identifier	taxonID	type	CVterm	format	language	title	description	additionalInformationURL	UsageTerms	rights	Owner	contributor	creator	bibliographicCitation
2A7987F0FFC20720FCB2FB62F6EDF924.text	2A7987F0FFC20720FCB2FB62F6EDF924.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Echiniscidae Thulin 1928	<div><p>Family: Echiniscidae Thulin, 1928</p><p>Genus: Echiniscus C.A.S. Schultze, 1840 (amended by Gąsiorek et al., 2017)</p><p>Species: Echiniscus africanus Murray, 1907</p><p>Table 3, Figs. 3 and 4</p><p>The specimens from Celebes and Madagascar fully correspond phenotypically with a single individual originating from terra typica in South Africa and examined by us (Gąsiorek et al., 2022), with a sole exception of the number of spines on the posterior margin of the scapular plate. Originally reported as having six spines (Murray, 1907), this character was later disclosed as variable (Gąsiorek &amp; Vončina, 2019; Gąsiorek et al., 2022), and the Malayan specimens conform with this, having either two (Figs. 3A and B, 4) or even no spines (Fig. 3C). Other trunk appendages may also be absent (Fig. 3C). The observations on the variability of chaetotaxy in E. africanus fully agree with initial data for a likely population of this species from Hawaii (Tsaliki et al., 2018). The sister relationship between E. africanus and E. lapponicus (Fig. 2) confirms</p><p>◂ Fig. 2 Updated phylogeny of the Echiniscus -like genera in the Bayesian and Maximum Likelihood tree based on nuclear markers (almost identical topologies, see the text for details). Species currently classified within Kristenseniscus given in colour, and newly added Echiniscus spp. —in bold. The scale refers to the Bayesian tree and represents substitutions per position; posterior probability values are provided as first at the nodes, followed by bootstrap values after slash; The asterisk (*) indicates maximal support, pound sign (#) indicates no support. Diploechiniscus oihonnae and Testechiniscus spitsbergensis constitute an outgroup a previously hypothesised relatedness based on the presence of centrodorsal spines, often crossed in a scissor-like manner (Fig. 4). This character can be now regarded as a synapomorphy for this clade.</p><p>Gąsiorek and Vončina (2019) have already stressed there are no morphological criteria for distinguishing between E. africanus and E. semifoveolatus . The latter species was reported from Japan and China (Qiao et al., 2013; Suzuki, micrometer) of selected morphological structures of adult females of Echiniscus africanus mounted in Hoyer’s medium 2017). Considering the fact that E. africanus is potentially a pantropical species (Africa, Hawaii, Madagascar, the Malay Archipelago), it is imperative to test the conspecificity of these two species, with a potential junior synonymy pertaining to E. semifoveolatus that we designate as nomen inquirendum herein.</p><p>sp the proportion between the length of a given structure and the length of the scapular plate,?—unknown</p><p>Species: Echiniscus minutus sp. nov.</p><p>Table 4, Figs. 5 and 6</p><p>Locus typicus and type material: 0°57′58″S, 119°46′25″E, 1309 m asl; Indonesia, Celebes, Sulawesi Tengah, Marowola Mts.; moss from tree bark. Holotype (mature female on the slide ID.374.17), seven paratypic females and one exuvia (slides ID.374.15–16, ID.485.02, ID.486.01, ID.517.01, 26, ID.564.01). All are deposited in the <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=119.77361&amp;materialsCitation.latitude=-0.9661111" title="Search Plazi for locations around (long 119.77361/lat -0.9661111)">Institute of Zoology</a> and <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=119.77361&amp;materialsCitation.latitude=-0.9661111" title="Search Plazi for locations around (long 119.77361/lat -0.9661111)">Biomedical Research</a> of the <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=119.77361&amp;materialsCitation.latitude=-0.9661111" title="Search Plazi for locations around (long 119.77361/lat -0.9661111)">Jagiellonian University in Kraków</a>, Poland.</p><p>Etymology: From Latin minutus = small, little. The name underlines the minute body size of adults of the new species, being one of the smallest representatives of Echiniscus . Adjective in the nominative singular.</p><p>Mature females (i.e. from the third instar onwards; measurements in Table 4). Body plump (Figs. 5, 6) and orange. Minute red eyes present. Primary and secondary clavae typical, of the Echiniscus - type; peribuccal cirri with well-developed cirrophores. Cirrus A very short (&lt;&lt;25% of the body length), with cirrophore. Body appendage configuration A -(B)- C -C d -D -D d -E, with all trunk appendages formed as robust spines with serrated edges (Figs. 5, 6). Spines B reduced, if present (Fig. 6). Spines C with broad bases, more robust than the remaining appendages (Fig. 5).</p><p>Dorsal plates strongly sclerotised and well-demarcated from each other, with the modified spinulosus type of sculpturing, consisting of typical pores located between aberrantly expanded epicuticular layer (Figs. 5, 6). Epicuticular layer forms striae in the median plates and the anterior portions of paired segmental plates and more or less separated granules in the scapular plate, lateral portions of paired segmental plates and the anterior margin of the caudal (terminal) plate (Figs. 5, 6). Both structures are atypical for the spinulosus morphogroup. The sculpturing vanishes in the centroposterior portions of paired segmental plates and in the lateralmost portions of the caudal plate. Pores are always without dark endocuticular rings. The cephalic plate consists of two weakly delineated halves, with a small, anterior chalice-like incision. The cervical (neck) plate is in the form of a narrow, grey belt, adjacent to the scapular plate; it can be unsculptured (Fig. 5) or sculptured, with minute pores (Fig. 6). The scapular plate is non-facetted, with a central weak sulcus/ incision (Fig. 5) and lateral sutures at the level of primary clavae demarcating unsculptured, rectangular portions (Figs. 5, 6). Three median plates: m1 and m3 are unipartite, whereas m2 is bipartite, with a wide transverse belt (Fig. 5). Two pairs of large segmental plates, their narrower anterior portions are separated from posterior portions by wide, non-sculptured transverse belts. The caudal (terminal) plate with short unsclerotised incisions, non-facetted (Figs. 5, 6).</p><p>Ventral cuticle smooth beside the gonoporal area, where a pair of grey, unsculptured genital plates flanks a hexapartite gonopore anteriorly to a trilobed anus, placed between legs IV. Pedal plates are present on all legs, small with minute pores on legs I–III, and unsculptured on legs IV (Fig. 6). Weak pulvini present on all legs (Fig. 5A). Spine I rudimental and exhibiting various stages of reduction: from a punctiform mass (Fig. 5A) to minute cone (Fig. 6). Dentate collar IV is composed of numerous short teeth. Papilla on leg IV is present (Figs. 5, 6). Claws short; claws IV higher than claws I–III. External claws on all legs are smooth. Internal claws with needle-like spurs positioned at ca. 25% of the claw height, tightly adjacent to the branches and homomorphic on all claws (Fig. 6, insert).</p><p>sp the proportion between the length of a given structure and the length of the scapular plate,?—unknown</p><p>Juveniles, larvae and eggs. Unknown.</p><p>Phylogenetic position: Echiniscus minutus sp. nov. is a sister species to the clade of the mostly tropical/Southern Hemisphere taxa (Fig. 2).</p><p>Phenotypic differential diagnosis: Small body size of adult females, a complex subtype of sculpturing (intermingled pores and granules) and the reduction of spine I make a combination of traits allowing for an easy identification of E. minutus sp. nov. There are several species that exhibit a similar morphotype of E. minutus sp. nov. but can be distinguished from it as follows:</p><p>• Echiniscus marcusi, a representative of Australian fauna (Pilato et al., 1989), being the only other member of the spinulosus morphogroup with well-developed striae in some elements of the dorsal armour, by the presence of epicuticular granules (present in E. minutus sp. nov. vs absent in E. marcusi), and by the presence of pores on legs (present in E. minutus sp. nov. vs absent in E. marcusi).</p><p>• Several species having typically strongly serrated trunk spines, such as E. duboisi or E. manuelae (Richters, 1902; da Cunha &amp; do Nascimento Ribeiro, 1962; Claxton, 1996; Gąsiorek &amp; Kristensen, 2018), exhibit a uniform, classical spinulosus sculpturing without the epicuticular matrix in the form of striae and granules.</p></div>	https://treatment.plazi.org/id/2A7987F0FFC20720FCB2FB62F6EDF924	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Gąsiorek, Piotr;Michalczyk, Łukasz	Gąsiorek, Piotr, Michalczyk, Łukasz (2025): Novel integrative data for Indomalayan echiniscids (Heterotardigrada): new species and old problems. Organisms Diversity & Evolution 25 (1): 81-116, DOI: 10.1007/s13127-023-00628-5, URL: https://doi.org/10.1007/s13127-023-00628-5
2A7987F0FFC8073AFCB2F97BF20DF928.text	2A7987F0FFC8073AFCB2F97BF20DF928.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Echiniscus pusae Marcus 1928	<div><p>Species: Echiniscus pusae Marcus, 1928</p><p>Table 5, Figs. 7 and 8</p><p>The dorsal plate sculpturing of E. pusae and E. africanus is almost identical; the main difference lies in the development of polygonal epicuticular edges of pores in paired segmental plates I–II: in E. africanus, these pores are poorly developed and only in the centroposterior plate portions are they comparable with those in the scapular and caudal plates (Fig. 4), whereas in E. pusae, pores are developed in the entire posterior plate portions and well-developed, similarly to the scapular and caudal plates (Figs. 7, 8). However, despite some morphological similarities, E. pusae is not directly related to the africanus complex (Gąsiorek et al., 2022), but was uncovered as sister to the virginicusperarmatus complex (Fig. 2). An evident epicuticular layer of ornamentation that overlaps with endocuticular layer of pillars and stiff spines in all trunk positions lend morphological support for such a grouping. One of the two main Echiniscus clades that comprises the africanus, blumicanadensis, granulatus-quadrispinosus, and virginicusperarmatus complexes can be characterised by a dominant dorsal plate sculpturing with large endocuticular pillars of polygonal shape and/or large epicuticular pores with polygonal edges. Among the taxa included in the analysis, only E. quadrispinosus Richters, 1902 diverges from this pattern, rendering its sculpturing as autapomorphic.</p><p>With reports from South and Central Africa (Bochnak et al., 2020; Gąsiorek &amp; Kristensen, 2018; Murray, 1907; Pilato et al., 2003), the Malay Archipelago ( locus typicus on Lombok, Marcus, 1928), and Australia (Claxton, 2004), E. pusae is likely another species with a broad geographic distribution in the tropics.</p><p>Genus: Kristenseniscus Gąsiorek et al., 2019</p><p>There are three clear morphotypes within Kristenseniscus (as currently defined, see Discussion and the problem of diphyly of the genus): (i) pronounced epicuticular matrix in the form of granules and ridges, the latter dividing scapular, paired segmental, and caudal plates into well-demarcated subplates; rare micropores sometimes present ( K. tessellatus, Fig. 9A); (ii) pronounced epicuticular matrix in the form of granules and ridges, the latter dividing only scapular and caudal plates into poorly-demarcated subplates; micropores absent ( K. limai, Fig. 9B); (iii) epicuticular matrix reduced (with the exception of posterior portions of median plates and anterior margins + centroposterior portions of paired segmental plates), micropores in all plates (the K. walteri complex; Figs. 9C, 10). In the walteri complex, the false subdivision of dorsal plates into subplates may be limited to posterior portions of paired segmental plates (Fig. 9C); the endocuticular layer can be complex, exhibiting well-developed pillars arranged in multangular groups (Fig. 9C, insert). Other species in the K. walteri complex have well-demarcated subplates, which may be regular (Fig. 10A) or with fuzzy margins (Fig. 10B).</p><p>Species: Kristenseniscus exanthema sp. nov.</p><p>Tables 6, 7 and 8, Figs. 11, 12 and 13</p><p>sp the proportion between the length of a given structure and the length of the scapular plate,?—unknown</p><p>Locus typicus and type material: 3°23′53″S, 129°15′41″E, 109 m asl; Indonesia, the Moluccas, <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=129.26138&amp;materialsCitation.latitude=-3.3980553" title="Search Plazi for locations around (long 129.26138/lat -3.3980553)">Seram</a>, <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=129.26138&amp;materialsCitation.latitude=-3.3980553" title="Search Plazi for locations around (long 129.26138/lat -3.3980553)">Tehoru</a>; moss and lichen from tree bark. Holotype (mature female on the slide ID.816.04), twelve paratypic females, seven juveniles and three larvae (slides ID.701.01–2, 4, 6, 8–10, ID.722.01–2, ID.816.03–4, 6–8, 10–12, ID.817.03). <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=129.26138&amp;materialsCitation.latitude=-3.3980553" title="Search Plazi for locations around (long 129.26138/lat -3.3980553)">All</a> are deposited in the <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=129.26138&amp;materialsCitation.latitude=-3.3980553" title="Search Plazi for locations around (long 129.26138/lat -3.3980553)">Institute of Zoology</a> and Biomedical Research of the Jagiellonian University in Kraków, Poland.</p><p>Etymology: From Latin exanthema = rash. The name refers to the morphology of dorsum, calling to mind the human bumpy hives (urticaria). A noun standing in apposition.</p><p>Mature females (i.e. from the third instar onwards; measurements in Table 6). Body plump (Figs. 11, 12) and dark orange. Minute red eyes present. Primary and secondary clavae typical of the Echiniscus - type; peribuccal cirri with well-developed cirrophores. Cirrus A of moderate length (30–50% of the body length), with cirrophore; flagellum sometimes bifid (Fig. 12A).</p><p>Dorsal plates are strongly sclerotised and well-demarcated from each other, with greatly modified tessellatus type of sculpturing, in which epicuticular granules are restricted mainly to anterior and posterior margins of paired segmental plates (Fig. 11), or otherwise may be absent (Fig. 12B). Instead, most of the surface of dorsal armour is sculptured with conspicuous micropores, and each micropore is surrounded with dark, usually multangular endocuticular matrix (Figs. 11, 12). The sculpturing vanishes in the anterior portions of median plate m2, anterior and lateralmost portions of paired segmental plates, the narrow patch anterior to the caudal (terminal) plate in the position of m3, and in a triangular zone of the caudal plate delimited by caudal incisions and lateral plate margins (Figs. 11, 12A). The cephalic plate is uniform. The cervical (neck) plate is in the form of a narrow, rectangular belt, adjacent to the scapular plate; always unsculptured (Figs. 11, 12). The scapular plate is not divided into subplates, without incisions or sutures. Two median plates: m1 unipartite, whereas m2 bipartite. Two pairs of large segmental plates, their posterior portions are divided into three subplates by two weakly marked semicircular sutures (Figs. 11, 12A). One pair of intersegmental lateral platelets embedded posteriorly to the scapular plate, and the second pair placed between paired segmental plates, both always unsculptured (Fig. 11). The caudal (terminal) plate non-facetted and not divided into subplates, analogously to the scapular plate (Figs. 11, 12).</p><p>Ventral cuticle smooth; a hexapartite gonopore placed anteriorly to a trilobed anus, lying between legs IV. Sculptured pedal plates and pulvini are absent. Spine I is very thin (Fig. 11). Dentate collar IV is composed of numerous short teeth. Papilla on leg IV is long (Figs. 11, 12A). Claws IV are higher than claws I–III. External claws on all legs are smooth. Internal claws with needle-like spurs positioned at ca. 20% of the claw height, tightly adjacent to the branches and homomorphic on all claws; spurs may be asymmetrically missing on one claw (Fig. 11A, inserts).</p><p>Juveniles (i.e. from the second instar onwards; measurements in Table 7). Clearly smaller than adult females. Qualitatively similar to adults, excluding the fact that epicuticular granules are more easily discernible (Fig. 13). Gonopore absent.</p><p>Larvae (i.e. the first instar; measurements in Table 8). Clearly smaller than juveniles. No larger morphological disparities regarding older instars. Gonopore and anus absent.</p><p>sp the proportion between the length of a given structure and the length of the scapular plate,?—unknown</p><p>Eggs. Up to two eggs per exuvia were found.</p><p>Phylogenetic position: Kristenseniscus exanthema sp. nov. is most closely related to Kristenseniscus cf. walteri 2 (Fig. 2 and SM.3).</p><p>Phenotypic differential diagnosis: Kristenseniscus exanthema sp. nov. is the only formally described member of Kristenseniscus that has uniform scapular and caudal plates, without subplates. Kristenseniscus cf. walteri 3 from Gunung Kinabalu (Borneo), not described in this work due to the lack of molecular data and the scarcity of individuals, shares this morphological state with K. exanthema sp. nov. but is easily distinguishable based on the endocuticular morphology resembling leopard spots under PCM (Fig. 9C, insert).</p><p>Species: Kristenseniscus limai (da Cunha &amp; do Nascimento Ribeiro, 1964)</p><p>Amendments to the morphology of the species (based on the populations listed in Table 1 that correspond with the morphotype depicted by da Cunha and do Nascimento Ribeiro): Anterior and posterior portions of paired segmental plates weakly demarcated (Figs. 14, 15A). Three pairs of intersegmental lateral platelets: the first embedded micrometer) of selected morphological structures of juveniles of Kristenseniscus exanthema sp. nov. (type series)</p><p>mounted in Hoyer’s medium posteriorly to the scapular plate, the second placed between paired segmental plates, and the third embedded posteriorly to the second paired segmental plate (Fig. 14). Subplates of the scapular plate developed only in its anterior half, with poorly developed epicuticular granules (Figs. 14A, C, 15B) or no granules (Fig. 14B). Each posterior portion of a paired segmental plate with a slightly semicircular, lateral epicuticular ridge formed by merged granules (Figs. 14A–C), not visible in juveniles (Fig. 14D). Caudal plate with four centrally positioned subplates and three epicuticular ridges: one at the posterior plate margin and two lateral (Figs. 14A–C, 15C). Only pedal plate IV was sculptured. Claws anisonych, as claws IV are on average slightly higher than claws I–III, with small and homomorphic spurs (Fig. 14D, insert).</p><p>sp the proportion between the length of a given structure and the length of the scapular plate</p><p>Species: Kristenseniscus tessellatus (Murray, 1910)</p><p>Amendments to the morphology of the species (based on the populations listed in Table 1 that correspond with the morphotype depicted by Murray): Two pairs of intersegmental lateral platelets: the first embedded posteriorly to the scapular plate and the second placed between paired segmental plates (Fig. 16A and fig. 1 in Dastych, 1997a). The pattern of subdivision of the scapular, paired segmental and caudal plates by epicuticular ridges into subplates is stable and well-identifiable due to the strong sclerotisation of the micrometer) of selected morphological structures of larvae of Kristenseniscus exanthema sp. nov. (type series)</p><p>mounted in Hoyer’s medium plates (Figs. 16 and 17). Claws heteronych, as IV are higher than claws I–III, with heteromorphic spurs, which are positioned slightly higher on claw IV branches and more divergent from them than spurs on claw I–III branches (Fig. 18).</p><p>sp the proportion between the length of a given structure and the length of the scapular plate,?—unknown</p><p>Genus: Pseudechiniscus Thulin, 1911 (amended by Vecchi et al., 2016)</p><p>Subgenus: Meridioniscus Gąsiorek et al., 2023</p><p>Species: Pseudechiniscus (Meridioniscus) celebesiensis sp. nov.</p><p>Pseudechiniscus (M.) sp. 1 in Gąsiorek et al. (2021a)</p><p>Tables 9 and 10, Figs. 19 and 20</p><p>Locus typicus and type material: 1°44′59″S, 120°32′16″E, 701 m asl; Indonesia, Celebes, Sulawesi Tengah, Terjun Saluopa; moss from tree bark. Holotype (mature female on the slide ID.411.05), seven paratypic females, three juveniles and one larva (slides ID.411.01–7, mounted together with a female of Pseudechiniscus (M.) angelusalas Roszkowska et al., 2020 and four females of P. (M.) quadrilobatus Iharos, 1969. All are deposited in the Institute of Zoology and Biomedical Research of the Jagiellonian University in Kraków, Poland.</p><p>Etymology: The name signifies terra typica. An adjective in the nominative singular.</p><p>Mature females (i.e. from the third instar onwards; measurements in Table 9). Body is small and slender (Fig. 19), orange. Minute black eyes are present in alive specimens, dissolving during mounting in Hoyer’s medium. Primary and secondary clavae elongated (dactyloid); peribuccal cirri with well-developed cirrophores. Cirrus A very short (&lt;25% of the body length), with cirrophore.</p><p>Dorsal plates weakly sclerotised, but well-demarcated from each other, with the modified Pseudechiniscus type of sculpturing, in which typical endocuticular pillars form ornamented belts (Fig. 19A). The space between these belts of pillars is completely smooth. Pillars are unequal in size: the largest can be found in the scapular and caudal (terminal) plates, medium-sized—in the centromedian portions of the remaining plates, and the smallest—in the lateral portions of paired segmental and intersegmental plates. Pillars are randomly joined by minute striae visible under 1000 ×magnification in LCM. The cephalic plate is uniform, with a posterior incision. The cervical (neck) plate clearly delimited from both cephalic and scapular plates as a belt of pillars (Fig. 19A). The scapular plate divided into two parts by a central transverse suture. Two unipartite median plates m1–2. Two pairs of segmental plates, poorly divided into anterior and posterior portions. One pair of intersegmental lateral platelets embedded posteriorly to the scapular plate, and the second pair placed between paired segmental plates. The pseudosegmental plate IV’ merged indistinctly with the caudal plate on the lateral sides of the body, only in the median line there is a sclerotised border with a pair of robust teeth-like projections. Incisions absent in the caudal plate (Fig. 19A).</p><p>Ventral cuticle with a system of epicuticular thickenings (Figs. 19B, 20) that replaced the usual pattern composed of endocuticular pillars (the largest remaining accumulation of pillars is present in the subcephalic zone; Fig. 20). A hexapartite gonopore placed anteriorly to a trilobed anus that lies between legs IV. Pedal plates and pulvini are absent, instead patches of pillars are present centrally on each leg. Spine I and dentate collar IV are absent. Papilla on leg IV is present (Fig. 19B). Claws I–IV of equal heights, very slender and spurless (Fig. 19A, insert).</p><p>Juveniles (i.e. from the second instar onwards; measurements in Table 10). Clearly smaller than adult females. Qualitatively alike adults. Gonopore absent.</p><p>Larvae (i.e. the first instar; measurements in Table 10). Clearly smaller than juveniles. No larger morphological disparities regarding older instars. Gonopore and anus absent.</p><p>Eggs. Unknown.</p><p>Phylogenetic position: Pseudechiniscus (M.) celebesiensis sp. nov. is one of the most basal unveiled lineages within the subgenus Meridioniscus according to the analyses presented in Gąsiorek et al. (2021a) ( Pseudechiniscus (M.) sp. 1 therein).</p><p>Phenotypic differential diagnosis: There are two known Pseudechiniscus (Meridioniscus) species exhibiting teeth on the posterior margin of the pseudosegmental plate IV’ and spurless claws, but P. (M.) celebesiensis sp. nov. can be differentiated from:</p><p>• P. (M.) bidenticulatus, with locus typicus in Java (Bartoš, 1963), by the distribution of endocuticular pillars in dorsal plates (arranged in ornamented belts in P. (M.) celebesiensis sp. nov. vs uniformly distributed in P. (M.) bidenticulatus), and by the teeth IV’ morphology (robust and prominent in P. (M.) celebesiensis sp. nov. vs minute and weakly developed in P. (M.) bidenticulatus);</p><p>• P. (M.) yunnanensis, described from tropical southern China (Wang, 2009), by the distribution of endocuticular pillars in dorsal plates (arranged in ornamented belts and joined by tiny striae in P. (M.) celebesiensis sp. nov. vs uniformly distributed and joined by large striae in P. (M.) yunnanensis), and by the presence of median plate m3 (absent in P. (M.) celebesiensis sp. nov. vs present in P. (M.) yunnanensis).</p></div>	https://treatment.plazi.org/id/2A7987F0FFC8073AFCB2F97BF20DF928	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Gąsiorek, Piotr;Michalczyk, Łukasz	Gąsiorek, Piotr, Michalczyk, Łukasz (2025): Novel integrative data for Indomalayan echiniscids (Heterotardigrada): new species and old problems. Organisms Diversity & Evolution 25 (1): 81-116, DOI: 10.1007/s13127-023-00628-5, URL: https://doi.org/10.1007/s13127-023-00628-5
2A7987F0FFDD0736FF75FA0EF722FF67.text	2A7987F0FFDD0736FF75FA0EF722FF67.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Kristenseniscus	<div><p>Diphyly of Kristenseniscus</p><p>The evident non-monophyly of Kristenseniscus (Fig. 2) should be solved by an erection of a new genus for K. limai . Despite an overall similarity, this species consistently differs from K. tessellatus and the K. walteri complex in two important characters: (1) the presence of three pairs of intersegmental platelets inserted between the segmental (scapular, paired segmental and caudal) plates, instead of two pairs in the latter, as clearly the third pair, positioned between the second paired segmental plate and caudal plate, is absent in Kristenseniscus s.s. (Fig. 21); and (2) the subdivision of plates into false subplates is limited to the scapular and caudal plates, but it is ancestrally present in all segmental plates in Kristenseniscus s.s. (Fig. 21). The intersegmental platelets of K. tessellatus were drawn in the original description (Murray, 1910), and later shown based on the material from Hawaii (Dastych, 1997a). Their relatively strong sclerotisation may have caused the effect of “nimbus” described for empty exuviae (Murray, 1910). The abovementioned new genus is not established due to nomenclatorial problems with its probable type species (see below) and a need for sequencing some enigmatic echiniscid taxa, such as Zealandiscus palmai (Dastych, 1997b), which is similar to K. limai in the development of large epicuticular granules on the dorsal plates (Gąsiorek et al., 2021c), potentially signifying a close phylogenetic affinity of the two species.</p></div>	https://treatment.plazi.org/id/2A7987F0FFDD0736FF75FA0EF722FF67	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Gąsiorek, Piotr;Michalczyk, Łukasz	Gąsiorek, Piotr, Michalczyk, Łukasz (2025): Novel integrative data for Indomalayan echiniscids (Heterotardigrada): new species and old problems. Organisms Diversity & Evolution 25 (1): 81-116, DOI: 10.1007/s13127-023-00628-5, URL: https://doi.org/10.1007/s13127-023-00628-5
2A7987F0FFDE0732FCCDFF3DF0D9F97E.text	2A7987F0FFDE0732FCCDFF3DF0D9F97E.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Kristenseniscus	<div><p>Sympatry of Kristenseniscus in the Malay Archipelago</p><p>The genus Kristenseniscus has a pantropical/subtropical distribution, but the exact biogeography of its members is yet to be recognised. The oldest described species, K. tessellatus, has confirmed records mainly from the Pacific Ocean basin (Murray, 1910; Utsugi, 1993; Dastych, 1997a; Claxton, 2004; Li et al., 2008; Suzuki, 2017; see Suzuki et al., 2018 for a summary), reaching Japan in the North, and only two reports from the Indian Ocean basin (Maucci &amp; Durante Pasa, 1980; Pilato &amp; Binda, 1990; Pilato &amp; Lisi, 2003). In Gąsiorek et al. (2019b) and in the present study, individuals originating from the previously unsurveyed areas, that should be inhabited by this species and incorporated within its geographic range, were studied. The molecular diversity of K. tessellatus specimens from Taiwan, Borneo, Celebes and the Moluccas is low (Fig. 2) and supports the high dispersal potential of this taxon. At the same time, the species is phenotypically very stable and we found no discrepancies between the abovementioned individuals and a Japanese specimen provided by Atsushi Suzuki.</p><p>In Gąsiorek et al. (2019b), we signalised problems with the identity of Kristenseniscus limai (da Cunha &amp; do Nascimento Ribeiro, 1964) by designating it as nomen inquirendum. Although far from being elaborative, the original description does stress the presence of “carination and facetting” in the scapular and caudal (terminal) plates of the African type series, also depicted in a standard way of that time, i.e. neither very detailed nor too general. As it so happens, two years later Schuster and Grigarick (1966), most likely unaware of the Portuguese contribution, described K. kofordi (Schuster &amp; Grigarick, 1966) from Santa Cruz Island belonging to the Galápagos archipelago, showing in greater detail the dorsal sculpturing as consisting of very large epicuticular granules and ridges forming subplates in the scapular and caudal plates (short epicuticular ridges are present also in the lateralmost portions of paired segmental plates in this species). While studying the former Echiniscus tessellatus group (= Kristenseniscus), Pilato and Lisi (2003) analysed the Madagascan material collected by Maucci (1993), dissecting three morphotypes that correspond with K. tessellatus, K. kofordi and K. walteri . Beasley and Cleveland (1996), who reported K. limai from the Indomalayan part of China, noted the similarity between K. limai and K. kofordi, but they enumerated alleged differences between the two species: the body size, the cirrus A length and the regularity of dorsal sculpturing. Importantly, K. kofordi is not a larger species according to the present state of knowledge, as Beasley and Cleveland (1996) claimed. da Cunha and do Nascimento Ribeiro (1964) clearly stated that the largest individuals of K. limai attained 142 µm in length, whereas Schuster and Grigarick (1966) determined the body size range of K. kofordi as 120–190 µm. The adult individuals originating from the Sulawesian populations we examined ranged between 130–190 µm (N = 20). We are also sceptical about the presumed disparity in the cirrus A length, as da Cunha and do Nascimento Ribeiro (1964) wrote it measured for 14–15 µm, and Schuster and Grigarick (1966) provided only a value for the holotype (22 µm). Putatively longer cirrus A in K. kofordi may be a derivative of significantly larger body size of a specimen measured (190 µm), as well as a consequence of unreliable morphometric data provided by the authors since Schuster and Grigarick (1966) have already been shown to overestimate measurements of cirrus A in the case of Echiniscus cavagnaroi (Meyer, 2016) . The Sulawesian examples ranged 16–25 µm in this criterion. Finally, the comparison of the “regularity” of sculpturing between the two species seems to be a result of an insufficient sample size being at the disposal of Beasley and Cleveland (1996). Based on the Sulawesian animals, it becomes clear that a species corresponding with the morphotype of K. kofordi (Figs. 9B, 14 and 15) is somehow intermediate in the stability of the development of dorsal sculpturing between very stable and invariant K. tessellatus (Fig. 9A) and a very variable K. walteri (Figs. 9C and 10) complex. Especially the lateral epicuticular ridges may be reduced in this species, leading to the morphotype drawn by da Cunha and do Nascimento Ribeiro (1964) as K. limai . In conclusion, in our opinion, there are no morphological differences between K. limai and K. kofordi; however, a formal status of a junior synonym cannot be assigned to K. kofordi, until new DNA barcodes for Afrotropical and Neotropical populations will confirm or reject this hypothesis. Before this happens, we are inclined to consider K. limai as a pantropical species, with reports from Angola (da Cunha &amp; do Nascimento Ribeiro, 1964), Venezuela (Grigarick et al., 1983), Costa Rica (Kaczmarek &amp; Michalczyk, 2010), Mexico (Pilato &amp; Lisi, 2006), Colombia (Lisi et al., 2017), Alabama (Christenberry, 1979), Florida (Meyer, 2006), Louisiana (Hinton &amp; Meyer, 2007), the Galápagos archipelago (Schuster &amp; Grigarick, 1966), tropical part of China (Beasley &amp; Cleveland, 1996) and the Malay Archipelago (present study).</p><p>sp the proportion between the length of a given structure and the length of the scapular plate</p><p>sp the proportion between the length of a given structure and the length of the scapular plate Fig. 19 General morphology of</p><p>Pseudechiniscus celebesiensis</p><p>sp. nov. (PCM): A holotype,</p><p>dorsal view; B holotype, ventral view. Insert shows claws II of a paratype. Scale bars in μm</p><p>Our study revealed at least four species of the walteri complex inhabiting the Malay Archipelago ( K. exanthema sp. nov. and K. cf. walteri 1–3), of which three are characterised with DNA barcodes (Fig. 2). Unfortunately, it is not possible to ascertain whether K. walteri s.s. is present in the Malay Archipelago without molecular data for populations of this species from its terra typica, i.e. Madagascar (Pilato &amp; Lisi, 2003). However, morphotypes present in the very limited type series (only a holotype and two paratypes) of K. walteri raise concerns whether this species was not described based on specimens representing two different biological species, taking into account that the morphological discrepancies in the Malay examples are minute (Fig. 10) and that the sculpturing on the scapular plate of the paratype (Fig. 2a in Pilato &amp; Lisi, 2003) is different from that of the holotype (Fig. 2b in Pilato &amp; Lisi, 2003). Moreover, multiple Kristenseniscus species inhabit similar habitats in the Malay Archipelago, and often more than one species is found in the same sample (Table 1). We hypothesise that the geographic range of K. walteri may be limited to Madagascar. In contrast, K. limai, having a pantropical distribution, is probably the most widespread species in the genus. Furthermore, K. tessellatus exhibits a broad distribution, encompassing the Pacific Ocean basin, but not reaching Afrotropical and Neotropical regions. Thus, the walteri complex seems to have radiated in the Malay Archipelago in sympatry with its congeneric species (the walteri morphotype(s) are unknown for tropical Australia and continental Indomalayan Asia). The evolutionary causes for this phenomenon are yet to be unravelled, but several species living in the same islands and likely nowhere else in the world indicate that this complex may be a weak dispersalist compared with its more distant relatives: K. tessellatus and K. limai . Analogous differentiation in dispersal potential within a complex of relatively close kin has been recently documented for a eutardigrade genus Paramacrobiotus (Guidetti et al., 2019) . A summary of Kristenseniscus distribution is presented in Fig. 22.</p><p>The walteri complex clearly represents an advanced state regarding the dorsal sculpturing within the genus: the pseudodivision of plates into subplates shows various stages of reduction (i.e. the return to the uniform, solid plates), reaching completely uniform scapular and caudal plates in K. exanthema sp. nov. Simultaneously, micropores are fully developed and dominate the entire dorsal armour (Figs. 11, 12, 13). Lastly, we pinpoint a morphological peculiarity of the entire genus: in specimens with fully extended hind legs, an elongated (dactyloid) papilla IV is apparent (Figs. 9A, 11A, 12A). Such a shape of this structure is atypical for the Echiniscus -like genera, which usually exhibit a tubby papilla IV.</p></div>	https://treatment.plazi.org/id/2A7987F0FFDE0732FCCDFF3DF0D9F97E	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Gąsiorek, Piotr;Michalczyk, Łukasz	Gąsiorek, Piotr, Michalczyk, Łukasz (2025): Novel integrative data for Indomalayan echiniscids (Heterotardigrada): new species and old problems. Organisms Diversity & Evolution 25 (1): 81-116, DOI: 10.1007/s13127-023-00628-5, URL: https://doi.org/10.1007/s13127-023-00628-5
