig. 6 B). Legs IV with a patch of coarse granulation tapering centrally on the caudal segment and around the claws, clearly visible with DIC (Fig. 5 D), and SEM (Fig. 6 D). Pulvinus, gibbosities, and garter-like structures on the legs absent. Rigid buccal tube of Macrobiotus - type (Pilato & Binda 2010) with ventral lamina, ten peribuccal lamellae and six sensory lobes. OCA of hufelandi - type (Kaczmarek & Michalczyk 2017) with an anterior band of teeth faintly visible as fine granules with DIC (Fig. 4 C – D) and as small cones with SEM (Fig. 6 A); a second band of larger granular teeth and a third row of posterior transverse ridges. Third row non-uniform; ventral side comprising two lateral teeth and a divided median tooth split into two (Fig. 4 D, F), and dorsal side with a single, fused elongated tooth (Fig. 4 C). Stylet furca typical of the genus and stylet support insertion point at 75 – 81 % of the buccal tube. Bulbous muscular pharynx with three apophyses and three rows of two rodlike macroplacoids with the length series 1> 2 and a microplacoid (Fig. 4 B). First macroplacoid with slight central constriction. Second macroplacoid with deeper pre-terminal constriction. Comma-shaped microplacoid situated closer than its own length to the macroplacoid row. Claws of hufelandi - type (Bertolani & Pilato 1988) of similar size on all legs. Large lunulae on all claws, and distinctly larger on legs IV (max. width on leg I – III 6 – 8 µm; leg IV 7 – 10 µm). Lunulae smooth on legs I – III and serrated on legs IV, visible with DIC (Fig. 5 B – D), and SEM (Fig. 6 B, D – E). Cuticular bars absent. Gonochoristic. Males with sperm-filled testis and females with a spermatheca filled with bundles of sperm observed (Fig. 5 A). Ornamented eggs are spherical and laid freely, bare diameter 62.3 – 78.2 µm. Eggs of hufelandi - type (Kaczmarek & Michalczyk 2017; Figs 7 A, 8 A) with processes shaped as inverted goblets with strait or lightly sigmoidal trunks and terminal disks (Fig. 7 A – C). Surface of the terminal disks smooth, concave with a deep hollow and cogwheel-shaped edges including 10 – 11 evenly spaced indentations (Figs 7 D, 8). Process height 6.0 – 7.3 µm; base diameter 6.0 – 7.2 µm; terminal disk diameter 3.2 – 4.8 µm; and inter-process distance 2.9 – 4.2 µm. 23 – 25 processes on the egg circumference. The egg chorion is wrinkled and appears solid in DIC (Fig. 7 E – F), but minute (<0.25 µm in diameter) pores present around the bases of the processes are visible with SEM (Fig. 8 B – C). Larger wrinkles surrounding the base of each process appear as a crown of thickenings in DIC (Fig. 7 E – F). DNA sequences Sequences for M. vattenrikense sp. nov. were attained for all four molecular markers and using four adult animals. ITS 2 was represented by a single haplotype; 18 S and 28 S were represented by two haplotypes (uncorrected p-distance between haplotypes 0.06 % and 0.11 %, respectively), and COI was represented by three haplotypes (uncorrected p-distances between haplotypes 0.2 – 2.6 %): – 18 S haplotype 1: specimen 22 - 003; 1769 bp; GenBank accession number PX 093663; – 18 S haplotype 2: specimen 22 - 101; 1769 bp; GenBank accession number PX 093664; – 28 S haplotype 1: specimens 22 - 003, 22 - 017; 921 bp; GenBank accession number PX 093649; – 28 S haplotype 2: specimen 22 - 101; 921 bp, GenBank accession number PX 093650; – COI haplotype 1: specimen 22 - 017; 658 bp, GenBank accession number PX 093653; – COI haplotype 2: specimen 22 - 101; 658 bp; GenBank accession number PX 093654; – COI haplotype 3: specimens 22 - 003, 22 - 004 - 1; 658 bp; GenBank accession number PX 093655; – ITS 2: specimens 22 - 003, 22 - 017, 22 - 101; 443 bp; GenBank accession number PX 093644.	en	Hulterström, Jens, Guidetti, Roberto, Jönsson, K. Ingemar, Atherton, Sarah (2025): Integrative taxonomy supports the description of two new species of Macrobiotidae (Tardigrada, Eutardigrada) from Kristianstads Vattenrike Biosphere Reserve in Sweden. European Journal of Taxonomy 1030: 1-52, DOI: 10.5852/ejt.2025.1030.3135, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3135/13991
7D3DFF4AFFE1FFA1582FFC7CCB0A2368.taxon	diagnosis	Morphological differential diagnosis Macrobiotus vattenrikense sp. nov. has hufelandi - type OCA, serrated lunules on legs IV, and eggs with inverted goblet-shaped processes with terminal disks showing cogwheel-like indented margins and a deep central hollow, wrinkled chorions and very small pores surrounding the processes. Based on these characteristics, M. vattenrikense is most similar to five other species of Macrobiotus (note, the pores on the egg may be easily overlooked with light microscopy / clearly visible only with SEM, and so species with similar egg morphologies that have not been observed with SEM are included): Macrobiotus halophilus Fontoura, Rubal & Veiga, 2017 – the animals of M. vattenrikense sp. nov. differ from animals of M. halophilus in the wider buccal tube (external width pt 16.7 – 20.9 compared with pt 13.2 – 16.5 in M. halophilus); the lack of a deep constriction in the first macroplacoid; the paired ventromedial teeth and single fused dorsal tooth in the third band of the OCA (compared with an undivided ventromedial tooth and three distinct dorsal teeth in M. halophilus); the shorter claws (posterior claw IV pt 23.8 – 33.3 compared to 33.4 – 42.0 in M. halophilus); and the lack of male gibbosities. The eggs of M. vattenrikense are differentiated by the larger processes (height 6.0 – 7.3 µm compared to 3.9 – 5.9 µm in M. halophilus). In addition, the very small pores that are present surrounding the egg processes of M. vattenrikense have not been documented for M. halophilus, although eggs of the latter species have not been observed with SEM. Macrobiotus marlenae Kaczmarek & Michalczyk, 2004 – the animals of M. vattenrikense sp. nov. differ from the animals of M. marlenae by the granulation present on all legs (compared to granulation only on legs IV in M. marlenae); the shallower constriction of the first macroplacoid; the subterminal constriction in the second macroplacoid; and the claws with larger lunules in M. vattenrikense The eggs of M. vattenrikense have more numerous (23 – 25 on the circumference compared to 16 in M. marlenae) processes that are shorter (6.0 – 7.3 µm compared to 8.4 – 8.8 µm in M. marlenae), more closely spaced (inter-process distance 2.9 – 4.2 µm compared with 4.5 – 6.5 µm in M. marlenae), and with much smaller terminal disks (diameter 3.2 – 4.8 µm compared to 9.5 – 11.4 µm in M. marlenae). In addition, the very small pores that are present surrounding the egg processes of M. vattenrikense have not been documented for M. marlenae, although eggs of the latter species have not been observed with SEM. Macrobiotus ovovittatus Stec, 2024 – the animals of M. vattenrikense sp. nov. are smaller than animals of M. ovovittatus (335 – 530 µm compared to 570 – 879 µm in M. ovovittatus) and have a shorter first macroplacoids (pt 21.4 – 27.8 compared to pt 28.4 – 34.4 in M. ovovittatus) with shallower median incision. The new species is further distinguished by the paired ventromedial teeth and single fused dorsal tooth in the third band of OCA (compared with an undivided ventromedial tooth and three distinct dorsal teeth in M. ovovittatus), and the lack of cuticular bars below the claws of the first three pairs of legs (bars present in M. ovovittatus). The eggs of M. vattenrikense are smaller than those of M. ovovittatus (diameter without the processes 62.3 – 78.2 µm compared to 100.6 – 129.8 µm in M. ovovittatus) with fewer processes on the circumference (23 – 25 compared to 28 – 34 in M. ovovittatus), and smaller processes (height 6.0 – 7.3 µm compared to 9.5 – 13.5 µm in M. ovovittatus; diameter at the base 6.0 – 7.2 µm compared to 9.4 – 13.6 µm in M. ovovittatus) with smaller (diameter 3.2 – 4.8 µm compared to 6.1 – 8.7 µm in M. ovovittatus), solid terminal disks (covered by multiple light-refracting dots in M. ovovittatus). Macrobiotus persimilis Binda & Pilato, 1972 (following the redescription of Bertolani et al. 2023) – the animals of M. vattenrikense sp. nov. differ from animals of M. persimilis in the lack of a deep constriction in the first macroplacoid; the shorter macroplacoids (first / second macroplacoid pt 21.4 – 27.8 / 16.7 – 20.9 compared to pt 32.0 – 33.8 / 22.8 – 24.2 in M. persimilis) and macroplacoid row (pt 37.1 – 52.8 compared to pt 54.7 – 57.2 in M. persimilis); the longer microplacoids (pt 8.6 – 11.6 compared to pt 7.7 – 8.0 in M. persimilis); the paired ventromedial teeth and single fused dorsal tooth in the third band of the OCA (compared with an undivided ventromedial tooth and three distinct dorsal teeth in M. persimilis); and the shorter claws (external claw III / IV pt 23.3 – 33.3 / 25.6 – 30.6 compared to 35.9 – 38.3 / 35.9 – 37.9 in M. persimilis). The eggs of M. vattenrikense are differentiated by the larger processes (height 6.0 – 7.3 µm compared to 3.7 – 5.3 µm in M. persimilis; diameter at the base 6.0 – 7.2 µm compared to 3.6 – 5.1 µm in M. persimilis). In addition, the very small pores that are present surrounding the egg processes of M. vattenrikense have not been documented for M. persimilis, although eggs of the latter species have not been observed with SEM. Macrobiotus polonicus Pilato, Kaczmarek, Michalczyk & Lisi, 2003 – the animals of M. vattenrikense sp. nov. differ from the animals of M. polonicus by the presence of granulation lateral to the claws on legs I – III (absent in M. polonicus); the absence of lateral gibbosites on the hind legs (present in M. polonicus); the absence of sclerotized areas near the lunulae of legs I – III (present in M. polonicus). The eggs of M. vattenrikense differ by the smaller diameter of the process terminal disks (3.2 – 4.8 µm compared with 4.9 – 6.3 µm in M. polonicus) and by the minute pores surrounding each process (lack of egg pores for M. polonicus confirmed with SEM). Macrobiotus trunovae Biserov, Pilato & Lisi, 2011 – the animals of M. vattenrikense sp. nov. differ from animals of M. trunovae by the paired ventromedial teeth and single fused dorsal tooth in the third band of the OCA (compared with an undivided ventromedial tooth and three distinct dorsal teeth in M. trunovae); the more anteriorly inserted stylet supports (pt 75.0 – 81.0 compared to 81.7 – 82.4 in M. trunovae); the smaller first macroplacoid (pt 21.4 – 27.8 compared to 28.3 – 31.4 in M. trunovae) with a much narrower constriction; and the similar sizes of the claws on all legs (compared to distinctly smaller claws on legs I – III than on legs IV in M. trunovae). The eggs of M. vattenrikense are smaller (diameter without processes 62.3 – 78.2 µm compared to 134.3 µm in M. trunovae) with fewer (23 – 25 on the egg circumference compared to 32 in M. trunovae) and smaller (length 6.0 – 7.3 µm compared to up to 10.9 µm in M. trunovae; diameter at the base 6.0 – 7.2 µm compared to 9.1 – 9.9 µm in M. trunovae) processes. In addition, the very small pores that are present surrounding the egg processes of M. vattenrikense have not been documented for M. trunovae, although eggs of the latter species have not been observed with SEM.	en	Hulterström, Jens, Guidetti, Roberto, Jönsson, K. Ingemar, Atherton, Sarah (2025): Integrative taxonomy supports the description of two new species of Macrobiotidae (Tardigrada, Eutardigrada) from Kristianstads Vattenrike Biosphere Reserve in Sweden. European Journal of Taxonomy 1030: 1-52, DOI: 10.5852/ejt.2025.1030.3135, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3135/13991
7D3DFF4AFFF8FFA7584BF906C8EA2050.taxon	description	urn: lsid: zoobank. org: act: C 91427 F 4 - F 50 C- 4 A 1 F-A 475 - 825777258 AE 5 Figs 9 – 11	en	Hulterström, Jens, Guidetti, Roberto, Jönsson, K. Ingemar, Atherton, Sarah (2025): Integrative taxonomy supports the description of two new species of Macrobiotidae (Tardigrada, Eutardigrada) from Kristianstads Vattenrike Biosphere Reserve in Sweden. European Journal of Taxonomy 1030: 1-52, DOI: 10.5852/ejt.2025.1030.3135, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3135/13991
7D3DFF4AFFF8FFA7584BF906C8EA2050.taxon	diagnosis	Diagnosis Paramacrobiotus without eyespots. Cuticle with fine dorsal granulation visible only in SEM, and medium-coarse granulation surrounding the claws of the legs I – IV visible with DIC and SEM. OCA with three bands of teeth: first composed of numerous small granules, second of a single row of vertical ridges, and third of three ventral and three dorsal transverse crests. Three rod-like macroplacoids and a distant microplacoid present. Claws of hufelandi - type with smooth lunulae. Freely laid eggs with sculptured areolae of richtersi - type; 10 – 12 reticulated processes around the circumference, conical with a mostly smooth apex.	en	Hulterström, Jens, Guidetti, Roberto, Jönsson, K. Ingemar, Atherton, Sarah (2025): Integrative taxonomy supports the description of two new species of Macrobiotidae (Tardigrada, Eutardigrada) from Kristianstads Vattenrike Biosphere Reserve in Sweden. European Journal of Taxonomy 1030: 1-52, DOI: 10.5852/ejt.2025.1030.3135, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3135/13991
7D3DFF4AFFF8FFA7584BF906C8EA2050.taxon	etymology	Etymology The species is proudly in dedication to Marc “ Marchelmon ” Hulterström, brother of the first author.	en	Hulterström, Jens, Guidetti, Roberto, Jönsson, K. Ingemar, Atherton, Sarah (2025): Integrative taxonomy supports the description of two new species of Macrobiotidae (Tardigrada, Eutardigrada) from Kristianstads Vattenrike Biosphere Reserve in Sweden. European Journal of Taxonomy 1030: 1-52, DOI: 10.5852/ejt.2025.1030.3135, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3135/13991
7D3DFF4AFFF8FFA7584BF906C8EA2050.taxon	materials_examined	Material examined A total of 38 animals and 24 eggs observed, including: 16 animals and 15 eggs mounted in Hoyer’s fluid, 10 animals and 5 eggs fixed for SEM, and 11 animals and 4 eggs used for DNA analysis. Type material Holotype SWEDEN • Skåne, Sånnarna; 55 ° 55 ′ 41.6 ″ N, 14 ° 15 ′ 11.1 ″ E; 8 m a. s. l.; 22 Mar. 2021; S. Atherton, R. Guidetti and K. I. Jönsson leg.; moss on calcareous-rich sand and rock; SMNH, slide SMNH-Type- 10017. Paratypes SWEDEN · 23 specs, 15 eggs; same collection data as for holotype; SMNH, slides SMNH-Type- 10018 to SMNH-Type- 10029, SEM stub SMNH-Type- 10030 • 3 specs, 5 eggs; same collection data as for holotype; GenBank nos: PX 093656 to PX 093658 (COI), PX 093645, PX 093646 (ITS), PX 093665 (18 S), PX 093651 (28 S); MUSN, slides 22 - 030, 22 - 037 and 22 - 103.	en	Hulterström, Jens, Guidetti, Roberto, Jönsson, K. Ingemar, Atherton, Sarah (2025): Integrative taxonomy supports the description of two new species of Macrobiotidae (Tardigrada, Eutardigrada) from Kristianstads Vattenrike Biosphere Reserve in Sweden. European Journal of Taxonomy 1030: 1-52, DOI: 10.5852/ejt.2025.1030.3135, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3135/13991
7D3DFF4AFFF8FFA7584BF906C8EA2050.taxon	description	Description Morphometric measurements and statistics given in Tables 1 and 2 (raw morphometric data is given in Supp. file 11). Paramacrobiotus with body length 238 – 710 µm (Fig. 9 A). Eyespots absent. Cuticle transparent in live animals and after fixation in Hoyer’s fluid. Dorsal body cuticle with even and very fine granulation, visible only with SEM (Fig. 10 A – B). A patch of larger granulation lateral to the claws on legs I – III and coarse granulation fully surrounding the claws on legs IV visible with DIC and SEM (Figs 9 F, 10 B – D). Cuticular pores, gibbosities, and papillae absent. Mouth antero-ventral; ten peribuccal lamellae present. Buccal apparatus with a rigid tube of Macrobiotus - type (Pilato & Binda 2010) with ventral lamina. OCA comprising three bands of teeth (Fig. 9 C – D). First band a field of small granular teeth situated closely behind the peribuccal lamellae. Second band one row of vertical ridges, uniform in shape on both ventral and dorsal side. Third band situated close to the second, with three ventral and three dorsal teeth, with the lateral teeth wider (larger along the left / right axis) and shorter (smaller along the anterior / posterior axis) than the median tooth for each side. Ventral teeth thinner (smaller along the left / right axis) than dorsal teeth (Fig. 9 C). Latero-dorsal teeth triangular, narrowing away from the median tooth (Fig. 9 D). No additional granular teeth observed between the second and third band of OCA. Globular pharyngeal bulb with triangular pharyngeal apophyses, three rod-shaped macroplacoids with length sequence 2 <1 <3, and a microplacoid (Fig. 9 B). First macroplacoid drop-like. Second macroplacoid oval. Third macroplacoid with a subterminal constriction, ending in posterior bulbs. Microplacoid heart-shaped, with antero-lateral wings, and situated distant from the third macroplacoid, further than its length. Double-claws of hufelandi - type (Bertolani & Pilato 1988). Main claw branches with evident accessory points. Smooth lunulae on all claws (Figs 9 E, 10 B – D). Paired muscle attachments below claw present. Cuticular bars absent. Gonochoristic. Males with sperm filled testis observed, as well as females with eggs. Ornamented eggs laid freely, richtersi - type (Kaczmarek et al. 2017; Fig. 11 A – B), diameter without processes 67.0 – 92.4 µm. Processes conical, short (10 – 16.5 µm) and wide (21.3 – 30.0 µm), with a smooth rounded apex (Fig. 11 A – E). 10 – 12 processes on the circumference (Fig. 11 B). Processes smooth in SEM (Fig. 11 A) but labyrinthine layer visible in DIC as a reticular pattern of small, evenly distributed meshes (Fig. 11 C). Processes surrounded by a single ring of 12 areolae (Fig. 11 A, G). Areolae with sculpturing comprising dotlike indentations, faintly visible fixed in Hoyer’s with DIC and clearly visible with SEM (Fig. 11 F). DNA sequences Sequences for P. marchelmoni sp. nov. were attained for all four molecular markers using 11 animals and four eggs. 18 S and 28 S were represented by one haplotype; ITS 2 was represented by two haplotypes (uncorrected p-distances between haplotypes 0.81 %), and COI was represented by three haplotypes (uncorrected p-distances between haplotypes 0.16 – 0.31 %): – 18 S: specimens 22 - 004, 22 - 006, 22 - 007, 22 - 019, 22 - 034, 22 - 110, 22 - 113; 1302 bp, GenBank accession number PX 093665; – 28 S: specimens 22 - 004, 22 - 019, 22 - 029, 22 - 110, 23 - 015; 921 bp, GenBank accession number PX 093651; – COI haplotype 1: specimens 22 - 004, 22 - 006, 22 - 007, 22 - 019, 22 - 026, 22 - 029, 22 - 034, 22 - 102, 22 - 108; 636 bp, GenBank accession number PX 093656; – COI haplotype 2: specimens 22 - 106, 22 - 110, 22 - 113, 23 - 041; 636 bp, GenBank accession number PX 093657; – COI haplotype 3: specimen 23 - 015, 636 bp, GenBank accession number PX 093658; – ITS 2 haplotype 1: specimens 22 - 029, 22 - 102; 370 bp; GenBank accession number PX 093645; – ITS 2 haplotype 2: specimens 22 - 004, 22 - 006, 22 - 007, 22 - 019, 22 - 034, 22 - 110, 22 - 113; 370 bp, GenBank accession number PX 093646.	en	Hulterström, Jens, Guidetti, Roberto, Jönsson, K. Ingemar, Atherton, Sarah (2025): Integrative taxonomy supports the description of two new species of Macrobiotidae (Tardigrada, Eutardigrada) from Kristianstads Vattenrike Biosphere Reserve in Sweden. European Journal of Taxonomy 1030: 1-52, DOI: 10.5852/ejt.2025.1030.3135, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3135/13991
7D3DFF4AFFF8FFA7584BF906C8EA2050.taxon	diagnosis	Morphological differential diagnosis Based on the presence of a microplacoid, P. marchelmoni sp. nov. is a member of the richtersi morphogroup (Kaczmarek et al. 2017). Species of this morphogroup may be difficult or sometimes impossible to distinguish morphologically, with distinguishing features frequently limited to a few differences in egg shell morphologies or reproductive biology (Guidetti et al. 2019 a). Indeed, P. marchelmoni is very similar to ten species of this group by the following criteria: richtersi - type of egg, with areolae with sculpturing (Kaczmarek et al. 2017), and egg processes with a blunt apex without a cap-like structure. However, P. marchelmoni can be differentiated from each by the following: Paramacrobiotus arduus Guidetti, Cesari, Bertolani, Altiero & Rebecchi, 2019 – the animals of P. marchelmoni sp. nov. lack cuticular bars below the claws of the first three pairs of legs. The eggs of the new species are larger (diameter without processes 67.0 – 92.4 µm compared to 55.3 – 62.3 µm in P. arduus), with processes that are surrounded by more areolae (12 compared with 5 “ double ” areolae in P. arduus) and that are much wider at the base (diameter 21.3 – 30.0 µm compared to 10.4 – 16.3 µm in P. arduus) with a more rounded and less narrow apex than those of P. arduus. Paramacrobiotus celsus Guidetti, Cesari, Bertolani, Altiero & Rebecchi, 2019 – the animals of P. marchelmoni sp. nov. lack cuticular bars below the claws of the first three pairs of legs, and the microplacoid has a different shape (heart-shaped with antero-lateral wings compared to drop-shaped in P. celsus). Additionally, the eggs of P. marchelmoni have processes with wider bases (diameter 21.3 – 30.0 µm compared to 14.3 – 18.2 µm in P. celsus). Paramacrobiotus depressus Guidetti, Cesari, Bertolani, Altiero & Rebecchi, 2019 – the animals of P. marchelmoni sp. nov. have much more evident granulation on all legs compared with P. depressus and lack cuticular bars below the claws of the first three pairs of legs. The microplacoid has a different shape (heart-shaped with antero-lateral wings compared to comma-shaped in P. depressus). The eggs are larger in P. marchelmoni (diameter without processes 67.0 – 92.4 µm compared to 56.2 – 66.2 µm in P. depressus) with much wider (21.3 – 30.0 µm compared to 12.4 – 15.2 µm in P. depressus) egg processes. Paramacrobiotus fairbanksi (following the redescription of Guidetti et al. 2019 a) – animals of P. marchelmoni sp. nov. differ from animals of P. fairbanksi in the heart-shaped microplacoid (compared to a comma-shape microplacoid in P. fairbanksi), and the new species is gonochoristic (males are presumed absent for P. fairbanski). The eggs of P. marchelmoni have more areolae surrounding each process (12 compared with 5 – 6 “ double ” areolae in P. fairbanksi) and processes with wider bases (21.3 – 30.0 µm compared to 10.9 – 20.8 µm in P. fairbanksi) and smoother, more rounded apexes. Paramacrobiotus gerlachae (Pilato, Binda & Lisi, 2004) – the animals of P. marchelmoni sp. nov. differ from the animals of P. gerlachae by the absence of cuticular bars near the lunules of legs I – III (present in P. gerlachae); by the cuticular granulation on legs I – III (absent in P. gerlachae); and by the more distinct granulation on leg IV (granulation very faint in P. gerlachae). The egg processes are wider in P. marchelmoni (base diameters 21.3 – 30 µm compared with 16.8 – 18.7 µm in P. gerlachae). Paramacrobiotus halei (Bartels, Pilato, Lisi & Nelson, 2009) – animals of P. marchelmoni sp. nov. differ from animals of P. halei by the absence of the cuticular tubercles (present in P. halei); the narrower buccal tube (pt 18.1 – 23.8 compared with pt 26.5 – 28.6 in P. halei); and the longer internal claw II (pt 21.4 – 30.0 compared with pt 20.4 – 21.1 in P. halei). The eggs of P. marchelmoni are smaller (full diameter 92.5 – 115.0 µm compared with 117.0 – 121.0 µm in P. halei) with processes with smoother, more rounded apexes. Paramacrobiotus pius Lisi, Binda & Pilato, 2016 – animals of P. marchelmoni sp. nov. differ from animals of P. pius in the lack of cuticular bars below the claws on the first three pair of legs (present in P. pius), the longer third macroplacoid (pt 18.3 – 29.4 compared to 14.2 – 14.6 in P. pius) and the longer macroplacoid / placoid rows (pt 52.1 – 75.7 / 69.5 – 95.2 compared to pt 41.4 – 43.1 / 53.0 – 56.2 in P. pius). The eggs of P. marchelmoni have processes with smooth apices (compared to apices with short spines in P. pius) and are surrounded by more numerous areolae (12 compared to 5 “ double ” areolae in P. pius). Paramacrobiotus richtersi (Murray, 1911) (following the redescription of Guidetti et al. 2019 a) – the animals of P. marchelmoni sp. nov. differ from the animals of P. richtersi in the undivided medioventral crest in the third band of the OCA and the lack of cuticular bars below the claws of the first three pairs of legs (bars present in P. richtersi). The eggs of P. marchelmoni differ in the processes, which are shorter (10.0 – 16.5 µm compared to 17.1 – 22.1 µm in P. richtersi) and wider (diameter 21.3 – 30.0 µm compared to 17.1 – 21.2 µm in P. richtersi) and thus have a stouter appearance than those of P. richtersi. Paramacrobiotus sklodowskae (Michalczyk, Kaczmarek & Węglarska, 2006) – the animals of P. marchelmoni sp. nov. differ from the animals of P. sklodowskae by the absence of eyespots (present in P. sklodowskae); by the more anteriorly inserted stylet support (pt 72.3 – 80.6 compared with pt 81.8 – 85.2 in P. sklodowskae); and by the larger third macroplacoid (pt 18.3 – 29.4 compared to pt 16.7 – 18.0 in P. sklodowskae). The eggs are differentiated by the process surfaces, which are always smooth in P. marchelmoni (ring folds present in P. sklodowskae). Paramacrobiotus spatialis Guidetti, Cesari, Bertolani, Altiero & Rebecchi, 2019 – the animals of P. marchelmoni sp. nov. possess an undivided medioventral crest in the third band of the OCA (subdivided into three round teeth in P. spatialis). The eggs of P. marchelmoni have wider processes (21.3 – 30.0 µm compared to 15.2 – 20.4 µm in P. spatialis) with smooth apices (compared to apices with tubercles as in P. spatialis).	en	Hulterström, Jens, Guidetti, Roberto, Jönsson, K. Ingemar, Atherton, Sarah (2025): Integrative taxonomy supports the description of two new species of Macrobiotidae (Tardigrada, Eutardigrada) from Kristianstads Vattenrike Biosphere Reserve in Sweden. European Journal of Taxonomy 1030: 1-52, DOI: 10.5852/ejt.2025.1030.3135, URL: https://europeanjournaloftaxonomy.eu/index.php/ejt/article/download/3135/13991
