New Oripodoidea (Acari, Oribatida) from Myanmar - BioOne

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Jul 6, 2017 - The oribatid mite taxa (Acari, Oribatida), superfamily Oripodoidea, families Scheloribatidae and Caloppiidae, genera Scheloribates Berlese ...
Systematic & Applied Acarology 22(7): 1022–1036 (2017) http://doi.org/10.11158/saa.22.7.10 Article

ISSN 1362-1971 (print) ISSN 2056-6069 (online)

http://zoobank.org/urn:lsid:zoobank.org:pub:A28A00D7-C6D5-4BE9-9AB4-F4394865C38B

New Oripodoidea (Acari, Oribatida) from Myanmar SERGEY G. ERMILOV Tyumen State University, Tyumen, Russia. E-mail: [email protected]

Abstract The oribatid mite taxa (Acari, Oribatida), superfamily Oripodoidea, families Scheloribatidae and Caloppiidae, genera Scheloribates Berlese, 1908 and Zetorchella Berlese, 1916, and the species Zetorchella orbiculata (Wen & Zhao, 1994), are recorded in Myanmar for the first time. Two new oripodoid species are described: Scheloribates natmaensis sp. nov. (Scheloribatidae) is most similar to S. yamaeensis Nakamura, Hashimoto, Nishi, Nakamura & Fujikawa, 2015, but differs by the larger body size, narrowly triangular rostrum, clavate, rounded distally bothridial setae and longitudinal stria on the notogaster; Zetorchella taungensis sp. nov. (Caloppiidae) is most similar to Z. nortoni Ermilov, Sidorchuk & Rybalov, 2010, but differs by the larger body size, longer notogastral setae, bothridial setae shorter than humeral notogastral setae c and notogastral setae c and p1 similar in length. Key words: mites, record, new species, systematics, morphology, Scheloribates, Zetorchella

Introduction This work is part of my study (Ermilov 2017) of the Myanmar oribatid mites (Acari, Oribatida) received from the collection of the Senckenberg Museum of Natural History (Görlitz, Germany) and includes data on the superfamily Oripodoidea. During taxonomic identification, I found three species of oripodoids; of these, two species are new to science, one belonging to the genus Scheloribates Berlese, 1908 (Scheloribatidae), from the nominative subgenus, and the other to the genus Zetorchella Berlese, 1916 (Caloppiidae), and one species, Zetorchella orbiculata (Wen & Zhao, 1994), which has been described from southern China. The main goal of the paper is to describe and illustrate the two new species. The superfamily Oripodoidea and listed families, genera and species are recorded for the first time in Myanmar. The genus Scheloribates comprises several subgenera (based on different opinions; e.g. Subías 2004, updated 2016; 2017; Ermilov & Anichkin 2014) and about 250 species and 20 subspecies, which have a cosmopolitan distribution collectively. The genus Zetorchella comprises 22 species, which are distributed in the Ethiopian, Neotropical, Oriental and southern Palaearctic regions (Subías 2004, updated 2017).

Material, methods, terminology and abbreviations The collection locality and habitat for each studied oripodoid species are given in the respective "Material examined" section. Specimens were mounted in lactic acid on temporary cavity slides for measurement and illustration. The body length was measured in lateral view, from the tip of the rostrum to the posterior edge of the ventral plate. Notogastral width refers to the maximum width of the notogaster (behind 1022

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pteromorphs in Scheloribates). Lengths of body setae were measured in lateral aspect. All body measurements are presented in micrometers. Formulas for leg setation are given in parentheses according to the sequence trochanter–femur–genu–tibia–tarsus (famulus included). Formulas for leg solenidia are given in square brackets according to the sequence genu–tibia–tarsus. Drawings were made with a camera lucida using a Leica transmission light microscope “Leica DM 2500”. Morphological terminology used in this paper follows that of F. Grandjean: see Travé & Vachon (1975) for references, Norton (1977) for leg setal nomenclature, and Norton & Behan–Pelletier (2009), for overview. The following abbreviations are used on the figures: lam—lamella; tlam—translamella; slam— sublamella; plam—prolamella; Al—sublamellar porose area; tu—tutorium; kf—lateral keel-shaped ridges; ro, le, in, bs, ex—rostral, lamellar, interlamellar, bothridial and exobothridial setae, respectively; Ad—sejugal porose area; hpr—humeral process; hr—humeral ridge; D— dorsophragma; c, la, lm, lp, h, p—notogastral setae; Sa, S1, S2, S3—notogastral sacculi; Aa, A1, A2, A3, A4—notogastral porose areas; ia, im, ip, ih, ips—notogastral lyrifissures; gla—opisthonotal gland opening; f—notogastral furrow; str—strium; cs—circumgastric scissure; csb—circumgastric sigillar band; h, m, a—subcapitular setae; or—adoral seta; v, l, d, cm, acm, ul, sul, vt, lt—palp setae; cha, chb—cheliceral setae; Tg—Trägårdh’s organ; Pd I, Pd II—pedotecta I, II, respectively; 1a, 1b, 1c, 2a, 3a, 3b, 3c, 4a, 4b, 4c—epimeral setae; dis—discidium; cp—circumpedal carina; g, ag, an, ad—genital, aggenital, anal and adanal setae, respectively; iad—adanal lyrifissure; Amar—marginal porose area; p.o.—preanal organ; p.a.—leg porose area; ω, σ, φ—solenidia; ɛ—leg famulus; v, ev, bv, l, d, ft, tc, it, p, u, a, s, pv, pl—leg setae.

Descriptions Scheloribates (Scheloribates) natmaensis sp. nov. (Figs 1–12) Diagnosis Body size: 879–962 × 597–664. Posterior part of notogaster longitudinally striate. Rostrum narrowly triangular, pointed or slightly blunted. Prolamellae complete. Translamellar line represented by two rudimentary lines. Rostral, lamellar and interlamellar setae long, setiform, barbed. Bothridial setae of medium size, clavate, barbed, erect. Notogastral setae setiform, smooth, p2 and p3 longer than others. Notogastral sacculi with long openings. Subcapitular setae setiform, h and m slightly barbed, a with short cilia mediodistally; h longest and thickest. Epimeral and anogenital setae short, setiform, hardly barbed. Circumpedal carinae very short. Marginal porose area band-like. Leg claws serrate on dorsal side, lateral claws each with small tooth ventrodistally. Leg tarsi I with 19 setae (l” absent). Description Measurements. Body length: 946 (holotype: male), 879–962 (four paratypes: two females and two males); notogaster width: 664 (holotype), 597–630 (four paratypes). No clear differences between females and males in body size. Integument (Figs 1, 3–6). Body color light brown to brown. Body surface microporose (visible under high magnification, × 1000). Lateral parts of prodorsum between sublamellae and acetabula I, II microgranulate. Posterior part of notogaster with several longitudinal stria. Subcapitular genae foveolate, diameter of foveolae up to 2. Prodorsum (Figs 1, 3). Rostrum narrowly triangular, pointed or slightly blunted. Lamellae located dorso-laterally, as long as half of prodorsum (measured in lateral view). Prolamellae 2017

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complete, lineate, curving backwards distally. Sublamellae thin, little shorter than lamellae. Translamellar line represented by two very small, poorly visible, rudimentary lines near lamellae. Sublamellar porose areas rounded (12–20) or oval (12–20 × 10–16). Rostral (127–131), lamellar (184–192) and interlamellar (233–258) setae setiform, barbed, ro slightly thinner than others. Bothridial setae (82–94) clavate, barbed, erect, with longer stalks and shorter heads. Anterocentral transverse ridge (between rostral and lamellar setae) not visible. Lateral keel-shaped ridges well developed. Exobothridial setae (90–102) setiform, thin, barbed. Sejugal porose areas elongate oval, transversely oriented. Dorsophragmata short.

FIGURE 1. Scheloribates natmaensis sp. nov., adult: dorsal view. Scale bar 100 μm.

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FIGURE 2. Scheloribates natmaensis sp. nov., adult: ventral view (gnathosoma and legs not shown). Scale bar 100 μm.

Notogaster (Figs 1–5). Anterior notogastral margin slightly convex medially. Anterior part of notogaster with several indistinct longitudinal furrows. Ten pairs of notogastral setae present, all setiform, thin, smooth, p2 and p3 (28–32) longer than others (16–20). Four pairs of sacculi with long openings and elongated channels, distance S1–S1 shorter than S2–S2. Setae lm inserted posteromedial to Sa, lp medial to S1. All lyrifissures distinct, im located anterolateral to S1 and distanced from them, ip posterior and close to S3, ih anterolateral to p3 and distanced from them, ips posterolateral and close to p3. Opisthonotal gland openings located posterior to im. Circumgastric scissure and circumgastric sigillar band distinct.

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FIGURES 3–5. Scheloribates natmaensis sp. nov., adult: 3—anterior part of body, lateral view (gnathosoma and legs not shown); 4—posterior part of body, lateral view; 5—posterior view. Scale bar 100 μm.

Gnathosoma (Figs 6–8). Subcapitulum longer than wide (176–180 × 131–135), with one pair of lateral tubercles. Subcapitular setae setiform, h (53–57) and m (36–41) slightly barbed, a (36–41) with short cilia mediodistally; h thickest. Two pairs of adoral setae (16–20) setiform, heavily barbed. Palps (length 98–106) with setation 0-2-1-3-9(+ω). Postpalpal setae (8) spiniform, smooth. Chelicerae (length 180–188) with two setiform, barbed setae, cha (53–61) longer than chb (36–41). Trägårdh’s organ elongate triangular. 1026

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FIGURES 6–10. Scheloribates natmaensis sp. nov., adult: 6—subcapitulum, ventral view; 7—palp, right, antiaxial view; 8—chelicera, left, paraxial view; 9—trochanter, femur and genu of leg II, right, antiaxial view; 10—trochanter, femur and genu of leg III, left, antiaxial view. Scale bar 50 μm.

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FIGURES 11–12. Scheloribates natmaensis sp. nov., adult: 11—leg I, right, antiaxial view; 12—leg IV, left, antiaxial view. Scale bar 50 μm.

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Epimeral and lateral podosomal regions (Figs 2, 3). Epimeral setal formula: 3-1-3-3. Setae setiform, thin, hardly barbed, 1b (65–69) longer than 1a, 2a, 3a (28–32) and other setae (45–53). Setae 1c inserted ventrolaterally on pedotecta I. Pedotecta I and II represented by small laminae, Pd II quadrangular in ventral view. Discidia well-developed, elongate triangular, rounded distally. Circumpedal carinae very short, directed to region of the acetabula IV. Anogenital region (Figs 2, 4, 5). Four pairs of genital (g1, 32–36; g2–g4, 28), one pair of aggenital (28–32), two pairs of anal (24–28) and three pairs of adanal (28–32) setae setiform, thin, hardly barbed. Adanal lyrifissures located close and parallel to anal plates. Adanal setae ad1 in postanal, ad2 in posterolateral, ad3 in preanal positions. Marginal porose area present, thin, band-like. Ovipositor elongated (298 × 61), blades (114) shorter than length of distal section (beyond middle fold; 184). Each of the three blades with four smooth setae, ψ1 ≈ τ1 (61–65) setiform, longer than thorn-like ψ2 ≈ τa ≈ τb ≈ τc (24–28). Six coronal setae spiniform (4). Legs (Figs 9–12). Median claw thicker than laterals, all serrate on dorsal side; lateral claws each with small tooth ventrodistally. Tibiae I, II with one triangular tubercle ventrobasally. Dorsoparaxial porose areas on femora I–IV and trochanters III, IV and ventral porose areas in basal parts of tarsi and distal parts of tibiae well visible. Formulas of leg setation and solenidia: I (1-5-3-4-19) [1-2-2], II (1-5-2-4-15) [1-1-2], III (2-3-1-3-15) [1-1-0], IV (1-2-2-3-12) [0-1-0]; homology of setae and solenidia indicated in Table 1. Solenidia ω1 on tarsi I, ω1 and ω2 on tarsi II and σ on genua III thickened, blunt-ended, other solenidia longer, setiform. Famuli short, straight, slightly dilated and truncated distally, inserted posterior to solenidia ω2. Material examined Holotype (male) and four paratypes (two females and two males): Myanmar (Burma), Southern Chin State, road to South of Nat Ma Taung National Park, 21°10'7.5''N, 93°54'53.5''E, 2543 m a.s.l., pristine primary forest with rich understory, leaf litter, Winkler extraction, 16.V.2014 (collected by Peter Jäger). TABLE 1. Leg setation and solenidia of adult Scheloribates natmaensis sp. nov. and Zetorchella taungensis sp. nov. Leg

Tr

Fe

Ge

Ti

Ta

I

v’

d, (l), bv”, v”

(l), v’, σ

(l), (v), φ1, φ2

(ft), (tc), (it), (p), (u), (a), s, (pv), v’, (pl), l”*, ɛ, ω1, ω2

II

v’

d, (l), bv”, v”

(l), σ

(l), (v), φ

(ft), (tc), (it), (p), (u), (a), s, (pv), ω1, ω2

III

l’, v’

d, l’, ev’

l’, σ

l’, (v), φ

(ft), (tc), (it), (p), (u), (a), s, (pv)

IV

v’

d, ev’

d, l’

l’, (v), φ

ft”, (tc), (p), (u), (a), s, (pv)

Roman letters refer to normal setae, Greek letters to solenidia (except ɛ = famulus). Single prime (’) marks setae on the anterior and double prime (”) setae on the posterior side of a given leg segment. Parentheses refer to a pair of setae. Tr—trochanter, Fe—femur, Ge—genu, Ti—Tibia, Ta—tarsus. *—seta l” absent on tarsi I in S. natmaensis.

Type deposition The holotype and one paratype are deposited in the collection of the Senckenberg Museum of Natural History, Görlitz, Germany. Three paratypes are deposited in the collection of the Tyumen State University Museum of Zoology, Tyumen, Russia. Etymology The specific name natmaensis refers to the [Nat Ma] Taung National Park, Myanmar, where the new species was collected. 2017

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Remarks Scheloribates natmaensis sp. nov. can be ascribed to the striolatus-group (Ermilov & Starý 2017). Representatives of this group have an entire or partially striate notogaster. The new species is morphologically most similar to S. yamaeensis Nakamura, Hashimoto, Nishi, Nakamura & Fujikawa, 2015 from Japan in having a striate notogaster in the posterior part only, but differs by the larger body size (879–962 × 597–664 vs. 457–564 × 293–400), narrowly triangular rostrum (vs. broadly rounded), clavate, rounded distally bothridial setae (vs. fusiform, pointed distally) and longitudinal stria on notogaster (vs. stria oriented transversely).

Zetorchella taungensis sp. nov. (Figs 13–25) Diagnosis Body size: 830–913 × 630–697. Rostral, lamellar and interlamellar setae long thickened, barbed. Bothridial setae of medium size, clavate, barbed. Notogastral setae bacilliform, barbed, p2, p3 of medium size, shortest, other setae long, with c and p1 slightly shorter. Subcapitular setae setiform, h and m barbed, a with short cilia mediodistally; m longest and thickest. Epimeral and anogenital setae setiform, slightly barbed. Three pairs of minute marginal porose areas present. Leg claws serrate on dorsal side, lateral claws each with small tooth ventrodistally. Description Measurements. Body length: 846 (holotype: male), 830–913 (four paratypes: three females and one male); notogaster width: 630 (holotype), 630–697 (four paratypes). No clear differences between females and male in body size. Integument (Figs 13–18). Body color brown to dark brown. Body surface densely microgranulate, granules larger (up to 1) in lateral parts of the body and posterior part of the ventral plate. Anterior part of prodorsum reticulate. Notogaster, ventral plate, anal plates, subcapitular mentum foveolate (diameter of foveoles up to 12). Subcapitular genae and interlamellar region microfoveolate. Prodorsum (Figs 13, 15). Rostrum rounded. Lamellae located dorso-laterally, shorter than half of prodorsum (measured in lateral view). Sublamellae thin, distinctly shorter than lamellae. Translamella complete, straight, thin. Sublamellar porose areas rounded (6). Tutoria as long as lamellae. Rostral (160–176), lamellar (205–225) and interlamellar (232–237) setae thickened, barbed, ro slightly thinner than others. Bothridial setae (94–114) clavate, barbed, with longer stalks and shorter heads. Exobothridial setae (49–57) setiform, barbed. Sejugal porose areas not visible. Dorsophragmata short, slightly elongated longitudinally. Notogaster (Figs 13–17). Humeral processes quadrangular. Humeral ridges strong. Ten pairs of notogastral setae present, all bacilliform, barbed, c and p1 (143–155) longer than p2, p3 (82–94), other setae longer (180–205). Five pairs of porose areas minute, rounded, Aa (4) larger than others (2). All lyrifissures distinct, im located anterolateral to h3, ip between p1 and p3, ih and ips anterior to p3. Opisthonotal gland openings located posterolateral to h3. Circumgastric scissure distinct, circumgastric sigillar band not visible. Gnathosoma (Figs 18–20). Subcapitulum longer than wide (201–209 × 164–180). Subcapitular setae setiform, h (57–61) and m (73–82) barbed, a (32–36) with short cilia mediodistally; m thickest. Two pairs of adoral setae (20–24) setiform, heavily barbed. Palps (length 118–127) with setation 02-1-3-9(+ω). Postpalpal setae (8) spiniform, hardly barbed. Chelicerae (length 221–229) with two setiform, barbed setae, cha (77–81) longer than chb (53). Trägårdh’s organ tapered. 1030

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FIGURE 13. Zetorchella taungensis sp. nov., adult: dorsal view. Scale bar 100 μm.

Epimeral and lateral podosomal regions (Figs 14, 15). Epimeral setal formula: 3-1-3-3. Setae setiform, slightly barbed, 1a, 2a, 3a (32–41) shorter than other setae (57–69). Setae 1c inserted ventrolaterally on pedotecta I. Pedotecta I and II represented by small laminae, Pd II quadrangular, slightly bifurcate in ventral view. Discidia well-developed, elongate triangular, rounded distally. Circumpedal carinae long, directed to pedotecta II and forming right angles in medioposterior parts. Anogenital region (Figs 14, 16, 17). Six pairs of genital (36–41), one pair of aggenital (36–41), two pairs of anal (36–41) and three pairs of adanal (41–49) setae setiform, slightly barbed. Adanal lyrifissures located close and parallel to anal plates. Adanal setae ad1 in posterolateral, ad2 and ad3 in paraanal positions. Three pairs of minute (2) marginal porose areas present. Ovipositor elongated (343 × 61), blades (118) shorter than length of distal section (beyond middle fold; 225). Each of the three blades with four smooth setae, ψ1 ≈ τ1 (73) setiform, longer than thorn-like ψ2 ≈ τa ≈ τb ≈ τc (16). Six coronal setae spiniform (4). 2017

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FIGURE 14. Zetorchella taungensis sp. nov., adult: ventral view (gnathosoma and legs not shown). Scale bar 100 μm.

Legs (Figs 21–25). Median claw thicker than laterals, all serrate on dorsal side; lateral claws each with small tooth ventrodistally. Dorso-paraxial porose areas on femora I–IV and trochanters III, IV and ventral porose areas in basal parts of tarsi and distal parts of tibiae well visible. Formulas of leg setation and solenidia: I (1-5-3-4-20) [1-2-2], II (1-5-2-4-15) [1-1-2], III (2-3-1-3-15) [1-1-0], IV (1-2-2-3-12) [0-1-0]; homology of setae and solenidia indicated in Table 1. All solenidia setiform. Famuli short, straight, slightly dilated and truncated distally, inserted posterior to solenidia ω2. Material examined Holotype (female) and four paratypes (three females and one male): Myanmar (Burma), Southern Chin State, Nat Ma Taung National Park, road Kampetlet-Mindat, 21°12'33.8''N, 94°01'26.8''E, 2150 m a.s.l., disturbed primary forest, Winkler extraction, 11.V.2014 (collected by Peter Jäger).

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Type deposition The holotype and one paratype are deposited in the collection of the Senckenberg Museum of Natural History, Görlitz, Germany. Three paratypes are deposited in the collection of the Tyumen State University Museum of Zoology, Tyumen, Russia.

FIGURES 15–17. Zetorchella taungensis sp. nov., adult: 15—anterior part of body, lateral view (gnathosoma and legs not shown); 16—posterior part of body, lateral view; 17—posterior view. Scale bar 100 μm.

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FIGURES 18–23. Zetorchella taungensis sp. nov., adult: 18—subcapitulum, ventral view; 19—palp, right, antiaxial view; 20—chelicera, left, paraxial view; 21—posterior part of femur IV, left, antiaxial view; 22— femur (without basal part) and genu of leg II, right, antiaxial view; 23—trochanter, femur and genu of leg III, left, antiaxial view. Scale bars 50 μm (18–20, 22, 23), 25 μm (21). 1034

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FIGURES 24–25. Zetorchella taungensis sp. nov., adult: 24—leg I, left, paraxial view; 25—leg IV, left, antiaxial view. Scale bar 50 μm.

Etymology The specific name taungensis refers to the Nat Ma [Taung] National Park, Myanmar, where the new species was collected. Remarks Zetorchella taungensis sp. nov. is morphologically most similar to Z. nortoni Ermilov, Sidorchuk & Rybalov, 2010 from Ethiopia in having well-developed bothridial setae with long stalks and bacilliform notogastral setae of medium size, but differs by the larger body size (830–913 × 630–697 2017

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vs. 713–813 × 514–564), longer notogastral setae, lm and lp reaching insertions of lp and h1, respectively (vs. shorter, not reaching), bothridial setae shorter than humeral notogastral setae c (vs. longer) and notogastral setae c, p1 similar in length (vs. c distinctly longer). Record Zetorchella orbiculata (Wen & Zhao, 1994). Distribution: southern China. Material (two specimens) examined: Myanmar (Burma), Southern Chin State, Nat Ma Taung National Park, road KampetletMindat, 21°12'33.8''N, 94°01'26.8''E, 2150 m a.s.l., disturbed primary forest, Winkler extraction, 11.V.2014 (collected by Peter Jäger).

Acknowledgements I cordially thank Dr. Elizabeth A. Hugo-Coetzee (National Museum, Bloemfontein, South Africa) and two anonymous reviewers for their valuable comments; Dr. Peter Jäger (Senckenberg Research Institute, Frankfurt am Main, Germany), who collected materials from Myanmar and transferred it from the collection of the Senckenberg Research Institute for our study, and Prof. Dr. Jochen Martens (Johannes Gutenberg Universität, Mainz, Germany) for support in the field.

References Berlese, A. (1908) Elenco di generi e specie nuovi di Acari. Redia, 5, 1–15. Berlese, A. (1916) Centuria prima di Acari nuovi. Redia, 12, 19–67. Ermilov, S.G. (2017) New faunistic and taxonomical findings of oribatid mites (Acari, Oribatida) of the family Otocepheidae from Myanmar. Systematic and Applied Acarology, 22(7), 948–961. http://doi.org/10.11158/saa.22.7.4 Ermilov, S.G. & Anichkin, A.E. (2014) A new species of Scheloribates (Scheloribates) from Vietnam, with notes on taxonomic status of some taxa in Scheloribatidae (Acari, Oribatida). International Journal of Acarology, 40(1), 109–116. https://doi.org/10.1080/01647954.2014.885564 Ermilov, S.G., Sidorchuk, E.A. & Rybalov, L.B. (2010) Zetorchella nortoni, a new species of oribatid mite from Ethiopia (Acari: Oribatida: Caloppiidae). Acarina, 18(1), 61–65. Ermilov, S.G. & Starý, J. (2017) A new species of Scheloribates (Acari, Oribatida, Scheloribatidae) from Vietnam, with key to the striolatus-group. Ecologica Montenegrina, 10, 14–21. Nakamura, Y.-N., Hashimoto, S., Nishi, Y., Nakamura, Y. & Fujikawa, T. (2015) Two new species of Eremellidae and Scheloribatidae (Acari, Oribatida) from the Kuma district of southern Japan. Acarologia, 55(2), 171–187. https://doi.org/10.1051/acarologia/20152159 Norton, R.A. (1977) A review of F. Grandjean's system of leg chaetotaxy in the Oribatei (Acari) and its application to the family Damaeidae. In: Dindal, D.L. (eds.), Biology of Oribatid Mites. Syracuse, SUNY College of Environmental Science and Forestry, pp. 33–61. Norton, R.A. & Behan-Pelletier, V.M. (2009) Suborder Oribatida. Chapter 15. In: Krantz, G.W. & Walter, D.E. (eds.), A Manual of Acarology. Texas Tech University Press, Lubbock, pp. 430–564. Subías, L.S. (2004) Listado sistemático, sinonímico y biogeográfico de los ácaros oribátidos (Acariformes: Oribatida) del mundo (excepto fósiles). Graellsia, 60 (número extraordinario), 3–305. Online version accessed in February 2016, 593 pp, in February 2017, 598 pp. http://escalera.bio.ucm.es/usuarios/bba/cont/docs/RO_1.pdf Travé, J. & Vachon, M. (1975) François Grandjean. 1882–1975 (Notice biographique et bibliographique). Acarologia, 17(1), 1–19. Wen, Z. & Zhao, X. (1994) The first investigation on the oribatid mites of the Yunnan province, China (Acari: Oribatida). Acta Arachnologica Sinica, 3(1), 71–80. Submitted: 20 May 2017; accepted by Lizel Hugo-Coetzee: 19 Jun. 2017; published: 6 Jul. 2017 1036

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