taxonID	type	description	language	source
9323FE1EFF9AD312FF464271FC6EFDBC.taxon	description	The Echiniscus brunus complex	en	Dey, Pritam K., Gąsiorek, Piotr, Michalczyk, Łukasz (2024): Convergent evolution of dark, ultraviolet-absorbing cuticular pigmentation in a new Afro-Oriental Echiniscus brunus species complex (Heterotardigrada: Echiniscidae). Zoological Journal of the Linnean Society 200 (1): 34-59, DOI: 10.1093/zoolinnean/zlad132, URL: http://dx.doi.org/10.1093/zoolinnean/zlad132
9323FE1EFF9AD312FF464271FC6EFDBC.taxon	diagnosis	Diagnosis: Unappendaged Echiniscus spp. of caramel colour in live individuals (e. g. Figs 3, 5; compare also with Figs 4, 6) caused by an interplay of orange pigments in the body cavity and dark brown pigmentation of the dorsal and pedal plates, and pulvini on legs. In specimens preserved in Hoyer’s medium, i. e. in which orange pigments dissolve asser several days, specimens are brown owing to the cuticular plate pigmentation (Figs 5, 9, 10, 12, 14). The dark cuticular pigmentation is UV absorbent (Fig. 7). All dorsal and (well-developed) pedal plates are with pores and without intracuticular pillars (sculpture of the E. spinulosus type; Figs 5, 9, 10, 12, 14). A median anterior suture has approximately from one-third to one-half of the scapular plate; it is visible in PCM and BFM as a bright incision and as a stripe of mostly smooth cuticle in SEM (Figs 3 E, 5, 7 B, 9 A – C, 10 A, E, F, 11 A, 12 A, 13 A, 14 A). In some specimens, a crescent-shaped thickening on the caudal plate is present, beưer visible in PCM (Figs 5, 10 A, E, 12 A, 14 A), but also detectable in SEM (Figs 11 A, B, 13 A, B) and UVM (Fig. 7 B). A pair of subcephalic ventral plates is present (Figs 10 C, 12 C, 14 C). Differential diagnosis: Because of the combination of distinct coloration (caramel / brown) and the lack of trunk appendages, species of the new complex are easily differentiable from all other Echiniscus spp. However, because of the presence of the dark cuticular pigmentation and the lack of trunk appendages, species of the E. brunus complex may be mistaken for Viridiscus at first sight. Thus, for accurate phenotypic identification, a careful examination of cuticular pigmentation (brown in the new complex vs. dark green to almost black in Viridiscus), dorsal plate endocuticle (homogeneous and solid in the new complex vs. sponge layer in Viridiscus) and scapular plate morphology (with a conspicuous anterior suture the new complex vs. no suture observed in Viridiscus) is needed. Composition: Echiniscus brunus (India) and two undescribed candidate species, E. aff. brunus sp. can. 1 and (India) E. aff. brunus sp. can. 2 (Tanzania), genetically distinct but without a sufficient number of specimens to warrant a proper characterization of their morphological intraspecific variability and formal species descriptions.	en	Dey, Pritam K., Gąsiorek, Piotr, Michalczyk, Łukasz (2024): Convergent evolution of dark, ultraviolet-absorbing cuticular pigmentation in a new Afro-Oriental Echiniscus brunus species complex (Heterotardigrada: Echiniscidae). Zoological Journal of the Linnean Society 200 (1): 34-59, DOI: 10.1093/zoolinnean/zlad132, URL: http://dx.doi.org/10.1093/zoolinnean/zlad132
9323FE1EFF9AD318FC46442FFA86F8FA.taxon	description	(Figs 3, 5, 7 B, 8 B, 9 A, 10, 11; Tables 2 – 4; Supporting Information, SM. 04) Type locality and type series: 11 ° 24 ʹ 55 ″ N, 76 ° 47 ʹ 35 ″ E, 2110 m a. s. l.; India, Tamil Nadu, Nilgiris, Kothagiri. Holotype (adult female) on slide IN. 391.02; 26 paratypes (13 adult females, nine juveniles, one exuvia and three larvae) on slides IN. 391.01 – IN. 391.13; 20 paratypes on SEM stub no. 22.05; three hologenophores preserved on slides IN. 391.01 – IN. 391.3, all deposited in the Department of Invertebrate Evolution of the Jagiellonian University.	en	Dey, Pritam K., Gąsiorek, Piotr, Michalczyk, Łukasz (2024): Convergent evolution of dark, ultraviolet-absorbing cuticular pigmentation in a new Afro-Oriental Echiniscus brunus species complex (Heterotardigrada: Echiniscidae). Zoological Journal of the Linnean Society 200 (1): 34-59, DOI: 10.1093/zoolinnean/zlad132, URL: http://dx.doi.org/10.1093/zoolinnean/zlad132
9323FE1EFF9AD318FC46442FFA86F8FA.taxon	etymology	Etymology: An epithet, in Latin brunus = brown, referring to the caramel / brown colour of the new species, the first formally described representative of the complex of brown echiniscids. An adjective in the nominative singular.	en	Dey, Pritam K., Gąsiorek, Piotr, Michalczyk, Łukasz (2024): Convergent evolution of dark, ultraviolet-absorbing cuticular pigmentation in a new Afro-Oriental Echiniscus brunus species complex (Heterotardigrada: Echiniscidae). Zoological Journal of the Linnean Society 200 (1): 34-59, DOI: 10.1093/zoolinnean/zlad132, URL: http://dx.doi.org/10.1093/zoolinnean/zlad132
9323FE1EFF9AD318FC46442FFA86F8FA.taxon	description	Adult females (i. e. from the third instar onwards; measurements and statistics are in Table 2). Body medium in size and plump (Figs 3, 5, 10 A, F, 11 A, B); dark orange pigments dominant in the ventral portion and limbs, but brown pigments prevalent within dorsal plates in living animals and specimens freshly mounted in Hoyer’s medium (Fig. 5 A, D). Immediately asser the mount, specimen appears orange to crimson red with caramel – brown dorsal plates under PCM, while under BFM the body appears slightly greenish in colour, with orange pigment inside the body cavity (Fig. 5 A, D). Several hours asser mounting, the body cavity becomes transparent, with a few droplets of yellowish orange pigments remaining inside the body cavity, but the dorsal and pedal plates appear more brown than caramel under PCM, while body cavity appears transparent with brown dorsal and pedal plates (Fig. 5 B, E). Asser several days when all the yellow and orange pigmentation dissolve, the body cavity appears transparent, with brown dorsal plates and pedal plates (Figs 5 C, F, 10). Large red granular eye spots are present in all individuals and vanish asser mounting. Buccal apparatus is short, rigid, with large cochlear stylet furcae and no stylet supports. Peribuccal cirri with cirrophores, cephalic papilla (secondary clava) elongated and conical (Figs 5 D – F, 10, 11). Primary clava small and tightly adjacent to the cirrophore of cirrus A (Figs 5 D – F, 9 A, 10 A, F, 11 A, B), which is extremely short, in some specimens even shorter than cirrus externus (Fig. 10 F). Sculpturing of body plates comprises two types of pores: (i) more sparsely arranged and circular in shape with thick borders, dominant in most plate portions (Figs 10, 11); and (ii) tightly arranged and polygonal in shape, usually with thin borders, present only in anterior portions of paired segmental plates, lateralmost portions of the scapular, paired segmental, intersegmental and caudal (terminal) plates and pedal platelets (Figs 10, 11). Pores without dark intraporal rings. Cephalic plate small, with a clearly marked anterior chalice-like incision (Figs 5, 7 B, 10 A, E, F, 11 A); cervical plate usually not identifiable, only sometimes a weak rectangular belt visible in LM (Figs 5, 7 B, 10 A, F). Scapular plate large, with the central poreless suture that can reach as far as to 80 % of the plate (Figs 3 E, 5, 7 B, 9 A, 10 A, F, 11 A); lateralmost plate portions demarcated by straight or slightly V-shaped sutures / incisions at the level of cirrus A (Figs 5, 7 B, 10 A, F). Always only two unipartite, triangular median plates, m 1 and m 2, with m 2 being slightly larger. Posterior margins of median plates ossen with more sparsely distributed pores (Fig. 11 A, B). Paired segmental plates I and II with posterior portions three to four times broader than anterior ones; a poreless belt separates anterior and posterior plate portions (Figs 5, 10 A, B, F, 11 B, C). Two pairs of large triangular intersegmental plates: the first positioned between the scapular plate and paired segmental plates I, and the second between plates I and II; a single pair between plate II and the caudal plate (Figs 5, 10 A, B, F, 11 B, C). The caudal plate is the largest element of the armour. In most specimens, a crescent-shaped thickening on the caudal plate is present, usually with fewer pores present in its central part compared with the remainder of the caudal plate (Figs 5, 10 A, F, 11 A, B). Venter densely granulated with minute endocuticular pillars, as typical for Echiniscus; a pair of subcephalic plates with large pores present (Fig. 13 D). Pulvini thin, but identifiable; pedal plates very large and intensely sculptured (Figs 5, 10 A, D – F, 11 B, D). First leg pair with a minute spine embedded on a pedal platelet (Figs 5, 10 A, E, 11 B, D). Fourth leg pair with a small papilla and a fringe composed of many tiny, separate teeth (Figs 5, 10 A, F, 11 B). Claws large and massive; internal ones with typical spurs divergent from main branches and positioned at ~ 20 % – 25 % of the branch. Juveniles (i. e. the second instar; measurements and statistics are in Table 3). ºualitatively similar to sexually mature females. Gonopore lacking. Larvae (i. e. the first instar; measurements and statistics are in Table 4). ºualitatively similar to later life stages, already with two pronounced types of pores (Fig. 10 E). Two claws per leg, each with divergent and evident spurs (Fig. 10 E). Gonopore and anus lacking. DNA sequences: A set of five markers was obtained: 18 S rRNA (OR 520116 – 7), 28 S rRNA (OR 520110 – 1), ITS- 1 (OR 520101 – 3), ITS- 2 (OR 520122 – 4) and COI (OR 506701 – 3). Uncorrected pairwise distances between the new species and the two candidate species within the complex were as follows: 18 S rRNA, 0.2 % (both E. aff. brunus sp. can. 1 and E. aff. brunus sp. can. 2); 28 S rRNA, 0.1 % – 0.3 % (E. aff. brunus sp. can. 2 and E. aff. brunus sp. can. 1, respectively); ITS- 1, 3.1 % – 4.4 % (E. aff. brunus sp. can. 1 and E. aff. brunus sp. can. 2, respectively); ITS- 2, 3.4 % – 4.9 % (E. aff. brunus sp. can. 2 and E. aff. brunus sp. can. 1, respectively); and COI, 2.0 % – 3.7 % (E. aff. brunus sp. can. 2 and E. aff. brunus sp. can. 1, respectively). See Supporting Information, SM. 03 for matrices with genetic distances.	en	Dey, Pritam K., Gąsiorek, Piotr, Michalczyk, Łukasz (2024): Convergent evolution of dark, ultraviolet-absorbing cuticular pigmentation in a new Afro-Oriental Echiniscus brunus species complex (Heterotardigrada: Echiniscidae). Zoological Journal of the Linnean Society 200 (1): 34-59, DOI: 10.1093/zoolinnean/zlad132, URL: http://dx.doi.org/10.1093/zoolinnean/zlad132
9323FE1EFF9AD318FC46442FFA86F8FA.taxon	discussion	Remarks: Found with Milnesium sp., Ramazzottius sp. and Macrobiotus sp. in the sample. In several specimens, developmental malformations were observed, in which various dorsal plates merged into a single large plate (for an example, see Fig. 10 F).	en	Dey, Pritam K., Gąsiorek, Piotr, Michalczyk, Łukasz (2024): Convergent evolution of dark, ultraviolet-absorbing cuticular pigmentation in a new Afro-Oriental Echiniscus brunus species complex (Heterotardigrada: Echiniscidae). Zoological Journal of the Linnean Society 200 (1): 34-59, DOI: 10.1093/zoolinnean/zlad132, URL: http://dx.doi.org/10.1093/zoolinnean/zlad132
9323FE1EFF9AD318FC46442FFA86F8FA.taxon	diagnosis	Differential diagnosis: Because of the distinct brown, UV-absorbing cuticular pigmentation (also clearly visible in specimens mounted in Hoyer’s medium) and the presence of an anterior suture in the scapular plate, the new species cannot be mistaken for any other formally described echiniscid.	en	Dey, Pritam K., Gąsiorek, Piotr, Michalczyk, Łukasz (2024): Convergent evolution of dark, ultraviolet-absorbing cuticular pigmentation in a new Afro-Oriental Echiniscus brunus species complex (Heterotardigrada: Echiniscidae). Zoological Journal of the Linnean Society 200 (1): 34-59, DOI: 10.1093/zoolinnean/zlad132, URL: http://dx.doi.org/10.1093/zoolinnean/zlad132
9323FE1EFF90D305FC75411BFE77F933.taxon	description	(Figs 12, 13) Material examined: Eight individuals (for details, see Table 1). Shortened description: Morphotype almost identical to that of E. brunus, except for the presence of the same type of circular cuticular pores with thick borders on all parts of the dorsal and pedal plates in E. aff. brunus sp. can. 1 (Figs 12 B, D and 13 C – D) vs. pores on dorsal and pedal plates different in E. brunus sp. nov. (Figs 10 B, D and 11 C, D). Molecular phylogenetic analysis suggests it is a closely related sister, potentially cryptic or pseudocryptic, species to E. brunus (see Fig. 2 and genetic distances in the description of E. brunus above). DNA sequences: A set of five markers was obtained: 18 S rRNA (OR 520118 – 9), 28 S rRNA (OR 520112 – 3), ITS- 1 (OR 520104 – 7), ITS- 2 (OR 520125 – 8) and COI (OR 506704 – 7). Uncorrected pairwise distances to E. brunus can be found in the description of the new species above, whereas genetic p-distances to E. aff. brunus sp. can. 2 are as follows: 18 S rRNA, 0.0 %; 28 S rRNA, 0.4 %; ITS- 1, 5.1 %; ITS- 2, 3.7 %; and COI, 3.6 % (see also Supporting Information, SM. 03).	en	Dey, Pritam K., Gąsiorek, Piotr, Michalczyk, Łukasz (2024): Convergent evolution of dark, ultraviolet-absorbing cuticular pigmentation in a new Afro-Oriental Echiniscus brunus species complex (Heterotardigrada: Echiniscidae). Zoological Journal of the Linnean Society 200 (1): 34-59, DOI: 10.1093/zoolinnean/zlad132, URL: http://dx.doi.org/10.1093/zoolinnean/zlad132
9323FE1EFF90D305FC75411BFE77F933.taxon	discussion	Remarks: Foundwith Echiniscus cf. scabrospinosus, Pseudechiniscus sp., Milnesium sp., Dianea sp., Ramazzottius sp., Diphascon sp., Minibiotus sp. and Macrobiotus sp. in the sample. Compare Figures 10 D, 11 D with Figures 12 D, 13 D for a potential difference in the shape of pedal plate pores between E. brunus and Echiniscus aff. brunus sp. can. 1.	en	Dey, Pritam K., Gąsiorek, Piotr, Michalczyk, Łukasz (2024): Convergent evolution of dark, ultraviolet-absorbing cuticular pigmentation in a new Afro-Oriental Echiniscus brunus species complex (Heterotardigrada: Echiniscidae). Zoological Journal of the Linnean Society 200 (1): 34-59, DOI: 10.1093/zoolinnean/zlad132, URL: http://dx.doi.org/10.1093/zoolinnean/zlad132
9323FE1EFF8DD306FE8F40D2FF09FA54.taxon	description	(Fig. 14)	en	Dey, Pritam K., Gąsiorek, Piotr, Michalczyk, Łukasz (2024): Convergent evolution of dark, ultraviolet-absorbing cuticular pigmentation in a new Afro-Oriental Echiniscus brunus species complex (Heterotardigrada: Echiniscidae). Zoological Journal of the Linnean Society 200 (1): 34-59, DOI: 10.1093/zoolinnean/zlad132, URL: http://dx.doi.org/10.1093/zoolinnean/zlad132
9323FE1EFF8DD306FE8F40D2FF09FA54.taxon	materials_examined	Material examined: Four individuals (for details, see Table 1).	en	Dey, Pritam K., Gąsiorek, Piotr, Michalczyk, Łukasz (2024): Convergent evolution of dark, ultraviolet-absorbing cuticular pigmentation in a new Afro-Oriental Echiniscus brunus species complex (Heterotardigrada: Echiniscidae). Zoological Journal of the Linnean Society 200 (1): 34-59, DOI: 10.1093/zoolinnean/zlad132, URL: http://dx.doi.org/10.1093/zoolinnean/zlad132
9323FE1EFF8DD306FE8F40D2FF09FA54.taxon	description	Shortened description: Except for pore density in the dorsal plates (Fig. 14 A), morphotype is identical to that of E. brunus and E. aff. brunus sp. can. 1 (Fig. 14 B – D). In contrast to the two Indian species, cuticular pores in E. aff. brunus sp. can. 2 are more numerous and thus more densely arranged (see Fig. 14 A). Despite clear genetic differences between E. aff. brunus sp. can. 2 and the two remaining species in the complex, we refrain from describing it as a new species because of the low number of available specimens, which precludes a detailed description of intraspecific morphological variability. DNA sequences: A set of five markers was obtained: 18 S rRNA (OR 520120 – 1), 28 S rRNA (OR 520114 – 5), ITS- 1 (OR 520108 – 9), ITS- 2 (OR 520129 – 30) and COI (OR 506708 – 9). For p-distances, see the description of E. brunus and the note about E. aff. brunus sp. can. 1 above (see also Supporting Information, SM. 03). Remarks: Found together with E. belloporus, E. tantulus, E. tristis and several species of macrobiotids and milnesiids in the samples.	en	Dey, Pritam K., Gąsiorek, Piotr, Michalczyk, Łukasz (2024): Convergent evolution of dark, ultraviolet-absorbing cuticular pigmentation in a new Afro-Oriental Echiniscus brunus species complex (Heterotardigrada: Echiniscidae). Zoological Journal of the Linnean Society 200 (1): 34-59, DOI: 10.1093/zoolinnean/zlad132, URL: http://dx.doi.org/10.1093/zoolinnean/zlad132
