Ermilov, Sergey G. ¹ and Frolov, Andrey V. ²

¹✉ Institute of Environmental and Agricultural Biology (X-BIO), Tyumen State University, Tyumen, Russia.
²Zoological Institute of Russian Academy of Sciences, Saint Petersburg, Russia.

2022 - Volume: 62 Issue: 3 pages: 798-810

ZooBank LSID: 7038FCA4-E123-4EA6-9622-0EE84CCC9DFA

Original research

Keywords

Abstract

Introduction

The oribatid mite (Acari, Oribatida) fauna of Madagascar is insufficiently studied (e.g. Balogh 1962; Mahunka 1993, 2010; Ermilov and Starý 2020). Our work is based on materials which were collected by swiping bushes during a field trip in central Madagascar in 2022. The collected specimens belong to four species (it is theoretically possible that some species are arboreal) of oribatid mites; of these, two species from the genera Afreremella Mahunka, 1973 (Eremellidae) and Lucoppia Berlese, 1908 (Oribatulidae) are new to science. The main goal of this paper is to describe these two new species and a new subgenus within Afreremella.

The oribatid mite genus Afreremella was proposed by Mahunka (1973) with Afreremella luisiae Mahunka, 1973 as type species. At present, the genus comprises only one (type) species, which is known from Zimbabwe; hence, Afreremella is recorded in Madagascar for the first time. The supraspecies system of Eremellidae and the support for independence of Afreremella were discussed by Ermilov and Frolov (2019), however, Subías (2004, online version 2022) synonymized this genus with Eremella (Licnocepheus) Woolley, 1969 without explanations.

The oribatid mite genus Lucoppia was proposed by Berlese (1908) with Notaspis burrowsi Michael, 1890 (= Zetes lucorum Koch sensu Berlese, 1892) as type species. At present, the genus comprises two species, which are distributed collectively in the Holarctic region, Chile and tropics (except Africa); hence, Lucoppia is recorded in the Afrotropical region for the first time. Subías (2004, online version 2022) included also Lucoppia nicora Dzhaparidze, 1986 (see Dzhaparidze and Gomelauri 1986) in Lucoppia, however, we previously support the placement of this species within Zygoribatula Berlese, 1916 according to opinion of Murvanidze et al. (2018).

The additional goals of this paper are: to present the revised generic diagnoses for Afreremella and Lucoppia; to provide an identification key to known representatives of Lucoppia; and to discuss the taxonomic status of the genus Archeremella Balogh and Mahunka, 1974.

Material and methods

Material — All mites were collected in one locality by swiping bushes at 1–2 m height with a beat entomological net and extracted from the net with an entomological aspirator and stored in 96% ethanol.

Observation and documentation — Specimens were mounted in lactic acid on temporary cavity slides for measurement and illustration. Body length was measured in lateral view, from the tip of the rostrum to the posterior edge of the notogaster. Notogastral width refers to the maximum width of the notogaster in dorsal view. 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''.

Terminology — Morphological terminology used in this paper follows that of Grandjean: see Travé and Vachon (1975) for references; Norton (1977) for leg setal nomenclature; and Norton and Behan-Pelletier (2009) for overview.

Abbreviations — Prodorsum: lam = lamella; cos = costula; slam = sublamella; tlam = translamella; tcos = transcostula; tu = tutorium; ro, le, in, ex, bs = rostral, lamellar, interlamellar, exobothridial, and bothridial seta, respectively; Ad = dorsosejugal porose area; D = dorsophragma; P = pleurophragma. Notogaster: c, da, la, dm, lm, dp, lp, h, p = setae; Aa, A1, A2, A3 = porose areas; ia, im, ip, ih, ips = lyrifissures; gla = opisthonotal gland opening. Gnathosoma: a, m, h = subcapitular setae; or = adoral seta; d, l, cm, acm, ul, su, lt, vt, inf, sup = palp setae; ω = palp solenidion; cha, chb = cheliceral setae; Tg = Trägårdh's organ. Epimeral and lateral podosomal regions: 1a, 1b, 1c, 2a, 3a, 3b, 4a, 4b, 4c = epimeral setae; z = aperture of supracoxal gland; Am = humeral porose area; PdI, PdII = pedotectum I and II, respectively; dis = discidium; cir = circumpedal carina. Anogenital region: g, ag, an, ad = genital, aggenital, anal, and adanal seta, respectively; iad = adanal lyrifissure; Ap = postanal porose area; p.o. = preanal organ. Legs: Tr, Fe, Ge, Ti, Ta = trochanter, femur, genu, tibia, and tarsus, respectively; p.a. = porose area; ɛ = famulus; d, l, v, ev, bv, ft, tc, it, p, u, a, s, pv, pl = setae; ω, σ, φ = solenidia.

Taxonomy

Family Eremellidae

Genus Afreremella Mahunka, 1973

Type species: Afreremella luisiae Mahunka, 1973

Generic traits (adult) of Afreremella

With character states of Eremellidae (Balogh 1961; Norton and Behan-Pelletier 2009). Measurements: Medium-sized (length about 500). Integument: Dorsal and ventral sides areolate. Prodorsum: Rostrum rounded. Costulae and transcostula of medium length, simple, forming trapezoid structure. Tutorium well developed, illusory forming triangular projection in dorsal aspect. Rostral, lamellar and interlamellar setae medium-sized to long; ro setiform, le and in fusiform or thick; ro inserted dorsolaterally on prodorsum, le on transcostula, in in interbothridial region. Exobothridial seta present, short. Bothridial seta with short stalk and globular or spoon-shaped head. Bothridium cup-shaped. Dorsophragma and pleurophragma present. Notogaster: With 10 or 12–13 (if dorsocentral setae da, dm/dp developed) pairs of notogastral setae; posterior seta p₁ short, broadly phylliform; p₂ and p₃ shortest, dilated distally; others medium-sized or long, fusiform or thick; humeral region with one seta (c₂). Gnathosoma. Subcapitulum diarthric. Palp setation: 0–2–1–3–9(+ω); solenidion bacilliform, attached to eupathidium. Axillary saccule absent. Chelicera chelate-dentate. Epimeral and lateral podosomal regions: Epimeral setal formula: 3–1–3–3. Pedotectum I represented by medium-sized lamina, pedotectum II represented by small lamina. Discidium developed. Anogenital region: Six pairs of genital, one pair of aggenital, two pairs of anal, and three pairs of adanal setae; ad₁ and ad₂ dilated distally, others setiform; an₁ located close to inner margin of anal plate. Legs: All legs homotridactylous. Tibia I with dorsodistal apophysis bearing solenidion φ₁. Solenidia ω₁ of tarsus I and ω of tarsus II inserted on tubercles; tarsus II with one solenidion. Porose area present on tarsi I–IV, femora I–IV, trochanters III, IV.

Subgeneric traits (adult) of Afreremella (Afreremella) Mahunka, 1973

Notogaster with 10 pairs of notogastral setae; dorsocentral part of notogaster without setae (da, dm, dp absent).

Subgeneric traits (adult) of Afreremella (Arboreremella) n. subgen

Type species: Afreremella (Arboreremella) madagascarensis n. sp.

Notogaster with 12–13 pairs of notogastral setae; dorsocentral part of notogaster with setae (da, dm and usually also dp present).

Etymology

The name Arboreremella refers to the arboreal place of origin +'eremella', a common suffix for generic names in Eremellidae.

Afreremella (Arboreremella) madagascarensis n. sp.

ZOOBANK: 28C0F62D-8A0C-4FFD-AA96-45A97703D478

(Figures 1–3)

Diagnosis — Body length: 547–564. Rostral seta setiform, barbed; lamellar and interlamellar setae thick, heavily barbed; bothridial seta globular, roughened; in longest. Notogastral setae (except posterior setae) long, thick, heavily barbed. Epimeral and anogenital setae (except dilated distally, heavily barbed ad₁, ad₂) setiform, slightly barbed. Adanal lyrifissure diagonal.

Description of adult — Measurements – Body length: 547 (holotype, female), 547–564 (three paratype females); body width: 348 (holotype), 348–365 (three paratype females).

Integument – Body light brown to brown. Surface and setae partially covered by thin layer of gel-like cerotegument. Prodorsum, notogaster, subcapitular mentum, epimeral and anogenital regions, genital and anal plates areolate (diameter of areola up to 12); areolae on genital plate indistinct. Lateral part of body partially microgranulate. Antiaxial side of tarsi I–IV and tibiae I, II partially with transverse rugosities; antiaxial side of leg femora I–IV, tibiae I, II and trochanters III, IV with rounded and elongated tubercles.

Prodorsum – Rostrum broadly rounded. Costula about 1/2 length of prodorsum, little not reaching transcostula. Transcostula long, laterally almost reaching tutorium. Tutorium bent mediodistally. Rostral seta (65–73) setiform, barbed, inserted on tubercle; lamellar (69–82) and interlamellar (102–114) setae thick, heavily barbed, inserted on tubercles; exobothridial seta (12–14) setiform, thin, slightly barbed; bothridial seta (32–36) globular, roughened.

Notogaster – Posterior notogastral margin slightly undulate. Thirteen (in holotype and in two paratypes) or twelve (in one paratype; dp absent) pairs of notogastral setae; p₁ (24–28) broadly phylliform, heavily barbed; p₂ and p₃ (14–16) dilated distally, heavily barbed; others (77–86) thick, heavily barbed; all setae inserted on tubercles. Opisthonotal gland opening and all lyrifissures (except ia not observed) well visible.

Gnathosoma – Subcapitulum size: 123–131 × 98–106. Subcapitular (a, h: 20–22; m: 28–32) and adoral (10) setae setiform, thin, slightly barbed; m slightly thicker than others. Palp (length: 73–82) with typical setation: 0–2–1–3–9(+ω). Postpalpal seta (4) spiniform, smooth. Chelicera (length: 135–139) with two setiform, barbed setae (cha: 41–45; chb: 24–28).

Epimeral and lateral podosomal regions – Epimeral setae (1b: 24; 3b: 20; others: 12) setiform, thin, slightly barbed. Discidium triangular, tubercle-like distally.

Anogenital region – Genital (g₁, g₂: 10–12; g₃–g₆: 12–14), aggenital (12–14) and anal (12–14) setae, and adanal seta ad₃ (12–14) setiform, thin, slightly barbed. Adanal setae ad₁ (12–14) and ad₂ (8–12) dilated distally, heavily barbed, inserted on tubercles. Adanal lyrifissure diagonal.

Legs – Median and lateral claws strong, slightly barbed on dorsal side. Proximoventral porose area on tarsi I–IV slightly visible; distoventral porose area on tibiae I–IV not observed; dorsoparaxial porose area on femora I–IV and on trochanters III, IV distinct. Formulas of leg setation and solenidia: I (1–4–3–4–20) [1–2–2], II (1–4–3–4–15) [1–1–1], III (2–2–1–3–15) [1–1–0], IV (1–2–2–3–12) [0–1–0]; homology of setae and solenidia indicated in Table 1. Solenidia φ₁ of tibia I and φ of tibia II very long, flagellate; ω₁ of tarsus I long, rod-like, coupled with seta ft″; ω₂ of tarsus I, φ₂ of tibia I and ω of tarsus II medium-sized, setiform; φ of tibiae III, IV and σ of genua I–III short, bacilliform, slightly bent mediodistally. Famulus short, erect, slightly swollen distally, inserted between solenidia ω₁ and ω₂. Some setae thick, heavily barbed.

Material examined — Holotype (female) and three paratypes (all females): Madagascar, Ankaratra Mt., 19°20′32″S, 47°18′03″E, 1980 m a.s.l., from bushes (swiping), 24.02.2022 (leg. A.V. Frolov).

Type deposition — The holotype is deposited in the collection of the Zoological Institute, Russian Academy of Sciences, Saint-Petersburg, Russia; three paratypes are deposited in the collection of the Tyumen State University Museum of Zoology, Tyumen, Russia. All specimens are preserved in 70% solution of ethanol with a drop of glycerol.

Etymology — The name madagascarensis refers to the country of origin, Madagascar.

Family Oribatulidae

Genus Lucoppia Berlese, 1908

Type species: Notaspis burrowsi Michael, 1890

Generic traits (adult) of Lucoppia

With character states of Oribatulidae (Balogh and Balogh 1984; Norton and Behan-Pelletier 2009). Measurements: Medium-sized to large (length about 450–750). Integument: Notogaster reticulate, foveolate or smooth. Prodorsum: Rostrum rounded or pointed. Lamella short, simple, without cusp, convergent; translamella short, thin; lamellae and translamella forming trapezoid structure. Sublamella and dorsosejugal porose area present. Prolamella and sublamellar porose area absent. Tutorium long, ridge-like. Rostral, lamellar and interlamellar setae long, setiform; ro inserted dorsally on prodorsum, le on the end of lamella, in in interbothridial region; le nearer to in than to ro. Exobothridial seta present, medium-sized. Bothridial seta short to medium-sized, globular or clavate to fusiform. Bothridium cup-shaped. Dorsophragma small. Notogaster: Anterior margin of notogaster distinct, convex medially. Pteromorph and pleural carina completely absent. Octotaxic system with four pairs of porose areas; Aa elongate oval, others rounded to oval. With fourteen pairs of long, stiff, barbed notogastral setae; humeral region with one seta (c₂). Gnathosoma. Subcapitulum diarthric. Palp setation: 0–2–1–3–9(+ω); solenidion bacilliform, attached to eupathidium. Axillary saccule absent. Chelicera chelate-dentate. Epimeral and lateral podosomal regions: Epimeral setal formula: 3–1–3–3. Apodemes II, III and sejugal apodeme well separated medially. Humeral porose area Am present, Ah absent. Pedotectum I represented by medium-sized lamina, pedotectum II represented by small lamina. Discidium and circumpedal carina present. Anogenital region: Four or five pairs of genital, one pair of aggenital, two pairs of anal, and three pairs of adanal setae. Adanal lyrifissure located in preanal position. Postanal porose area band-like. Legs: All legs heterotridactylous. Porose area present on tarsi I–IV, tibiae I–IV, femora I–IV, trochanters III, IV.

Lucoppia ankaratraensis n. sp.

ZOOBANK: AE6B6891-F377-44BB-BA3B-45861BA3DB81

(Figures 4–6)

Diagnosis — Body length: 415–481. Notogaster and anogenital region with small foveolae. Rostrum pointed. Rostral, lamellar and interlamellar setae long, setiform, erect, barbed; ro shortest and thinnest, in longest and thickest; bothridial seta short, globular, roughened. Notogastral setae long, stiff, erect (c, p₁–p₃ shorter than others). Epimeral, genital and aggenital setae short, setiform, thin, slightly barbed; four pairs of genital setae. Anal and adanal setae short, stiff, thin, slightly barbed. Leg tarsus I with 19 setae (l″ absent).

Description of adult — Measurements – Body length: 415 (holotype, male), 415 (one male paratype); 448–481 (six paratype females); body width: 249 (holotype), 249 (one male paratype); 282–298 (six paratype females).

Integument – Body light brown to brown. Surface covered by layer of dense microgranulate cerotegument. Notogaster and anogenital region sparsely foveolate (diameter of foveola up to 4). Region between costulae with indistinct transverse ridges. Antiaxial side of tarsi I–IV and paraxial side of tarsi I–IV, tibiae I–IV and femora I–IV partially with transverse rugosities.

Prodorsum – Rostrum with triangular projection, pointed. Lamella about 1/4 length of prodorsum, connected by thin, straight or slightly convex medially translamella. Tutorium well developed. Rostral seta (73–82) setiform, barbed, inserted on tubercle; lamellar (98–106) and interlamellar (123–131) setae setiform, erect, barbed, inserted on tubercles; in slightly thicker than le, le slightly thicker than ro; exobothridial seta (28–36) setiform, thin, slightly barbed; bothridial seta (28–32) globular, roughened. Dorsosejugal porose area oval (12 × 4), located posterior to in.

Notogaster – Anterior margin distinctly convex medially. Fourteen pairs of stiff, erect notogastral setae (c: 41–49; p₁: 49–65; p₂: 45–49; p₃: 41–45; others: 69–77). Porose area Aa elongate oval (24–26 × 8–10); Aa oval (14–16 × 8–10); A1 and A2 rounded (10–16). Opisthonotal gland opening and all lyrifissures well visible.

Gnathosoma – Subcapitulum size: 123–131 × 98–102. Subcapitular (a: 20; m, h: 28–32) and adoral (10) setae setiform, thin, slightly barbed. Palp (length: 77–86) with typical setation: 0–2–1–3–9(+ω). Postpalpal seta (8) spiniform, roughened. Chelicera (length: 123–131) with two setiform, barbed setae (cha: 36–38; chb: 24–26).

Epimeral and lateral podosomal regions – Epimeral setae (1a, 2a, 3a: 16–20; others: 28–32) setiform, thin, slightly barbed; 1c, 3c and 4c slightly thicker than others. Discidium triangular, tubercle-like distally. Circumpedal carina distinct.

Anogenital region – Four pairs of genital setae. Genital (16–20) and aggenital (28–32) setae setiform, thin, slightly barbed; anal (16–20) and adanal (16–24) setae stiff, thin, slightly barbed. Adanal lyrifissure diagonal. Postanal porose area distinct.

Legs – Median claw clearly thicker than lateral claws; all slightly barbed on dorsal side; lateral claw with tubercle distoventrally. Proximoventral porose area on tarsi I–IV, distoventral porose area on tibiae I–IV and dorsoparaxial porose area on femora I–IV and on trochanters III, IV distinct. Formulas of leg setation and solenidia: I (1–4–3–4–19) [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 2. Solenidia φ₁ of tibia I and φ of tibiae II–IV very long, setiform; φ of tibia IV swollen distally; σ of genu III short, bacilliform, slightly bent mediodistally; others medium-sized, rod-like. Famulus short, erect, slightly swollen distally, inserted between solenidion ω₂ and seta ft″.

Material examined — Holotype (male) and seven paratypes (one male and six females): Madagascar, Ankaratra Mt., 19°20′32″S, 47°18′03″E, 1980 m a.s.l., from bushes (swiping), 24.02.2022 (leg. A.V. Frolov).

Type deposition — The holotype is deposited in the collection of the Zoological Institute, Russian Academy of Sciences, Saint-Petersburg, Russia; seven paratypes are deposited in the collection of the Tyumen State University Museum of Zoology, Tyumen, Russia. All specimens are preserved in 70% solution of ethanol with a drop of glycerol.

Etymology — The name ankaratraensis refers to the place of origin, Ankaratra Mt.

Remarks — Distinctive characters of the new species from the other Lucoppia species can be found in the identification key below.

Key to known species of Lucoppia

1. Rostrum pointed; bothridial seta short, globular; notogaster and anogenital region foveolate; four pairs of genital setae; body length: 415–481
...... Lucoppia ankaratraensis n. sp. Distribution: Madagascar.

— Rostrum rounded; bothridial seta medium-sized, clavate to fusiform; notogaster and anogenital region not foveolate; five pairs of genital setae
...... 2

2. Notogaster with reticulate pattern; body length: 714
...... Lucoppia feideri Subías, 2010 (=Romanobates reticulatus Feider, Vasiliu and Călugăr, 1970). Distribution: eastern Mediterranean, Chile.

— Notogaster without reticulate pattern; body length: 620–750
...... Lucoppia burrowsi (Michael, 1890) (see also e.g. Weigmann 2006; Seniczak and Seniczak 2012). Distribution: Holarctic region, southeastern China, Hawaii, Mexico.

Discussion

Number and localization of notogastral setae have importance for supporting of the supraspecies status in Eremellidae. For example, Balogh and Mahunka (1974) described the eremellid subgenus Eremella (Archeremella) Balogh and Mahunka, 1974 which differs from the nominate subgenus by the presence of 14 pairs (versus 10 pairs) of notogastral setae and the localization of setae da, dm, dp in the dorsocentral part of the notogaster (versus da, dm, dp absent; dorsocentral notogastral part without setae). Later, Pérez-Íñigo (1981) suggested to raise the status of Archeremella to the generic level, which was supported further by other authors (e.g., Fujikawa 1991; Subías 2004; Mahunka 2008; Ermilov and Frolov 2019). Thus, following the analogy, Afreremella (Arboreremella) madagascarensis n. sp. could also theoretically belong to the new genus Arboreremella which differs from the related genus Afreremella by the presence of 12–13 pairs (versus 10 pairs) of notogastral setae including da, dm and usually also dp in dorsocentral notogastral part (versus da, dm, dp absent). However, the type species of Afreremella (A. luisiae) and A. madagascarensis are morphologically very similar in other important characters (e.g., body ornamentation; structure of costular-transcostular complex and tutorium; morphological differentiation of notogastral and adanal setae and their lengths), therefore, we have no reason to allocate a new genus, but, in our opinion, the extra number of notogastral setae and the presence of da, dm, dp in the dorsocentral notogastral part are sufficient to support a new subgenus.

Based on above listed explanations, we believe that the generic status of Archeremella in the current situation is not justified, therefore, we propose to maintain this taxon as subgenus within Eremella, as suggested by Balogh and Mahunka (1974) initially. Hence, we support the initial systematic placement of Eremella (Archeremella) leowae Balogh and Mahunka, 1974 and propose recombinations of two other known species from Archeremella to Eremella (Archeremella): E. (A.) africana (Pérez-Íñigo, 1981) comb. nov.; and E. (A.) bartlae (Mahunka, 2008) comb. nov.

Other records

Family Cymbaeremaeidae

Scapheremaeus anteriorugosus Mahunka, 2011: 3 ex. Distribution: Madagascar.

Family Humerobatidae

Humerobates africanus (Mahunka, 1984): 29 ex. Distribution: Afrotropical region.

Acknowledgements

The authors thank two anonymous reviewers for valuable comments. The study was performed in the framework of the Russian State research project 122031100272-3. Collecting in Madagascar was conducted under a permit No. 443/21/MEDD/SG/DGGE/DAPRNE/SCBE.Re; samples were exported to Russia under a permit No. 036N-EA02/MG21.

References

1. Balogh J. 1961. Identification keys of world oribatid (Acari) families and genera. Acta Zool. Acad. Sci. Hung., 7(3-4): 243-344.
2. Balogh J. 1962. New oribatids from Madagascar (Acari). Ann. Hist. Nat. Mus. Nat. Hung., 54: 419-427.
3. Balogh J., Balogh P. 1984. A review of the Oribatuloidea Thor, 1929 (Acari: Oribatei). Acta Zool. Hung., 30(3-4): 257-313. https://doi.org/10.1016/0305-4179(84)90004-4
4. Balogh J., Mahunka S. 1974. Oribatid species (Acari) from Malaysian soils. Acta Zool. Acad. Sci. Hung., 20(3-4): 243-264.
5. Berlese A. 1892. Acari, Myriapoda et Scorpiones hucusque in Italia reperta. Portici, Padova, Fascicolo LXIV, N. 1-10.
6. Berlese A. 1908. Elenco di generi e specie nuove di Acari. Redia, 5: 1-15.
7. Berlese A. 1916. Centuria terza di Acari nuovi. Redia, 12: 289-338.
8. Dzhaparidze N.I.W., Gomelauri L.A. 1986. Oribatid (Oribatei) and gamasid (Gamasoidea) mites in caves of Georgia Soobshcheniya Acad. Nauk Gruz. SSR, 124(3): 613-616.
9. Ermilov S.G., Frolov A.V. 2019. Contribution to the knowledge of the oribatid mite genus Eremella (Acari, Oribatida, Eremellidae). Acarologia, 59(2): 214-226. https://doi.org/10.24349/acarologia/20194326
10. Ermilov S.G. & Starý J. 2020. New species and records of Oppiidae (Acari, Oribatida) from the Montagne d'Ambre National Park (Madagascar). Syst. Appl. Acarol. 25(4): 697-706. https://doi.org/10.11158/saa.25.4.9
11. Feider Z., Vasiliu N., Calugar M. 1970. Trois especes nouvelles de la famille des Oribatulidae Thor, 1929 (Oribatei). Rev. Roum. Biol. Ser. Zool., 15(5): 293-313.
12. Fujikawa T. 1991. List of oribatid families and genera of the World. Edaphologia, 46: 1-132.
13. Mahunka S. 1973. Neue und interessante Milben aus dem Genfer Museum XI. Neue und wenig bekannte Oribatiden aus Rhodesien (Acari). Arch. Sci., 26(3): 205-225.
14. Mahunka S. 1984. Oribatids of the Eastern Part of the Ethiopian Region (Acari). V. Acta Zool. Hung., 30(1-2): 87-136.
15. Mahunka S. 1993. Oribatids from Madagascar I. (Acari: Oribatida). New and interesting mites from Geneva Museum LXXVI. Rev. suisse Zool., 100(2): 289-315. https://doi.org/10.5962/bhl.part.79862
16. Mahunka S. 2008. A new genus and some other data of oribatids from Thailand (Acari: Oribatida). Acta Zool. Acad. Sci. Hung., 54(2): 125-150.
17. Mahunka S. 2010. New and little known oribatid mites from Madagascar (Acari: Oribatida). I. Op. Zool. Budapest, 41(1): 47-56.
18. Mahunka S. 2011. New and little known oribatid mites from Madagascar (Acari: Oribatida), II. Acta Zool. Acad. Sci. Hung., 57(1): 1-21.
19. Michael A.D. 1890. On a collection of Acarina formed in Algeria. Proc. Sci. Meet. Zool. Soc. London, Part I: 414-425.
20. Murvanidze M., Cobanoglu S., Cilbircioglu C. 2018. New record of Oribatula (Zygoribatula) nicora (Djaparidze, 1986) comb. nov. from Turkey with a redescription of species. Turk. J. Zool., 42: 601-607. https://doi.org/10.3906/zoo-1806-24
21. 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. (Ed.). Biology of oribatid mites. Syracuse: SUNY College of Environmental Science and Forestry. pp. 33-61.
22. Norton R.A., Behan-Pelletier V.M. 2009. Oribatida. In: Krantz G.W., Walter D.E. (Eds). A Manual of Acarology (TX). Lubbock: Texas University Press. Chapter 15. pp. 430-564.
23. Pérez-Íñigo С. 1981. Resultados de la expedición Peris-Alvarez a la isla de Annobón (13) Oribatid mites (2nd series). Eos, LVII: 201-212.
24. Seniczak S., Seniczak A. 2012. Differentiation of external morphology of Oribatulidae (Acari: Oribatida) in light of the ontogeny of three species. Zootaxa, 3184: 1-34. https://doi.org/10.11646/zootaxa.3184.1.1
25. Subías L.S. 2004. Listado sistemático, sinonímico y biogeográfico de los ácaros oribátidos (Acariformes, Oribatida) del mundo (1758-2002). Graellsia, 60 (número extraordinario): 3-305. https://doi.org/10.3989/graellsia.2004.v60.iExtra.218
26. Subías L.S. 2010. Nuevos nombres de oribátidos (Acari: Oribatida). Bol. R. Soc. Esp. Hist. Nat. Sec. Biol., 104: 35-39.
27. Subías L.S. 2022. Listado sistemático, sinonímico y biogeográfico de los Ácaros Oribátidos (Acariformes: Oribatida) del mundo (excepto fósiles), 17ª actualización. 537 pp. Available from: http://bba.bioucm.es/cont/docs/RO_1.pdf (accessed March 2022).
28. Travé J., Vachon M. 1975. François Grandjean. 1882-1975 (Notice biographique et bibliographique). Acarologia, 17(1): 1-19.
29. Weigmann G. 2006. Hornmilben (Oribatida). Die Tierwelt Deutschlands. Teil 76. Keltern: Goecke and Evers. 520 pp.
30. Woolley T.A. 1969. A new and phoretic oribatid mite (Acarina: Cryptostigmata: Licnodamaeidae). Proc. Ent. Soc. Washington, 71(4): 476-481.

instructions