taxonID	type	description	language	source
03CDAC58FFD8FFBFA084FF6BFCD6D1D5.taxon	description	(Figs 2 – 12; 13 A, C, D) (Figs 1 − 8; 11; 12, not Figs 9; 10; 13; 14 in: Behan-Pelletier and Eamer 2005, see Morphological remarks below)	en	Ermilov, Sergey G., Makarova, Olga L. (2025): New records of Zachvatkinibates (Acari, Oribatida, Punctoribatidae) in Russia, with description of ontogenetic instars of amphi-Pacific littoral Z. nortoni. Zootaxa 5740 (1): 43-65, DOI: 10.11646/zootaxa.5740.1.6, URL: https://doi.org/10.11646/zootaxa.5740.1.6
03CDAC58FFD8FFBFA084FF6BFCD6D1D5.taxon	materials_examined	Material. Adults (26 specimens: 16 females and 10 males) and juvenile instars (four larvae, seven protonymphs, nine deutonymphs, and four tritonymphs) were collected in the Magadan Region (North-East Asia, Russia), in the vicinity of Ola Lagoon (Figs 1 A, B). Of them: five females, six males, and all juvenile instars originated from the western coast of the lagoon, 59 ° 34 ’ 33 ” N, 151 ° 20 ’ 49 ” E, marine debris of Zostera on the sandy seashore (Fig. 1 B), 22. IX. 2018 (O. L. Makarova); 11 females and four males from the same district, but from the northern coast of the lagoon, 59 ° 35 ’ 54 ” N 151 ° 21 ’ 38 ” E, in fucoid seaweeds, 26. VII. 2017 (M. S. Bizin). Three females and six males (in 70 % solution of ethanol with a drop of glycerol) are in the personal collection of the first author. Other adults (mounted in Hoyer-medium on three permanent slides) are kept in the collection of the Laboratory of Synecology, IPEE RAS.	en	Ermilov, Sergey G., Makarova, Olga L. (2025): New records of Zachvatkinibates (Acari, Oribatida, Punctoribatidae) in Russia, with description of ontogenetic instars of amphi-Pacific littoral Z. nortoni. Zootaxa 5740 (1): 43-65, DOI: 10.11646/zootaxa.5740.1.6, URL: https://doi.org/10.11646/zootaxa.5740.1.6
03CDAC58FFD8FFBFA084FF6BFCD6D1D5.taxon	description	Supplementary description of adult. Measurements. Body length: 570 – 575 (females), 510 – 540 (males); notogaster width: 315 (females), 270 – 300 (males). Integument. Body colour brown. Body surface microsculpturing tuberculate; interlamellar region, pedotectum I, dorsal side of tutorium, lateral parts of epimeres I, II, and antiaxial side of leg trochanters III, IV with dense, short or long, thin ridges (Figs 12 C, D, E); antiaxial side of leg femora I, II and paraxial side of leg femora III, IV with sparse, transverse ridges. Prodorsum (Figs 2 A; 3 A; 4 A, B; 5 C; 12 C, D; 13 A, C). Rostrum nearly rounded or slightly undulate, usually with two small lateral tubercles (sharp teeth absent; Figs 5 C; 12 D; 13 C). Rostral region with median bulge. Lamella (including cusp) about 2 / 3 length of prodorsum; cusp short, truncate; translamella usually thin (Fig. 13 A). Tutorium (including cusp) about 3 / 4 length of prodorsum; cusp broad, with three to six small teeth (Fig. 12 D). Genal tooth well defined, elongate triangular. Rostral (67 – 71), lamellar (79 – 86), and interlamellar (111 – 143) setae stiff, acute, barbed; basal part of ro covered by tutorium; bothridial seta (45 – 52) clavate, with short stalk and short, rounded distally, barbed head; exobothridial seta (22 – 26) setiform, roughened. Bothridium covered or not by anterior margin of notogaster in dorsal aspect. One pair of curved ridges (connected or not medially) developed posterior to insertions of interlamellar setae. Dorsosejugal porose area and dorsophragma not observed. Notogaster (Figs 2 A, B; 3 A; 4 A, B; 5 A, B; 13 D). Anterior margin of notogaster nearly straight or with very weak medial protrusion (Fig. 13 D). Pteromorph broadly rounded laterally, with distinct line of desclerotization, its lateral margin distinct but anterior margin wearing thin, membranous, often looking dentate (Fig. 13 D). Octotaxic system showing sexual dimorphism: male with two pairs of nearly circular porose areas Aa (22 – 26) and A 1 (56 – 67), and one unpair spacious porose area (fused two pairs of A 2 and A 3) covering posterior part of notogaster (expanding laterally of setae h 1 and h 2 and medially of setae p 3 and lyrifissures ih; setae p 1 and p 2 located on A 2 + A 3); female with four pairs of nearly circular porose areas (Aa: 22; A 1: 19 – 22; A 2: 15 – 22; A 3: 22 – 30). Ten pairs of notogastral setae (h 1: 56 – 60; p 1 – p 3: 26 – 34; others: 37 – 35) slightly stiff, acute, nearly smooth, roughened or shortly barbed distally; h 3 inserted close and between porose areas A 1 and A 2 + A 3 in males. Opisthonotal gland opening and all lyrifissures distinct. Posterior of notogaster of male extending well beyond margin of ventral plate, unlike that of female (Figs 4 A, B). Posterior notogastral tectum incomplete medially in both genders (Figs 5 A, B). Gnathosoma (Figs 5 D – F). Subcapitulum size: 112 – 116 × 90 – 94; subcapitular (a, h: 19; m: 30) and adoral (15) setae setiform, slightly barbed. Palp length: 82 – 86; setal formula: 0 - 2 - 1 - 3 - 9 (+ ω); postpalpal seta (7) spiniform. Axillary saccule distinct, slightly oblong. Chelicera length: 112 – 116; setae (cha: 45; chb: 30) setiform, barbed. Lateral podosomal and epimeral regions (Figs 2 B; 3 B; 4 A, B; 5 C). Humeral porose areas Am and Ah not observed, but unclear porosity slightly visible in humeral region. Pedotectum I concave dorsally, usually with one small tooth, rarely with two teeth. Custodium strong, reaching epimere I. Discidium triangular. Circumpedal carina distally fused to custodium. Epimeral setal formula: 3 - 1 - 3 - 3; setae 3 c (26 – 30) and 1 b, 1 c, 3 b (19) setiform, slightly barbed; 1 a, 2 a, 3 a, 4 a, 4 b, and 4 c (11 – 13) setiform, roughened; 1 c slightly thicker than other setae. Anogenital region (Figs 2 B; 3 B; 4 A, B; 5 A, B). Six pairs of genital (11 – 13), one pair of aggenital (15 – 19), two pairs of anal (13 – 19), and three pairs of adanal (15 – 22) setae setiform, roughened. Adanal lyrifissure located close and parallel to anterior half of anal plate. Postanal porose area narrowly oblong (37 – 41 × 5 – 7). Legs (Figs 6 A – D). Median claw distinctly thicker than lateral claws, all slightly barbed dorsally. Dorsoparaxial porose area on femora I – IV and on trochanters III, IV distinct; proximoventral porose area on tarsi I – IV and distoventral porose area on tibiae I – IV present, but poorly visible. Tibia II – IV with small dorsodistal triangular process. Formulas of leg setation and solenidia: I (1 - 5 - 3 - 4 - 20) [1 - 2 - 2], II (1 - 5 - 3 - 4 - 15) [1 - 1 - 2], III (2 - 2 - 1 - 3 - 15) [1 - 1 - 0], IV (1 - 2 - 2 - 3 - 12) [0 - 1 - 0]; homology of setae and solenidia as indicated in Table 1; famulus short, erect, slightly swollen distally, inserted between seta ft " and solenidion ω 2; seta s of tarsus I eupathidial; lateral seta l " of tibia II, III and l' of genu III thick; leg femur III with two setae (l' absent); solenidia ω 1 and ω 2 of tarsus II and σ of genu III slightly thickened, rounded distally, ω 1 and ω 2 of tarsus I and φ 2 of tibia I rod-like, other solenidia subflagellate; solenidion φ 2 of tibia I located on dorsodistal tubercle anterior to solenidion φ 1. Description of juvenile instars (Figs 7 – 11; 12 G). Measurements. Body length of LA: 225 – 255, PN: 285 – 315, DN: 345 – 375, TN: 450 – 480. Body width of LA: 105 – 120, PN: 135 – 150, DN: 180 – 195, TN: 240 – 255. Integument. Body colourless in LA and PN to pale yellow in DN and TN; prodorsal and gastronotic shields (macrosclerites) and legs light brown in PN, DN, and TN; posterolateral sclerite around opisthonotal gland opening brown in all instars. Cuticle densely microporose (micropores poorly visible, but distinct on macrosclerites in DN and TN, and on sclerite around opisthonotal gland opening in all instars); dorsal side of gastronotic region with strong, transverse fold posterior to setae c - row in PN, DN, and TN; lateral part of body and anogenital region (also sternal region and partially dorsal part of gastronotum in LA) with sparse or dense folds (variability present in specimens); subcapitular mentum, podosomal region, and apodemal regions partially microtuberculate in PN, DN, and TN; paraxial side of all leg femora with sparse, transverse ridges. Prodorsum (Figs 7 A – D; 9 A, B). Relatively short, about 1 / 2 (in LA) and 2 / 5 (in PN, DN, and TN) length of gastronotic region. Rostrum rounded. Dorsal side of prodorsum with large prodorsal shield (macrosclerite). Lateral prodorsal carina poorly visible. Rostral (LA: 30 – 34; PN: 30 – 37; DN: 34 – 41; TN: 41 – 49), lamellar (LA: 19 – 22; PN: 22 – 26; DN: 22 – 30; TN: 34 – 37), and interlamellar (LA: 41 – 45: PN: 45 – 49; DN: 49 – 52; TN: 64 – 67) setae stiff, acute or blunt, barbed; bothridial seta (LA: 19 – 22; PN: 22 – 26; DN: 26 – 30; TN: 26 – 34) clavate, with short stalk and short, rounded distally, barbed head; exobothridial seta (LA: 9 – 11; PN: 7; DN: 11 – 13, TN: 15) thin, acute or blunt, roughened. Transverse scale located posterior to bothridium. Gastronotic region (Figs 7 A – D; 8 A – D; 9 A – D). Dorsal side with large gastronotic shield (macrosclerite) bordered by strong fold anteriorly and poorly visible ecdysial cleavage line (line of dehiscence) laterally and posteriorly in all instars. Humeral organ present, positioned anterior to seta c 3 in LA versus lateral or anterolateral to seta c 3 in PN, DN, and TN. Posterolateral sclerite well developed. Larva with 12 pairs (of them, c 1, c 2, da, la, dm, lm, dp, lp, and h 1 located on macrosclerite), nymphs with 15 pairs (of them, da, la, dm, lm, dp, lp, h 1 – h 3, and p 1 located on macrosclerite) of stiff, acute or blunt, barbed setae (except setiform, roughened setae p 1 – p 3 in PN); seta h 3 thinner than other setae in LA versus setae p 1 – p 3 thinner than other setae in PN, DN, and TN. Seta c 2 with sclerite at the base in PN, DN, and TN. Length of gastronotic setae: LA: dp, lp, h 1: 30 – 37, c 3, dm: 22 – 26; h 3: 11 – 13, others: 19 – 22; PN: c 3, h 1, h 3: 22 – 26, c 2: 13 – 15, p 1 – p 3: 11 – 13, others: 15 – 19; DN: c 3: 30 – 34, h 1: 28 – 30, c 1, c 2: 22 – 24, p 1 – p 3: 15 – 17, others: 19; TN: c 3: 41 – 49, h 1: 34 – 37, c 1, c 2: 30, p 1 – p 3: 15 – 17, others: 22 – 24. Opisthonotal gland opening and all cupules distinct; ih and ips appearing in normal ontogenetic pattern. Gnathosoma (Figs 9 E – G). Subcapitulum size: LA: 45 – 49 × 56; PN: 56 – 60 × 64 – 67; DN: 71 – 75 × 82 – 90; TN: 82 – 86 × 90 – 94; length of subcapitular setae a, h: LA, PN: 11; DN: 15; TN: 19; length of subcapitular seta m: LA: 11; PN: 11 – 13; DN: 19; TN: 22; length of adoral seta: LA, PN: 6; DN: 9; TN: 11; all setae setiform, slightly barbed. Palp length: LA: 41; PN: 49; DN: 60; TN: 67; palp setal formula in all instars: 0 - 2 - 1 - 3 - 9 (+ ω); solenidion connected with eupathidium; length of postpalpal seta: LA, PN: 4; DN, TN: 6. Axillary saccule small, slightly oblong, poorly visible. Chelicera length: LA: 52 – 56; PN: 67; DN: 77 – 82; TN: 90 – 94; length of seta cha: LA: 19 – 22; PN: 22; DN: 30; TN: 34; length of seta chb: LA: 11 – 13; PN: 15 – 17; DN: 22; TN: 26; both setae setiform, barbed. Epimeral region (Figs 8 A – D; 9 A, B). Setal formulas for successive epimeres: LA: 3 - 1 - 2 (1 c as typical scale covering Claparède’s organ); PN 3 - 1 - 2 - 1; DN 3 - 1 - 2 - 2, TN 3 - 1 - 3 - 3; length of setae: LA: 9 – 11; PN: 11 – 13; DN: 13 – 19; TN: 19 – 22; all setae setiform, nearly smooth in LA and PN versus setiform, roughened in DN and TN. Anogenital region (Figs 8 A – D; 9 A – D). Ontogeny of genital (PN: 9 – 11; DN: 11; TN: 11 – 13), aggenital (DN: 11; TN: 11 – 13), anal (TN: 11 – 15), and adanal (DN: 11; TN: 11 – 15) setal formulas, PN to TN: 1 - 3 - 5, 0 - 1 - 1, 0 - 0 - 2, 0 - 3 - 3, respectively; all setae setiform, nearly smooth in LA and PN versus setiform, roughened in DN and TN; paraproctal setae absent. Cupule iad appearing in normal ontogenetic pattern. Legs (Figs 10 A – D; 11 A – D). Claw of each leg strong, slightly barbed dorsally. Ventroparaxial porose area on femora I – IV indistinct in LA and PN versus well visible in DN and TN; porose area on trochanters, tibiae, and tarsi not observed. Formulas of leg setation and solenidia: larva: I (0 - 2 - 2 - 3 - 16) [1 - 1 - 1], II (0 - 2 - 2 - 2 - 13) [1 - 1 - 1], III (0 - 2 - 1 - 1 - 13) [1 - 1 - 0]; protonymph: I (0 - 2 - 2 - 3 - 16) [1 - 1 - 2], II (0 - 2 - 2 - 2 - 13) [1 - 1 - 1], III (0 - 2 - 1 - 1 - 13) [1 - 1 - 0], IV (0 - 0 - 0 - 0 - 7) [0 - 0 - 0]; deutonymph: I (0 - 4 - 2 - 3 - 16) [1 - 2 - 2], II (0 - 4 - 2 - 3 - 13) [1 - 1 - 2], III (1 - 2 - 1 - 2 - 13) [1 - 1 - 0], IV (0 - 2 - 1 - 1 - 12) [0 - 1 - 0]; tritonymph: I (1 - 4 - 3 - 4 - 18) [1 - 2 - 2], II (1 - 4 - 3 - 4 - 15) [1 - 1 - 2], III (2 - 2 - 1 - 3 - 15) [1 - 1 - 0], IV (1 - 2 - 2 - 3 - 12) [0 - 1 - 0]; homologies of leg setae and solenidia as indicated in Table 1.	en	Ermilov, Sergey G., Makarova, Olga L. (2025): New records of Zachvatkinibates (Acari, Oribatida, Punctoribatidae) in Russia, with description of ontogenetic instars of amphi-Pacific littoral Z. nortoni. Zootaxa 5740 (1): 43-65, DOI: 10.11646/zootaxa.5740.1.6, URL: https://doi.org/10.11646/zootaxa.5740.1.6
03CDAC58FFCAFFA7A084FF23FB10D603.taxon	description	(Figs 13 B, E, I)	en	Ermilov, Sergey G., Makarova, Olga L. (2025): New records of Zachvatkinibates (Acari, Oribatida, Punctoribatidae) in Russia, with description of ontogenetic instars of amphi-Pacific littoral Z. nortoni. Zootaxa 5740 (1): 43-65, DOI: 10.11646/zootaxa.5740.1.6, URL: https://doi.org/10.11646/zootaxa.5740.1.6
03CDAC58FFCAFFA7A084FF23FB10D603.taxon	materials_examined	Material. 28 adults (18 females and 10 males), North-East Asia, Magadan Region, Taujsk village vicinity, 59 ° 44 ’ 04 ” N, 149 ° 23 ’ 16 ” E, sandy seashore (Figs 1 C, D), under halophilous plants (Seneciо pseudoarnica, Mertensia maritima, Leymus mollis), 24. VII. 2017 (M. S. Bizin). All adults (mounted in Hoyer-medium on three permanent slides) are kept in the collection of the Laboratory of Synecology IPEE RAS.	en	Ermilov, Sergey G., Makarova, Olga L. (2025): New records of Zachvatkinibates (Acari, Oribatida, Punctoribatidae) in Russia, with description of ontogenetic instars of amphi-Pacific littoral Z. nortoni. Zootaxa 5740 (1): 43-65, DOI: 10.11646/zootaxa.5740.1.6, URL: https://doi.org/10.11646/zootaxa.5740.1.6
03CDAC58FFCAFFA7A084FF23FB10D603.taxon	discussion	Until now, Z. erimo was known only from the type locality on the sandy beach of Hyakunin-hama, Erimo-Misaki Cape, Hokkaido, Northern Japan (41 ° 55 ′ 28 ″ N, 143 ° 14 ′ 54 ″ E). Our record represents the first occurrence of Z. erimo in Russia, located approximately 2, 000 km north of the original locality, but it was associated with the same biotope — sandy seashores. The female to male ratio was 9: 5. Oviparous females (two from 18) contained one and six eggs. Analysis of the gut contents revealed a diverse diet, dominated by hyphae and solitary spores of micromycetes, as well as fragments of unicellular algae, whereas fungal ascospores and conidia were present but rare. Given these new records, the fauna of Russia now include eight species of Zachvatkinibates. Below we present an identification key for the adults, as well as a summary of the distribution and habitats of all known Zachvatkinibates species in Russia. Identification keys for described larvae and tritonymphs are also given.	en	Ermilov, Sergey G., Makarova, Olga L. (2025): New records of Zachvatkinibates (Acari, Oribatida, Punctoribatidae) in Russia, with description of ontogenetic instars of amphi-Pacific littoral Z. nortoni. Zootaxa 5740 (1): 43-65, DOI: 10.11646/zootaxa.5740.1.6, URL: https://doi.org/10.11646/zootaxa.5740.1.6
03CDAC58FFC8FFA4A084FF23FA0ED431.taxon	discussion	Nymphal instars are similar in many morphological characteristics, therefore, the key can also be applied to protonymphal and deutonymphal instars (except characteristics that are specific to each instar, for example, body size, number of genital setae).	en	Ermilov, Sergey G., Makarova, Olga L. (2025): New records of Zachvatkinibates (Acari, Oribatida, Punctoribatidae) in Russia, with description of ontogenetic instars of amphi-Pacific littoral Z. nortoni. Zootaxa 5740 (1): 43-65, DOI: 10.11646/zootaxa.5740.1.6, URL: https://doi.org/10.11646/zootaxa.5740.1.6
03CDAC58FFCEFFA2A084FF23FC5BD3C7.taxon	discussion	In Russia, members of the genus Zachvatkinibates are distributed mainly in the southern regions of the country (Central Asian species — Z. conjunctus, Z. latilamellatus, Z. nemoralis) and in the Far East (Pacific littoral species — Z. erimo, Z. maritimus, Z. nortoni, Z. volgini). One littoral species, Z. quadrivertex, occurs in both the Far East and the European North of Russia (Table 2). According to Table 2 and available habitat data from the literature (e. g., Behan-Pelletier 1988; Behan-Pelletier & Eamer 2005; Weigmann 2009; Bayartogtokh 2010; Shimano & Aoki 2019), Z. conjunctus, Z. latilamellatus, and Z. nemoralis inhabit various inland biotopes, whereas Z. erimo, Z. maritimus, Z. nortoni, Z. quadrivertex, and Z. volgini are generally confined to coastal habitats. Of these, only two species with broad distributions, the amphi-Pacific Z. maritimus and the Holarctic Z. quadrivertex, are occasionally found in some coastal forests (Ryabinin 2015). This distribution pattern is likely influenced by maritime air masses. A similar case has been documented in Scotland, where the strictly littoral oribatid mite, Ameronothrus lineatus (Thorell, 1871), was recorded in moss and grasses up to 300 m inland from the shore (Colloff 1984). We compiled all available information on the body sizes and habitats of all 19 described species of the genus Zachvatkinibates (Table 3). The genus comprises nearly equal numbers of seaside (9) and inland (10) species. Preferred inland biotopes are very diverse (forests, meadows, steppes, barrens), but almost all of them are associated with mountains, and all woodland records are attributable to only evergreen (mainly coniferous) forests. Littoral species tend to be larger than inland species, and this difference in mean body size is statistically significant at p <0.01. Differences were also found in the proportions of the proterosoma that is expressed in the different relative length of the bothridial seta (Table 3); its head either reaches (in inland species) or does not reach the level of the translamella (in littoral species). Apparently, the proterosoma is somewhat elongated in littoral species. Littoral habitats are often characterized by high humidity and an abundance of easily decomposable organic matter, which usually stimulate the development of abundant microflora and invertebrates (Remmert 1960; Strenzke 1962). It is interesting to note that among predatory mesostigmatic mites, the strictly littoral species Dissoloncha superba (Hull, 1923) (Macrochelidae) and Vulgarogamasus immanis (Berlese, 1903) (Parasitidae) are among the largest representatives of their families and free-leaving mites in general, reaching lengths of up to 1,700 μm and 2,700 μm, respectively (Bregetova & Koroleva 1960; Tikhomirov 1977).	en	Ermilov, Sergey G., Makarova, Olga L. (2025): New records of Zachvatkinibates (Acari, Oribatida, Punctoribatidae) in Russia, with description of ontogenetic instars of amphi-Pacific littoral Z. nortoni. Zootaxa 5740 (1): 43-65, DOI: 10.11646/zootaxa.5740.1.6, URL: https://doi.org/10.11646/zootaxa.5740.1.6
