taxonID	type	description	language	source
001087910D06FF9AD399FE75FB44F80C.taxon	description	Locus typicus. Moss on carbonate bedrock from Homole Ravine in the Pieniny Mts. (49 ° 24 ' 17 '' N, 20 ° 32 ' 52 '' E; 595 m asl), Poland.	en	Gąsiorek, Piotr, Degma, Peter (2018): Three Echiniscidae species (Tardigrada: Heterotardigrada) new to the Polish fauna, with the description of a new gonochoristic Bryodelphax Thulin, 1928. Zootaxa 4410 (1): 77-96, DOI: 10.11646/zootaxa.4410.1.4
001087910D06FF9AD399FE75FB44F80C.taxon	materials_examined	Type material. Holotype (mature female, slide no. PL. 189.19), coll. Maciej Barczyk, 25 th May 2016, allotype (mature male, slide no. PL. 189.01), 30 paratypes (21 mature females, 7 males, one juvenile, and one larva; slides PL. 189.02 – 32) and additional 15 paratypes (both juveniles and adults) on SEM stubs. Found together with numerous Bryodelphax parvulus Thulin, 1928 (Fig. 24), Pseudechiniscus suillus (Ehrenberg, 1853), and single specimens of Testechiniscus spitsbergensis (Scourfield, 1897). All from the same moss sample. Holotype and most of paratypes deposited in the Institute of Zoology and Biomedical Research, Jagiellonian University, Kraków, Poland, two paratypes (slides PL. 189.10 – 11) deposited in the Department of Zoology, Comenius University in Bratislava, Slovakia, and two paratypes (slides PL. 189.22 – 23) deposited in the Zoological Museum, University of Copenhagen, Denmark. Additional material examined. Two specimens in a moss sample from Chuda Alp in the Tatra Mts. (slide PL. 258.06), coll. Piotr Gąsiorek, 24 th September 2016, deposited together with type material. 128 specimens (44 males, 43 females, 3 juveniles, and 38 specimens of unidentified sex due to unsuitable orientation of a specimen on a slide; slides 390 / 7 – 10, 390 / 18, 390 / 20, 390 / 23, 390 / 25, 390 / 27 – 30, and 390 / 32; deposited in the Department of Zoology, Comenius University in Bratislava, except for 13 specimens deposited in the Institute of Zoology and Biomedical Research, Jagiellonian University, Kraków — slide 390 / 18) and 9 specimens used for SEM analyses (not deposited) in a moss sample on a rock from Rajtopíky hill, the Branisko Mts. (48 ° 59 ' 42 '' N, 20 ° 51 ' 59 '' E; ca. 1036 m asl — on the hill top area), Eastern Slovakia, coll. Peter Degma, 26 th July 2003. Short description of holotype (Figs 1, 10; Table 3) Adult female; colour greyish; eyes not visible (after preparation); body length 137 µm (685); cephalic cirri and papillae, and clava not enlarged; cirrus A 26.5 µm (132.5); claws I – III of similar size, claw IV longer; dentate collar present, with three teeth; small papilla on leg IV. Dorsal plates with sparse pseudopores / pores; ventral plates not obvious. Female gonopore rosette-shaped. Short description of allotype (Figs 2, 12; Table 4) Adult male; colour greyish; eyes not visible (after preparation); body length 123 µm (628); cephalic cirri and papillae, and clava enlarged; cirrus A 20.7 µm (103.0); claws I – III of similar size, claw IV longer; dentate collar present, with three teeth; small papilla on leg IV not visible. Dorsal plates with sparse pseudopores / pores; suggestion of deeper faceting on scapular and caudal plates; ventral plates present. Male gonopore simple, circular. Details of the new species. Adults (i. e. from the third instar onwards; measurements and statistics for ♀♀ in Table 3, for ♂♂ in Table 4): Body greyish; eyes absent or not visible after preparation. Mouth opening surrounded by a ring of 10 papulae (visible in SEM only). Cephalic papillae greatly enlarged in males (compare Figs 1 and 2, 4), and clavae are also more prominent (Figs 13, 17 – 18). Internal cirri much shorter than external (Tables 3 – 4, Figs 1 – 4, 7 – 9). Cirri A thinner at the tip (visible only under SEM; Fig. 17). Dorsal plates covered with sparsely distributed pseudopores or pores of large and intermediate size (visible in PCM as bright dots with margins either blurred (pseudopores) or obvious (pores); Figs 1 – 2, 10, 12), true pores occur infrequently, being restricted to the posterior portion of the cephalic plate and the anterior portion of the scapular plate (Fig. 13) in the proximal body part, to the posterior parts of segmental plates (Fig. 14), and the anterior margin of the caudal plate in the distal body part. Pores rarely present on all plates (Figs 11, 15 – 16), but are often absent (Figs 6 – 7). Pseudopores / pores density similar on all dorsal plates, approximate mean density 10 – 13 pseudopores / pores per 100 µm 2 in females and 8 – 10 pseudopores / pores per 100 µm 2 in males (Table 5). Intracuticular pillars visible as fine dark dots under PCM (Figs 10 – 12). Scapular plate typically distinctly faceted with a median longitudinal fold and three smaller transverse folds (Figs 2, 12). In females, sometimes only pseudopore / pore rows mark borders of faint facets (Figs 1, 10 – 11). Paired plates divided into two unequal anterior and posterior parts by a transverse stripe (Figs 10 – 12, 14, 16). Caudal plate faceted with two evident longitudinal folds (Figs 1 – 2, 6, 10 – 12), most often the central portion of this plate is subdivided into two parts by a transverse V-suture (Figs 2, 6, 12). Median plates 1 and 2 with transverse division into two unequal parts (compare Figs 10 – 12, 14, 16). Median plate 3 with faint transverse suture, triangular in shape, and with a roundish posterior edge (Figs 14, 16). Poorly developed supplementary lateral platelets present at the levels of median plates (three pairs of platelets on each body side: a pair between scapular plate and first pair of the segmental plates, a pair between paired plates, and a pair between second pair of segmental plates and caudal plate; Figs 1, 10 – 12), devoid of pores / pseudopores (Fig. 7). Ventral cuticle typically with weakly developed plates. Ventral intracuticular pillars either absent or present and well-visible (compare Figs 4 – 5) at 100 x oil immersion, and always less evident than dorsal (compare Figs 5 and 11). Ventral plate configuration: VII / IX: (2) - (1) - 2 / 4 - 2 - 2 / 4 - 2 - 2 - 2 - 1. Two small subcephalic longitudinal plates just below the mouth opening and one subpharyngeal plate weakly outlined and usually only visible under SEM (compare Figs 5, 8 – 9). Typically, rows III – VIII / IX are discernible (Figs 4 – 5, 8 – 9, 21). Rarely, ventral plates invisible under PCM (e. g. holotype). Ventral granules absent or restricted to the plate surface (Fig. 5). Papilla on legs I absent, minute papilla on legs IV present (Fig. 1). Dentate collar on legs IV with 3 – 6 teeth (Figs 1 – 2, 20, arrowheads). External claws of all legs smooth, internal claws with tiny spurs pointing strongly downwards with very small gap between spur and claw base, making them almost invisible under PCM (however, always clear under SEM; Figs 19 – 20). Remarks on the sexual dimorphism: Sex differences in the new species are well-marked and embrace: longer, tubbier primary and secondary ♂ clavae (compare Figs 1 – 2 and Tables 3 – 4); stronger faceting of the ♂ scapular plate (compare Figs 10 – 11 and 12); significantly lower pseudopore / pore density on unpaired plates in ♂ (compare ranges presented in Table 5). Juveniles (i. e. the second instar, four-clawed without gonopore): In appearance like adults, but clearly smaller (75 µm) and with indiscernible ventral plates (Fig. 22). It is impossible to state firmly whether the species exhibits ontogenetic shifts in ventral armature or the lack of ventral plates in the first two life stages is inconstant. This issue certainly requires more studies, since changes in ventral armature occurring during development were recently detected by Gąsiorek et al. (2017). Larvae (i. e. the first instar, two-clawed without gonopore; measurements in Table 4): Median plates with poorly delineated margins, lacking pseudopores / pores; supplementary lateral platelets and pedal (leg) plates undeveloped. Pores / pseudopores present only on the anterior and posterior margins of the cephalic, scapular, segmental, and caudal plates (Fig. 3). Claws with spurs formed as in adults. Legs IV without dentate collar. Ventral plates absent (Fig. 23). Eggs unknown.	en	Gąsiorek, Piotr, Degma, Peter (2018): Three Echiniscidae species (Tardigrada: Heterotardigrada) new to the Polish fauna, with the description of a new gonochoristic Bryodelphax Thulin, 1928. Zootaxa 4410 (1): 77-96, DOI: 10.11646/zootaxa.4410.1.4
001087910D06FF9AD399FE75FB44F80C.taxon	etymology	Etymology: The name instabilis underlines the variability in the number of ventral plates (see Remarks), which is unusual for Bryodelphax. Moreover, often only some of the ventral plates rows are visible under the light microscope, thus an animal’s venter seems to be devoid of plates in its proximal part.	en	Gąsiorek, Piotr, Degma, Peter (2018): Three Echiniscidae species (Tardigrada: Heterotardigrada) new to the Polish fauna, with the description of a new gonochoristic Bryodelphax Thulin, 1928. Zootaxa 4410 (1): 77-96, DOI: 10.11646/zootaxa.4410.1.4
001087910D06FF9AD399FE75FB44F80C.taxon	discussion	Remarks. Polish and Slovak populations have same ventral plates configuration but in three mature females from the Slovak population, row V has two additional, smaller ventral plates placed in a more marginal position, and two of the specimens also have such additional plates in row III (Figs 5 and 21).	en	Gąsiorek, Piotr, Degma, Peter (2018): Three Echiniscidae species (Tardigrada: Heterotardigrada) new to the Polish fauna, with the description of a new gonochoristic Bryodelphax Thulin, 1928. Zootaxa 4410 (1): 77-96, DOI: 10.11646/zootaxa.4410.1.4
001087910D06FF9AD399FE75FB44F80C.taxon	diagnosis	Differential diagnosis. Bryodelphax instabilis sp. nov. can easily be distinguished from the other members of the weglarskae group on the basis of the scapular plate faceting and being gonochorous. Nevertheless, the new species should be compared with four most similar taxa having last six ventral plates rows configuration same as the new species, i. e. 2 - 2 - 2 - 2 - 2 - 1 or 2 - 4 - 2 - 2 - 2 - 1 (due to the instability in ventral plate arrangement). It differs specifically from:	en	Gąsiorek, Piotr, Degma, Peter (2018): Three Echiniscidae species (Tardigrada: Heterotardigrada) new to the Polish fauna, with the description of a new gonochoristic Bryodelphax Thulin, 1928. Zootaxa 4410 (1): 77-96, DOI: 10.11646/zootaxa.4410.1.4
001087910D06FF9AD399FE75FB44F80C.taxon	description	General remarks on Bryodelphax in Polish Pieniny Mts. and Tatra Mts. There are four sympatric Bryodelphax spp. that inhabit Poland and occur in the Pieniny Mts. (Dastych 1988); three (with the exception of B. weglarskae) are also present in the Tatra Mts. Our new discovery confirms an eucalciphil and subalpine preference for these Central European members of the genus. In Poland, Bryodelphax spp. are frequently found in close proximity. For example, in the Pieniny Mts. B. parvulus was found in the same moss cushions as B. instabilis sp. nov. or B. weglarskae; similarly, B. parvulus and B. tatrensis co-occur in mosses in the Tatra Mts.	en	Gąsiorek, Piotr, Degma, Peter (2018): Three Echiniscidae species (Tardigrada: Heterotardigrada) new to the Polish fauna, with the description of a new gonochoristic Bryodelphax Thulin, 1928. Zootaxa 4410 (1): 77-96, DOI: 10.11646/zootaxa.4410.1.4
001087910D0FFF9ED399F99DFACDFAD6.taxon	description	Locus typicus. Castlebar, Ireland.	en	Gąsiorek, Piotr, Degma, Peter (2018): Three Echiniscidae species (Tardigrada: Heterotardigrada) new to the Polish fauna, with the description of a new gonochoristic Bryodelphax Thulin, 1928. Zootaxa 4410 (1): 77-96, DOI: 10.11646/zootaxa.4410.1.4
001087910D0FFF9ED399F99DFACDFAD6.taxon	materials_examined	Material examined: One four-clawed juvenile, found together with very numerous Echiniscus granulatus (Doyère, 1840), E. reticulatus Murray, 1905, Pseudechiniscus suillus, Testechiniscus spitsbergensis, and Richtersius coronifer (Richters, 1903) (sample PL. 273; Kobylarzowy Coulouir).	en	Gąsiorek, Piotr, Degma, Peter (2018): Three Echiniscidae species (Tardigrada: Heterotardigrada) new to the Polish fauna, with the description of a new gonochoristic Bryodelphax Thulin, 1928. Zootaxa 4410 (1): 77-96, DOI: 10.11646/zootaxa.4410.1.4
001087910D0FFF9ED399F99DFACDFAD6.taxon	description	Description: Very small (162 µm), conspicuously pink coloured. Cirrus internus / externus length ratio 67 %. Cirrus A 48 µm long and very thick, lateral appendages B and E of equal lengths (20 µm), lateral cirri C and D markedly longer (60 and 76 µm, respectively). All dorsal appendages present, of which C d are the longest (44 µm vs B d 18 µm long and D d 33 µm long). In general, cirri in all B-positions are visibly thinner than other appendages. The difference is especially evident in the width of the dorsal appendage bases (1.9 µm in B d, 2.8 µm in C d, and 4.9 µm in D d, respectively). Plates with well-marked borders, lacking cervical (neck) and third median plates. Sculpture composed of polygons (blumi-canadensis type), with the largest polygons found on the small median plates and the central part of the caudal plate. Ventral cuticle smooth, except for the genital zone, where two large, trapezoid plates, in line with legs IV, occur (Fig. 29). Pedal (leg) plates invisible in PCM. Dentate collar on legs IV comprises seven long, parallel teeth. Large, slender claws; claws I – III markedly shorter than claws IV (11.9 – 13.1 µm, 32.5 – 35.8 % vs 16.6 µm, 45.4 %). Small spurs (2.1 µm, 5.7 %) positioned very close to the internal claw bases, and almost perpendicular to the claw branch.	en	Gąsiorek, Piotr, Degma, Peter (2018): Three Echiniscidae species (Tardigrada: Heterotardigrada) new to the Polish fauna, with the description of a new gonochoristic Bryodelphax Thulin, 1928. Zootaxa 4410 (1): 77-96, DOI: 10.11646/zootaxa.4410.1.4
001087910D0FFF9ED399F99DFACDFAD6.taxon	discussion	Remarks: New record for Poland. E. militaris is without doubt one of the most elusive Palaearctic echiniscid species (Ramazzotti & Maucci 1983). Since its original description in 1911, it has been recorded only a few times and in very small numbers from various European countries (McInnes 1994). Pilato (1977) suppressed all subspecies after analysing different populations of the species. As there is a very high variability among recorded specimens, it is very likely that a number of independent species are captured under the one specific name. Unfortunately, difficulties in obtaining an adequate number of specimens for a detailed taxonomic study, including DNA sequencing, will hinder clarification of the taxonomic status of this species and resolving whether this is a single species exhibiting considerable intraspecific variability or a group of morphologically similar species.	en	Gąsiorek, Piotr, Degma, Peter (2018): Three Echiniscidae species (Tardigrada: Heterotardigrada) new to the Polish fauna, with the description of a new gonochoristic Bryodelphax Thulin, 1928. Zootaxa 4410 (1): 77-96, DOI: 10.11646/zootaxa.4410.1.4
001087910D08FF9CD399FA48FD38FBD4.taxon	description	Locus typicus. Visby, Gotland, Sweden.	en	Gąsiorek, Piotr, Degma, Peter (2018): Three Echiniscidae species (Tardigrada: Heterotardigrada) new to the Polish fauna, with the description of a new gonochoristic Bryodelphax Thulin, 1928. Zootaxa 4410 (1): 77-96, DOI: 10.11646/zootaxa.4410.1.4
001087910D08FF9CD399FA48FD38FBD4.taxon	materials_examined	Material examined. Three adult specimens and one juvenile, found together with exceptionally abundant Echiniscus granulatus, E. testudo (Doyère, 1840), and Richtersius coronifer (PL. 247; Kościeliska Valley.	en	Gąsiorek, Piotr, Degma, Peter (2018): Three Echiniscidae species (Tardigrada: Heterotardigrada) new to the Polish fauna, with the description of a new gonochoristic Bryodelphax Thulin, 1928. Zootaxa 4410 (1): 77-96, DOI: 10.11646/zootaxa.4410.1.4
001087910D08FF9CD399FA48FD38FBD4.taxon	description	Description. Medium-sized (160 – 313 µm), dark-yellow. Cirrus A long (104 µm, cirrus A / body length ratio 33 %). Cirrus internus / externus length ratio 60 – 65 %. Plate sculpture composed of small and medium-sized pores (spinulosus type), with the largest pores found on the large median plates and on the caudal plate. Lateral and dorsal appendages at B, C, C d, D, D d, and E in the shape of long spines, with dorsal spines slightly longer than the lateral ones (38 – 48 µm vs 21 – 36 µm). Pedal (leg) plates very well-developed and large, those on leg IV forming a collar composed of eight small, blunt teeth. Claws I – III moderately shorter than claws IV (12.5 – 20.6 µm, 27.3 – 31.1 % vs 16.9 – 23.6 µm, 30.3 – 38.3 %). Spurs (1.5 – 3.9 µm, 3.4 – 5.8 %) positioned at ~ 30 % of the claw length.	en	Gąsiorek, Piotr, Degma, Peter (2018): Three Echiniscidae species (Tardigrada: Heterotardigrada) new to the Polish fauna, with the description of a new gonochoristic Bryodelphax Thulin, 1928. Zootaxa 4410 (1): 77-96, DOI: 10.11646/zootaxa.4410.1.4
001087910D08FF9CD399FA48FD38FBD4.taxon	discussion	Remarks. New record for Poland. Dastych (1988) expressed the opinion that the so-called spinulosus - quadrispinosus group was in need of a complete revision, and produced a schematic illustration of E. spinulosus (Doyère, 1840) with long lateral appendages. He also remarked that the lateral appendages of Polish E. spinulosus specimens varied significantly in length (Dastych 1988). Presented findings suggest that his records could embrace two distinct species: E. spinulosus s. s., which is a predominantly lowland species, and E. spiniger, more frequently found in colder and upland habitats. In E. spinulosus specimens from the locus typicus (Saint-Maur-des-Fossés, France), dorsal spines C d and D d are longer than the extremely short lateral spines (similar to the differences in length recorded for E. scabrospinosus Fontoura, 1982, see Figs 31 – 32). In E. spiniger, the lateral spines are longer and of similar or equal development to the dorsal appendages (Fig. 34). Scottish specimens of E. quadrispinosus Richters, 1902 of identical body size to Polish E. spiniger specimens have dorsal spines C d and D d distinctly shorter than in E. spiniger, and unlike in E. spiniger, spines D d are typically shorter than C d (Fig. 35). Moreover, all lateral appendages in E. quadrispinosus are developed as long cirri with broad bases (Ramazzotti & Maucci 1983). Pilato et al. (2008) showed that different species within spinulosus group can be separated on the basis of differences in plate sculpturing. An interesting pattern can be observed within members of the group: generally, species with short appendages exhibit large and sparsely distributed cuticular pores, whereas those with long spines or cirri possess comparatively smaller, more densely arranged pores (compare Figs 31 – 32 for E. spinulosus and E. scabrospinosus, and 33 – 35 for E. lichenorum Maucci, 1983, E. spiniger and E. quadrispinosus). Curiously, E. migiurtinus Franceschi, 1957 with reduced set of trunk appendages (exclusively stout spines D d present) seems to deviate from this rule, since it exhibits small, densely arranged pores (Fig. 30). However, there are also differences in the degree of pedal (leg) plate development between the representatives of the spinulosus group. Such differences could be a stable specific trait. Pedal (leg) plates can be: invisible in PCM on all legs (Fig. 32); visible only on legs IV (Figs 30, 33, 35, arrowheads); or present on all legs, and with strongly developed sculpture similar to that on the dorsal plates, which so far was detected only in E. spinulosus and E. spiniger (Figs 31, 34, arrowheads). It is clear that additional studies are needed to better understand the phylogeny and evolution of morphological traits within this species complex. For the Polish fauna, both E. spiniger and E. spinulosus should be considered as its indigenous components. The presence of E. testudo, a typical lowland species, within this sample, represents the first record for this species from the Polish Tatras (Dastych 1980).	en	Gąsiorek, Piotr, Degma, Peter (2018): Three Echiniscidae species (Tardigrada: Heterotardigrada) new to the Polish fauna, with the description of a new gonochoristic Bryodelphax Thulin, 1928. Zootaxa 4410 (1): 77-96, DOI: 10.11646/zootaxa.4410.1.4
001087910D0AFF9DD399FB00FB23FF3F.taxon	description	Terra typica. Greenland.	en	Gąsiorek, Piotr, Degma, Peter (2018): Three Echiniscidae species (Tardigrada: Heterotardigrada) new to the Polish fauna, with the description of a new gonochoristic Bryodelphax Thulin, 1928. Zootaxa 4410 (1): 77-96, DOI: 10.11646/zootaxa.4410.1.4
001087910D0AFF9DD399FB00FB23FF3F.taxon	materials_examined	Material examined. Eleven females, two males, and one juvenile, found together with Echiniscus granulatus, Testechiniscus spitsbergensis, and Hypsibius convergens (Urbanowicz, 1925) (PL. 247, 248, 252, and 274; Kościeliska Valley, Ciemniak hillside, Kobylarzowy Coulouir).	en	Gąsiorek, Piotr, Degma, Peter (2018): Three Echiniscidae species (Tardigrada: Heterotardigrada) new to the Polish fauna, with the description of a new gonochoristic Bryodelphax Thulin, 1928. Zootaxa 4410 (1): 77-96, DOI: 10.11646/zootaxa.4410.1.4
001087910D0AFF9DD399FB00FB23FF3F.taxon	description	Description. Small (females 152 – 201 µm, males 180 – 205 µm), light-orange. Black eyes present. Primary clavae long, secondary clavae large (females: 20.6 – 22.7 %, male: 35.1 %; Figs 36 – 37, arrowheads). Cirrus A very short (24.6 – 32.7 µm, cirrus A / body length ratio 14 – 20 %). All plates weakly sclerotized, covered with fine granulation of suillus type. Cephalic and caudal plates clearly facetted (Figs 36 – 37, empty arrowheads). Pseudosegmental plate always divided longitudinally. Pedal (leg) plates present. Spine on the leg I absent. Venter with developed sculpture consisting of intracuticular pillars. Claws I – IV 7.7 – 11.0 µm long (34.0 – 46.2 %), the internal claws with spurs, 1.2 – 2.1 µm long (4.7 – 9.3 %).	en	Gąsiorek, Piotr, Degma, Peter (2018): Three Echiniscidae species (Tardigrada: Heterotardigrada) new to the Polish fauna, with the description of a new gonochoristic Bryodelphax Thulin, 1928. Zootaxa 4410 (1): 77-96, DOI: 10.11646/zootaxa.4410.1.4
001087910D0AFF9DD399FB00FB23FF3F.taxon	discussion	Remarks. Additional Polish record. Dastych (1970) was the first to report this taxon from Poland (as P. suillus facettalis), but in his later monograph he synonymised it with P. suillus (Dastych 1988). Since one of the primary characters separating these two species is the presence of cephalic and caudal plate faceting, the status of P. facettalis as an element of the Polish fauna is restored. Kristensen (1987) gave two features that allow differentiation of the sexes in the genus Pseudechiniscus: body shape (males are more elongate than females) and enlarged male secondary clavae. There are two additional qualitative traits which can be used to distinguish the sexes in P. facettalis, i. e. males are more flattened than females (compare Figs 36 and 37), and the faceting of the caudal plate in males is not as deep as that observed in females (visible only when specimens are laterally or dorso-laterally oriented). Similar sexual dimorphism was observed in the two populations from Norway and Tunisia used for comparisons (see Table 2).	en	Gąsiorek, Piotr, Degma, Peter (2018): Three Echiniscidae species (Tardigrada: Heterotardigrada) new to the Polish fauna, with the description of a new gonochoristic Bryodelphax Thulin, 1928. Zootaxa 4410 (1): 77-96, DOI: 10.11646/zootaxa.4410.1.4
