taxonID	type	description	language	source
03E8C872FF9B0848FC49FBB628ACFE10.taxon	description	Dactylobiotus taiwanensis sp. nov. (Figs. 2 – 9, Tables 3 – 4) urn: lsid: zoobank. org: act: 2 F 270225 - 1415 - 4690 - B 8 E 8 - 7673 F 0 FF 4131 Material examined: 82 animals, 73 eggs mounted on microscope slides in Hoyer’s medium (some of the eggs were embryonated), six animals and two eggs examined in SEM and two specimens processed for DNA sequencing. Type locality: 25.0506021, 121.4628704; 3 m asl: Xinzhuang Touqian Sports Park, New Taipei City, Taiwan; debris from the leaf of Limnophila sp.; coll. Chih-Yu Pai; 1 December 2022. Etymology: The species is named after the country in which it was discovered. Type depositories: Holotype: slide TW. 001.11 and 46 paratypes (slides: TW. 001. *, where the asterisk can be substituted by any of the following numbers: 04, 05, 07 – 10, 12, 13) and 59 eggs (slides: TW. 001. *: 01, 14 – 17) are deposited at the Institute of Systematics and Evolution of Animals, Polish Academy of Sciences, Sławkowska 17, 31 - 016, Kraków, Poland, whereas 35 paratypes (slides: TW. 001. *: 03, 06) and 14 eggs (slide: TW. 001.02) deposited in the Biodiversity Research Center of Academia Sinica. 6 animals and 3 eggs prepared for SEM (UNICT-Stub N. 67) are deposited at the University of Catania, Italy. DNA voucher: Two exoskeletons mounted on permanent slides, labelled Dac. tai. _ TW. 001.01 and Dac. tai. _ TW. 001.02, are deposited at the Institute of Systematics and Evolution of Animals, Polish Academy of Sciences, Sławkowska 17, 31 - 016 Kraków, Poland. Description of the new species Animals (measurements and statistics in Table 3): Body transparent in juveniles and whitish in adults, but transparent after fixation in Hoyer’s medium (Fig. 2 A). In live specimens, eyes are present but they dissolve in Hoyer’s medium (out of 21 measured animals). In the dorso-lateral head region an area with minute pores (probably chemosensory function) can be identified in both sides of the head, but only with SEM (Fig. 3 A – B). Other than that cuticle is without typical pores but wrinkled with two flat, oval papillae present on the dorsum between legs III and IV in adults and juveniles (Figs. 2 B – C and 3 C – D). Granulation absent on all legs. Claws of the Dactylobiotus type with short basal portion and primary branches with distinct accessory points (Figs. 4 and 5). Lunules absent, but under PCM a robust semilunar cuticular connection is present between external / posterior and internal / anterior claws (Fig. 4). Under SEM this connection is visible as discontinuous, being composed of extended lunulelike thickenings under the claws on the lateral sides whereas its median portion is located within or under cuticle (Fig. 5). Claws on the first three pairs of legs similar in size but obviously larger on the hind legs. A cuticular thickening is present above claws I – III (Fig. 4 C and 5 A), which under PCM is visible a darkened continuous cuticular bar (Fig. 4 C). Under PCM the area above claws IV is darkened (4 D), being similar to the darkened area present in horseshoe structure connecting the anterior and the posterior claw in many species of the family Macrobiotidae. The cuticle of this area under SEM appears smooth when compared to the surrounding cuticle (Fig. 5 C). Mouth antero-ventral followed by ten short peribuccal lamellae, bucco-pharyngeal apparatus of the Macrobiotus type (Fig. 3 A, 6 A and 7). Under PCM, only the second and third bands of teeth are visible in the oral cavity armature (Fig. 6 B – C). However, in SEM three bands of teeth are clearly visible with the first band being situated at the base of peribuccal lamellae and composed of several rows of scattered small conical teeth arranged around the oral cavity (Fig. 7). The second band of teeth is situated below the ring fold, and comprises 4 – 6 rows of small cone-shaped teeth which are larger than those of the first band and increase in size towards the third band of teeth (Figs. 6 B – C and 7). The teeth of the third band are located within the posterior portion of the oral cavity, between the second band of teeth and the buccal tube opening (Figs. 6 B – C and 7 B – C). The third band of teeth is discontinuous and divided into dorsal and the ventral portions. Under PCM, the dorsal teeth are seen as three distinct transversal ridges whereas the ventral teeth appear as two separate lateral transverse ridges, between which a roundish median tooth is visible (Fig. 6 B – C). In SEM, both dorsal and ventral teeth are also clearly distinct (Fig. 7 B – C). Under PCM, in the lateral view of the buccal apparatus, a strengthening bar (ventral lamina) with an incision determining a ventral hook is clearly visible (Fig. 6 D). Pharyngeal bulb spherical, with triangular apophyses, two rod-shaped macroplacoids which sometimes have jagged edges (Fig. 6 E – G). The macroplacoid length sequence 2 <1. The first macroplacoid has a central constriction, whereas the second macroplacoid is only gently constricted sub-terminally (Fig. 6 E – G). Eggs (measurements and statistics in Table 4): Laid freely, whitish, spherical (Fig. 9 A). Processes in the shape of short and wide cones with apexes usually divided into multiple (typically three to six) short, nodular, finger-like apices (Figs. 8 and 9). Under SEM, apices usually covered with microgranulation (Fig. 9 C). The egg surface between the processes appears wrinkled; however, this is barely visible under PCM (Fig. 8 C, D), where most often the surface appears to be smooth (Fig. 8 A, B), whereas wrinkles are clearly distinguishable under SEM imaging (Fig. 9). Under PCM, the margins of the process bases appear serrated and are surrounded by a crown of faint, small thickenings / projections, usually with faintly visible pores (Fig. 8 A – D). Under SEM these dark projections are clearly visible as vertical thickenings present on basal portions of processes walls and each process base is surrounded by a line of around 25 small, but evident pores (Fig. 9). Eggs are sticky because they are covered by mucus which most likely enhances their adhesion to the substrate and maybe has also a protective function. This mucus is sometimes visible under SEM as a web of flexible filaments that cover the egg surface (Fig. 9 C).	en	Camarda, Daniele, Pai, Chih-Yu, Kristensen, Reinhardt Møbjerg, Stec, Daniel (2025): Positive association between PTN polymorphisms and schizophrenia in Northeast Chinese Han population. Zoological Studies 64 (14): 141-149, DOI: 10.6620/ZS.2025.64-14, URL: http://dx.doi.org/10.1097/YPG.0000000000000262
03E8C872FF9B0848FC49FBB628ACFE10.taxon	diagnosis	Differential diagnosis and PCA results Currently, there are four other species of Dactylobiotus that possess dorsal papillae: D. selenicus, D. dispar (Murray, 1907), D. parthenogeneticus, and D. grandipes. However, the new species differs specifically from these taxa as follows: Dactylobiotus dispar: The new species has two flat dorso-caudal papillae, whereas D. dispar exhibits two large conical papillae in the caudal region of the body. The new species also shows less pronounced constriction in the first macroplacoid. While D. dispar was described as having three macroplacoids, it is more likely to have only two, with the first one being profoundly constricted. Additionally, the eggs of the new species have interprocess distances smaller than the widths of the process bases, whereas in D. dispar, the interprocess distances are equal to the widths of the process bases.	en	Camarda, Daniele, Pai, Chih-Yu, Kristensen, Reinhardt Møbjerg, Stec, Daniel (2025): Positive association between PTN polymorphisms and schizophrenia in Northeast Chinese Han population. Zoological Studies 64 (14): 141-149, DOI: 10.6620/ZS.2025.64-14, URL: http://dx.doi.org/10.1097/YPG.0000000000000262
03E8C872FF9B0848FC49FBB628ACFE10.taxon	description	Dactylobiotus grandipes: The new species differs by having two flat dorso-caudal papillae compared to the two large conical papillae in D. grandipes. The posterior primary claws on leg IV have lower pt values in the new species (pt 50.4 – 66.5) than in D. grandipes (pt 70.7 – 89.2). The eggs of the new species have fewer processes around the circumference (38 – 42 compared to 50 – 57 in D. grandipes) and possess process bases surrounded by a crown of faint, small thickenings or projections, usually with faintly visible pores. These projections and pores are absent or not visible in D. grandipes. Dactylobiotus parthenogeneticus: The new species has egg process bases surrounded by numerous pores (approximately 25) that are faintly visible under light microscopy, but well-visible with SEM. In contrast, D. parthenogeneticus exhibits scarce, small, singular pores visible only with SEM. The egg processes of the new species have generally wider bases (5.0 – 7.2 µm compared to 3.1 – 5.2 µm in D. parthenogeneticus) and a smaller interprocess distance (1.2 – 2.7 µm compared to 2.0 – 4.8 µm in D. parthenogeneticus). Although the ranges of egg measurements slightly overlap, these measurements differentiate the species most effectively (see the PCA results section below). Dactylobiotus selenicus: The new species has conical egg processes, whereas D. selenicus has truncoconical or crater-shaped processes. The eggs of the new species also have fewer processes around the circumference (38 – 42 compared to approximately 60 in D. selenicus). Additionally, the claws of the new species exhibit evident accessory points, which are absent or not visible under PCM in D. selenicus. Among the four species, D. parthenogeneticus is the most similar to the new species. Given that an abundant population of D. parthenogeneticus was recently studied by Pogwizd and Stec (2020), we utilized their data to perform more detailed comparisons using PCA analysis. The PCA analysis revealed partial overlap in absolute and relative (pt) morphometric data for animals of D. taiwanensis sp. nov. and D. parthenogeneticus. However, it showed a clear separation (no overlap) in egg morphometric data between the two species (Fig. 10). For the absolute measurement dataset, PC 1 explained 87.89 % of the variance, while PC 2 explained 3.63 %. Almost all morphometric traits contributed similarly to PC 1, except for body length and ventral lamina length, which contributed more to PC 2. For relative measurements (pt values), the loadings of the traits were more dispersed compared to absolute measurements. Body length and ventral lamina length again contributed more to PC 2 than PC 1, with PC 1 explaining 52.88 % of the variance and PC 2 explaining 9.78 %. In the PCA analysis of egg morphometric data, the two species were clearly separated. The PC 1 explained 70.79 % of the variance, while PC 2 explained 13.12 %. The egg morphometric traits contributing most to this separation were the number of processes around the egg circumference (positive), the ratio of process base width to height (positive), and the interprocess distance (negative). Other traits contributed similarly to PC 1 and PC 2, except for process height, which contributed more to PC 2 than PC 1.	en	Camarda, Daniele, Pai, Chih-Yu, Kristensen, Reinhardt Møbjerg, Stec, Daniel (2025): Positive association between PTN polymorphisms and schizophrenia in Northeast Chinese Han population. Zoological Studies 64 (14): 141-149, DOI: 10.6620/ZS.2025.64-14, URL: http://dx.doi.org/10.1097/YPG.0000000000000262
03E8C872FF93084FFC28FEF62FE4F837.taxon	description	(Figs. 11 – 12) Material examined: Four animals and 50 eggs were extracted from the sample collected in Greenland. The DNA sequences were obtained from one embryonated egg (Table 2). Fifteen eggs were prepared and examined under SEM (SEM stub: TAR. 19), while the remaining specimens were mounted on microscope slides in Hoyer’s medium (Slides: GL. 004.01 – 05). Material depositories: All specimens are deposited at the Institute of Systematics and Evolution of Animals, Polish Academy of Sciences, Sławkowska 17, 31 - 016, Kraków, Poland. Remarks: The specimens and eggs found in this sample resemble those of D. ampullaceus (Thulin, 1911) and D. octavi. Although the eggs of D. ampullaceus exhibit processes similar to those observed in the newly discovered Greenlandic population, they lack the pores around the process bases (Thulin 1911) that are clearly visible under light microscopy in our population as well as in D. octavi. Consequently, the examination of the prepared specimens allowed their identification as D. cf. octavi, as all observed characteristics matched the original description of animals and eggs, except for some details in egg ornamentation. Guidetti et al. (2006) noted that the egg processes in D. octavi are crater-like, often featuring a small elevation at the center. Specifically, most processes are conical with introverted apices, while only a few are fully conical. In contrast, the eggs from the newly discovered population analyzed in this study are distinctly conical in shape (Figs. 11 and 12). Their distal parts form a broad cone, tapering proximally into a short, slender tip (Figs. 11 and 12). Notably, some processes appeared not fully extended, giving the impression of introverted apices (Fig. 12 A, C). Other investigated taxa: The examination of the type material for D. caldarellai and D. lombardoi, along with their original descriptions, revealed no reliable characters to differentiate these species from their congeners. Furthermore, as eggs are unknown for these species and the type specimens are in very poor condition (SM. 3), we designate them as nomina dubia with the following combinations: Dactylobiotus caldarellai Pilato and Binda, 1994 nom. dub., and Dactylobiotus lombardoi Binda and Pilato, 1999 nom. dub. The challenges surrounding the identification of these three Dactylobiotus taxa are further elaborated and discussed in detail in the corresponding discussion section below.	en	Camarda, Daniele, Pai, Chih-Yu, Kristensen, Reinhardt Møbjerg, Stec, Daniel (2025): Positive association between PTN polymorphisms and schizophrenia in Northeast Chinese Han population. Zoological Studies 64 (14): 141-149, DOI: 10.6620/ZS.2025.64-14, URL: http://dx.doi.org/10.1097/YPG.0000000000000262
03E8C872FF910853FC93FEF62ECAFE30.taxon	description	Animals - rela � ve values (pt) Eggs D. parthenogeneticus D. taiwanensis sp. nov. exhibit dorsal papillae (D. grandipes, D. selenicus, D. parthenogeneticus, and D. taiwanensis sp. nov.) cluster together to form a well-supported clade. However, the relationship of this clade with other Dactylobiotus taxa remains poorly supported in our analyses (Fig. 1).	en	Camarda, Daniele, Pai, Chih-Yu, Kristensen, Reinhardt Møbjerg, Stec, Daniel (2025): Positive association between PTN polymorphisms and schizophrenia in Northeast Chinese Han population. Zoological Studies 64 (14): 141-149, DOI: 10.6620/ZS.2025.64-14, URL: http://dx.doi.org/10.1097/YPG.0000000000000262
03E8C872FF8F0850FC93FF362F6EFAD3.taxon	description	Intraspecific variability in egg morphology has been previously documented in several other tardigrade genera, such as Bertolanius Özdikmen, 2008 (primarily variability in the apical portion of the processes; Dastych 1983), Ramazzottius Binda and Pilato, 1986 (mainly in process length and shape; Stec et al. 2016 2017; Vecchi and Stec 2024), and Paramacrobiotus Guidetti, Schill, Bertolani, Dandekar and Wolf, 2009 (notably in minor differences in the shape of the processes, sometimes even within the same egg; Guidetti et al. 2019). However, an extreme case was recently reported in the latter genus for Paramacrobiotus bifrons Brandoli, Cesari, Massa, Vecchi, Rebecchi and Guidetti, 2024, which exhibits two morphologically distinct egg forms. Regarding the considerable intraspecific variability of egg ornamentation in Dactylobiotus, this phenomenon has so far been reported only for D. ovimutans. Eggs of this species, thoroughly examined in a culture maintained under stable laboratory conditions by Kihm et al. (2020), exhibited variability in the number, size, and inflation of egg processes. This variability was unlikely to result from seasonality or the production of dormant or active eggs, given the consistent conditions under which the culture was maintained (Kihm et al. 2020). Thus, it cannot be excluded that a similar variability may also occur in D. octavi, given the correspondence in all egg characters between the type population and the newly examined population, except for the number and inflation of processes. Specifically, fewer and less inflated processes were observed in the type population from Greenland (Guidetti et al. 2006), whereas eggs with more processes, which were always well extended, were found in the Greenlandic population studied here. Interestingly, differences in the shape of egg processes, attributed to developmental stages, have also been reported for Paramacrobiotus derkai (Degma, Michalczyk and Kaczmarek 2008) and P. bifrons (Brandoli et al. 2024; Degma et al. 2008). This suggests that similar situations might occur in other genera as well. Given these uncertainties and the inability to compare DNA sequences of variable markers from the population studied here with those of D. octavi, we classify our population as D. cf. octavi until further data become available. After examining the type material of D. caldarellai and reviewing its original description, we concluded that this species is insufficiently diagnosed. Pilato and Binda (1994) described D. caldarellai based on two specimens collected from two different locations in Tierra del Fuego, without finding any eggs. The authors considered a morphologically identical population of D. ambiguus reported by Dastych (1984) on King George Island as conspecific with D. caldarellai. Their rationale was based on Dastych’s (1984) observation that the eggs of the newly found population of D. ambiguus differed from those of the population from the species’ type locality in Europe. Notably, the two locations (Tierra del Fuego (type locality of D. caldarellai) and King George Island) are more than 300 km apart. Given the morphological uniformity among animals of different species within the genus and the absence of egg description for D. caldarellai, it cannot be confidently determined whether these populations represent the same or different species. Furthermore, the suboptimal condition of the holotype and paratype of D. caldarellai hinders a detailed examination (SM. 3). Therefore, until further analyses and a potential integrative redescription based on material from the locus typicus are conducted, D. caldarellai should be considered a nomen dubium, as designated in the results section. For the second dubious species, D. lombardoi, the original description was based on two specimens also collected in Tierra del Fuego. In their study, Binda and Pilato (1999) provided a table of pt values derived from the measurements of a single specimen of D. lombardoi. These values fall within the pt range of the most similar species, D. parthenogeneticus, as reported in the same paper, with the exception of buccal tube width and ventral lamina length. However, it is likely that these small differences are the result of an insufficient number of measured specimens or variations in measurement techniques used by different authors. Importantly, when the original measurements from Binda and Pilato (1999) are compared with those provided for D. parthenogeneticus by Pogwizd and Stec (2020), the specimen falls perfectly within the newly reported measurement ranges. Although dorsal papillae are not mentioned in the original description of D. lombardoi, it has been suggested that this species can be distinguished from D. parthenogeneticus by the absence (D. lombardoi) or presence (D. parthenogeneticus) of these structures. However, despite the poor condition of the type series, it appears that dorsal papillae may indeed be present in specimens of D. lombardoi (SM. 3). Given the vague diagnosis of this species, which prevents its clear distinction from other congeners, the lack of a population with a sufficient number of specimens, the unknown egg morphology, and the poor condition of the type material, D. lombardoi should be considered a nomen dubium. This designation, as outlined in the results section, will remain until further analyses confirm whether it represents a distinct species. As a result, all species for which eggs have never been found (i. e., D. aquatilis, D. caldarellai, D. henanensis, D. kansae, D. lombardoi, D. macronyx), with the exception of D. haplonyx Maucci, 1981, are considered nomina dubia. Regarding D. haplonyx, although no specimens from the type series have been examined, it is important to note that the type series comprises individuals collected from multiple locations, with the holotype originating from a different locality than the paratypes (a total of 33 paratypes from five distinct locations, all different from that of the holotype). Furthermore, the species description lacks not only an account of the eggs but also distinctive diagnostic traits that clearly differentiate it from other species. Consequently, further analyses are necessary to assess its validity. Therefore, we consider it appropriate to designate this species as a nomen inquirendum with the following combination: Dactylobiotus haplonyx Maucci, 1981 nom. inq. Dichotomous key Given that egg ornamentation often comprises fundamental diagnostic characters for distinguishing Dactylobiotus taxa, the presented key includes only valid species for which eggs have been described (12 species), excluding from the key all the species designated as nomina dubia or nomina inquirenda.	en	Camarda, Daniele, Pai, Chih-Yu, Kristensen, Reinhardt Møbjerg, Stec, Daniel (2025): Positive association between PTN polymorphisms and schizophrenia in Northeast Chinese Han population. Zoological Studies 64 (14): 141-149, DOI: 10.6620/ZS.2025.64-14, URL: http://dx.doi.org/10.1097/YPG.0000000000000262
03E8C872FF8F0850FC93FF362F6EFAD3.taxon	description	** The measurement was obtained by proportionally scaling a drawing from the species description by Murray (1907), which reported the egg diameter, including processes, as 130 µm. Additionally, data from Thulin (1911) indicated that the process bases had a diameter of 9.5 µm. *** It must be stressed that further investigations are necessary to confirm whether the processes can also have a conical shape. For more details, please refer to the section on Dactylobiotus cf. octavi in this study, which discusses the shape of egg processes in Dactylobiotus octavi.	en	Camarda, Daniele, Pai, Chih-Yu, Kristensen, Reinhardt Møbjerg, Stec, Daniel (2025): Positive association between PTN polymorphisms and schizophrenia in Northeast Chinese Han population. Zoological Studies 64 (14): 141-149, DOI: 10.6620/ZS.2025.64-14, URL: http://dx.doi.org/10.1097/YPG.0000000000000262
