# Contribution to the knowledge of the oribatid mite
genus Discoppia (Acari, Oribatida, Oppiidae)

Sergey G Ermilov, Olga L Makarova

► To cite this version:

Sergey G Ermilov, Olga L Makarova. Contribution to the knowledge of the oribatid mite genus Discoppia (Acari, Oribatida, Oppiidae). Acarologia, 2023, 63 (3), pp.757-769. 10.24349/131f-vr01. hal-04142770

HAL Id: hal-04142770

https://hal.science/hal-04142770

Submitted on 27 Jun 2023

HAL is a multi-disciplinary open access
archive for the deposit and dissemination of sci-
entific research documents, whether they are pub-
lished or not. The documents may come from
teaching and research institutions in France or
abroad, or from public or private research centers.

L'archive ouverte pluridisciplinaire HAL, est
destinée au dépôt et à la diffusion de documents
scientifiques de niveau recherche, publiés ou non,
émanant des établissements d'enseignement et de
recherche français ou étrangers, des laboratoires
publics ou privés.

<figure><img src="image_1.png" /></figure>

Distributed under a Creative Commons Attribution 4.0 International License
<figure><img src="image_2.png" /></figure>

Acarologia
Open Science in Acarology

<figure><img src="image_3.png" /></figure>

Acarologia

A quarterly journal of acarology, since 1959
Publishing on all aspects of the Acari

All information:
http://www1.montpellier.inra.fr/CBGP/acarologia/
acarologia-contact@supagro.fr

<figure><img src="image_4.png" /></figure>

OPEN ACCESS

Acarologia is proudly non-profit,
with no page charges and free open access

Please help us maintain this system by
encouraging your institutes to subscribe to the print version of the journal
and by sending us your high quality research on the Acari.

Subscriptions: Year 2023 (Volume 63): 450 €
http://www1.montpellier.inra.fr/CBGP/acarologia/subscribe.php

Previous volumes (2010-2021): 250 € / year (4 issues)
Acarologia, CBGP, CS 30016, 34988 MONTFERRIER-sur-LEZ Cedex, France
ISSN 0044-586X (print), ISSN 2107-7207 (electronic)

The digitalization of Acarologia papers prior to 2000 was supported by Agropolis Fondation under the reference ID 1500-024 through the « Investissements d'avenir » programme
(Labex Agro: ANR-10-LABX-0001-01)

<figure><img src="image_5.png" /></figure>

<figure><img src="image_6.png" /></figure>

Acarologia is under **free license** and distributed under the terms of the
Creative Commons-BY
Acarologia

Open Science
in Acarology

# Contribution to the knowledge of the oribatid mite genus *Discoppia* (Acari, Oribatida, Oppiidae)

Sergey G. Ermilov IDª, Olga L. Makarova IDᵇ

## Original research

### ABSTRACT

The supplementary description of the oribatid mite species *Discoppia (Cylindroppia cylindrica* (Pérez-Íñigo, 1965) (Oribatida, Oppiidae) is presented, based on materials collected from deep layers of coastal sandy soils in the Republic of Dagestan, eastern Caucasus. The morphology of exobothridial setae, gnathosoma and ovipositor, and the identification of leg setae and solenidia are given for the first time for this species. The revised generic diagnosis, identification key, distribution, and habitats of all known representatives of *Discoppia* Balogh, 1983 are presented. Some taxonomic nuances within *Discoppia* are discussed, resulting in the following proposals: *Oppia bifidus* Bayoumi and Al-Khalifa, 1985 is a junior synonym of *D. (C.) cylindrica*; *Oppia agricola* Fujikawa, 1982 is not synonym of *D. (C.) cylindrica* and transferred to the genus *Microppia* Balogh, 1983; *Oppia tenuis* Hammer, 1958 is not representative of *Discoppia*; *Discoppia (Cylindroppia rostroincisa)* Subías and Rodríguez, 1986 **n. stat.** (instead of the existing subspecies status). Together with *D. (C.) cylindrica*, in the same samples, two rare Oppiidae species were found: *Oxyoppia (Oxyoppiella minuscula)* Hugo-Coetzee, Lotfollahi, Abbasi-Kalo and Movahedzade, 1917 (known only from the open habitats in Iran) is first recorded in Russia; *Graptoppia (Stenoppia boucheri)* Ermilov and Frolov, 2021 (known from passalid beetles collected in Ghana and Gabon, West Africa) is first found in the Palaearctic. Both findings in Dagestan are among the northernmost records of their semicosmopolitan subgenera.

**Keywords** taxonomy; morphology; generic diagnosis; identification key; distribution; habitat; new record; Caucasus; Russia

**Zoobank** http://zoobank.org/6B244B1D-D506-40AB-A69B-AF81458861A7

### Introduction

The oribatid mite genus *Discoppia* (Acari, Oribatida, Oppiidae) was proposed by Balogh (1983), with *Oppia limae* Balogh and Mahunka, 1974 as type species. Subías and Rodríguez (1986) proposed a subgenus *Discoppia* (*Cylindroppia*), with *Oppia minus cylindrica* Pérez-Íñigo, 1965 as type species. Presently, the genus comprises two subgenera with five species (*D. (Discoppia) limae* (Balogh and Mahunka, 1974); *D. (Cylindroppia) casuarina* (Abd-el-Hamid, Hussein, Bayoumi and Al-Assiuty, 1983); *D. (C.) cylindrica* (Pérez-Íñigo, 1965); *D. (C.) pentasetata* Subías and Rodríguez, 1986; and *D. (C.) rostroincisa* Subías and Rodríguez, 1986), which are distributed collectively in tropics and subtropics.

The type species of *Discoppia* (*Cylindroppia*) was described by Pérez-Íñigo (1965). However, its description is brief and contains significant errors, e.g. incorrect arrangement of dorsal notogastral setae (see Fig. 3d in Pérez-Íñigo 1965). Later, this author (Pérez-Íñigo 1971) provided correct figures. Subías and Rodríguez (1986), Arillo and Subías (1996), Ay

Received 23 April 2023

Accepted 16 June 2023

Published 26 June 2023

Corresponding author

Sergey G. Ermilov ID:
ermilovacari@yandex.ru

Academic editor

Pfingstl, Tobias

https://doi.org/10.24349/131f-vr01

ISSN 0044-586X (print)

ISSN 2107-7207 (electronic)

<figure><img src="image_7.png" /></figure>

Ermilov S. G. and Makarova O. L.

Licensed under
Creative Commons CC-BY 4.0

<footnote>ªTyumen State University, Tyumen, Institute of Environmental and Agricultural Biology (X-BIO), Russia.</footnote>

<footnote>ᵇSevertsov Institute of Ecology and Evolution, Russian Academy of Sciences, Moscow, Russia.</footnote>
and Ayyildiz (2019) presented the supplementary descriptions of this species, however, this
description remained incomplete (for example, morphology of exobothridial seta, gnathosoma,
and ovipositor, as well as identification of leg setae and solenidia are missed) and rather briefly
illustrated.

The main goals of the paper are: to present the supplementary description of *D. (C.) cylindrica* on the basis of mass materials from Caucasus; to summarize its main morphological traits, which will ensure the identification of this species in the future; to revise the generic diagnosis, which is short and incomplete so far; to discuss the systematic placement of *Oppia agricola* Fujikawa, 1982 and *O. tenuis* Hammer, 1958 which Subías (2022) included in *Discoppia*, as well as the synonymy of *Oppia bifidus* Bayoumi and Al-Khalifa, 1985 with *D. (C.) casuarina* (according to Subías 2022) and the taxonomic status of *D. (C.) cylindrica rostroincisa* Subías and Rodríguez, 1986; to provide an identification key to known species of *Discoppia*; to compile data concerning distribution and habitats of members of the genus. Additionally, some interesting findings of Oppiidae for Russia and Palaearctic are presented.

**Material and methods**

**Specimens**

Mites were found on the slope of the small sand drift (Fig. 1a) formed on the flat sandy beach of the Caspian Sea at the distance of 300 m from the shore line. The vegetation of small dune (two samples) and adjacent plane sandy surface (three samples) was dominated by the bindweed, *Convolvulus persicus* L. (Fig. 1d). Numerous individuals (231) of *D. (C.) cylindrica* were found only in two samples from the hill slope (Fig. 1b, c).

Samples (each of volume 250 ml) were taken from the depth of 20-30 cm and contained numerous small roots of the bindweed. Mites were extracted from cores in Moscow using laboratory eclectors during 10 days until their complete dryness without additional heating or lighting. The relative humidity of samples amounted 3.81%. The mites were fixed with 85% alcohol.

**Observation and documentation**

For measurement and illustration, specimens were mounted in lactic acid on temporary cavity slides. All measurements are in micrometers. Body length was measured in lateral view, from the tip of the rostrum to the posterior edge of the notogaster; other structures were oriented to avoid parallax errors. Notogastral width refers to the maximum in dorsal aspect. Setal lengths were measured perpendicular to their long axis, accounting for curvature. 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 DM 2500 light microscope.

**Terminology**

Morphological terminology used in this paper follows that of F. Grandjean (see Travé and Vachon 1975) for references, Ermilov *et al.* (2021), Ermilov and Rybalov (2022); Norton (1977) for leg setal nomenclature; Norton and Behan-Pelletier (2009) for overview.

**Abbreviations**

*Prodorsum:* *ro, le, in, bs, ex* = rostral, lamellar, interlamellar, bothridial, and exobothridial seta, respectively; *exv* = vestige of second exobothridial seta; *ms* = muscle sigillum. *Notogaster:* *c, la, lm, lp, h, p* = setae; *ia, im, ip, ih, ips* = lyrifissures; *gla* = opisthonotal gland opening. *Gnathosoma:* *a, m, h* = subcapitular setae; *or* = adoral seta; *sup, inf, d, l, cm, acm, ul, su, vt* = palp setae; *ω* = palp solenidion; *cha, chb* = cheliceral setae; *Tg* = Trägårdh's organ. *Epimeral region:* *1a-1c, 2a, 3a-3c, 4a-4c* = epimeral setae. *Anogenital region:* *g, ag, an, ad* = genital,
<figure><img src="image_8.png" /></figure>

**Figure 1** Locality and habitat of *Discoppia (Cylindroppia) cylindrica* (Pérez-Íñigo, 1965): a – Dagestan, 20 km N from Makhachkala City, vicinity of Shushiya village, sandy sea beach, slope of small dune; b, c – collection places under *Convolvulus persicus*, 4.XI.2022: points 1 and 2 respectively; d – flowering *C. persicus*, Makhachkala, 10.VI.2021.
<figure><img src="image_9.png" /><figcaption>**Figure 2** *Discoppia (Cylindroppia) cylindrica* (Pérez-Íñigo, 1965), adult: a – dorsal view (legs omitted); b – ventral view (gnathosoma and legs omitted). Scale bar 20 µm.</figcaption></figure>
**Description** — *Measurements* — Body length: 176–199 (n=100); body width: 67–90 (n=100).

*Integument* (Fig. 3a) – Body color light yellowish to light brownish. Body surface with dense cuticular microgranules (visible in dissected specimens under high magnification) which well visible between bothridium and acetabula II and III. Dorsal side sometimes covered by sparse cerotegumental microgranules.

*Prodorsum* (Figs 2a, 3a) – Rostrum rounded. Rostral (15), lamellar (9–11) and interlamellar (9–11) setae branched; exobothridial seta (26–30) setiform, barbed, often pressed to prodorsal surface and therefore poorly observed in dorsal aspect; bothridial seta (19–22) with unilaterally dilated, nearly disk-like head bearing about 10–12 long ciliae (Fig. 3c). One pair of muscle sigilla well visible, located close and anteriorly to interlamellar setae. Slight transverse ridge located between insertions of interlamellar setae.

*Notogaster* (Figs 2a, 3a, 3b) – All notogastral setae (13–15) branched; number of ciliae variable, from one to four (Fig. 3d). Lyrifissures *ia*, *im*, *ip*, *ih*, and *ips* comparatively long, well visible.

*Gnathosoma* (Figs 3e–3g) – Subcapitulum size: 45–49 × 35–37; all subcapitular setae (9) setiform, slightly roughened; both adoral setae (2) setiform, smooth. Palp length: 24–26; setation: 0–2–1–3–8(+ω); tarsal seta *l''* absent; solenidion long (2/3 of tarsus length); postpalpal seta (2) spiniform. Chelicera length: 45–49; setae (*cha*: 17–19; *chb*: 9–11) setiform, barbed.

*Epimeral region* (Figs 2b, 3a) – Epimeral border IV straight. Epimeral setae (1b, 3b, 3c, 4c: 9; others: 7) setiform, slightly roughened.

*Anogenital region* (Figs 2b, 3a, 3b, 3h) – Genital (7), aggenital (9–11), anal (9), and adanal (11–13) setae setiform, slightly roughened. Adanal lyrfissure short, distinct. Ovipositor (Fig. 3h) elongated (39–45 × 11–13), blades (13–15) shorter than length of distal section (beyond middle fold; 26–30); setae of all blades (ψ₁, τ₁: 13; ψ₂, τₐ, τₑ: 11) rod-like, smooth; first blade with four setae, second and third blades with three setae only (τᵇ absent); coronal setae absent.

*Legs* (Figs 4a–4d) – Claw of each leg with slight barb on dorsal side. Porose area on all segments not observed. Formulas of leg setation and solenidia: I (1–5–2–4–20) [1–2–2], II (1–5–2–4–16) [1–1–2], III (2–3–1–3–11) [1–1–0], IV (1–2–2–3–10) [0–1–0]; homology of setae and solenidia indicated in Table 1. Setae *p'* and *p''* setiform on tarsus I versus very short, spiniform on tarsi II–IV; famulus short, rod-like, located laterally to solenidion ω₁; seta *s* eupathidial on tarsus I; setae *a'* (*a''*), *it'* and *it''* absent on tarsus III, *a'* and *a''* absent on tarsus IV; many setae on segments branched; solenidia ω₁ and ω₂ on tarsi I and II, φ₂ on tibia I, φ on tibiae II and III, and σ on genua II and III short or medium-sized, slightly or distinctly bacilliform; solenidia φ₁ on tibia I, σ on genu I and φ on tibia IV long, subflagellate.

**Remarks** — Based on our supplementary description and on the available data (e.g., Pérez-Íñigo 1971; Subías and Rodríguez 1986) about adult *D. (C.) cylindrica*, we propose the following diagnostic morphological traits for this species: body length: 176–225; rostrum rounded; rostral, lamellar, interlamellar, and notogastral setae branched; exobothridial seta setiform, barbed; bothridial seta with unilaterally dilated, nearly disk-like head; epimeral border IV straight; epimeral and anogenital setae setiform, slightly roughened; leg setae *a'* (*a''*), *it'* and *it''* absent on tarsus III, *a'* and *a''* absent on tarsus IV; many leg setae branched.

<table><caption>**Table 1** Leg setation and solenidia of adult *Discoppia* (*Cylindroppia*) *cylindrica* (Pérez-Íñigo, 1965).</caption><thead><tr><th>Leg</th><th>Tr</th><th>Fe</th><th>Ge</th><th>Ti</th><th>Ta</th></tr></thead><tbody><tr><td>I</td><td>v'</td><td>d, (l), bv'', v''</td><td>(l), σ</td><td>(l), (v), φ₁, φ₂</td><td>(ft), (tc), (it), (p), (u), (a), s, (pv), (pl), l'', v', ε, ω₁, ω₂</td></tr><tr><td>II</td><td>v'</td><td>d, (l), bv'', v''</td><td>(l), σ</td><td>(l), (v), φ</td><td>(ft), (tc), (it), (p), (u), (a), s, (pv), l'', ω₁, ω₂</td></tr><tr><td>III</td><td>l', v'</td><td>d, l', ev'</td><td>l', σ</td><td>l', (v), φ</td><td>(ft), (tc), (p), (u), s, (pv)</td></tr><tr><td>IV</td><td>v'</td><td>d, ev'</td><td>d, l'</td><td>l', (v), φ</td><td>ft'', (tc), (p), (u), s, (pv)</td></tr></tbody></table>
<figure><img src="image_10.png" /><figcaption>**Figure 3** *Discoppia (Cylindroppia) cylindrica* (Pérez-Íñigo, 1965), adult: a – right lateral view (gnathosoma and legs omitted); b – posterior view; c – variability of bothridial seta; d – variability of notogastral setae; e – subcapitulum, ventral view; f – palp, right, antiaxial view; g – chelicera, right, antiaxial view; h – ovipositor. Scale bar 20 µm (a, b), 10 µm (c, g, h), 5 µm (c, d, f).</figcaption></figure>
<figure><img src="image_11.png" /><figcaption>**Figure 4** *Discoppia (Cylindroppia) cylindrica* (Pérez-Iñigo, 1965), adult: a – leg I, right, antiaxial view; b – leg II, right, antiaxial view; c – leg III, left, antiaxial view; d – leg IV, left, antiaxial view. Scale bar 10 µm.</figcaption></figure>
## Taxonomical proposals

1. *Oppia bifidus* was described by Bayoumi and Al-Khalifa (1985). The distinctive traits of the species are morphologically variable (branches of rostral and notogastral setae; bar-bulation of bothridial seta) or were not correct (presence of costulae), therefore, Subías (2004) rightly synonymized it with *D. (C.) cylindrica cylindrica*. However, later, Subías (2022) changed his opinion and synonymized *O. bifidus* with *D. (C.) casuarina*. *Discoppia* (*C*) *cylindrica* and *D. (C.) casuarina* are morphologically very similar differing only by the morphology of head of the bothridial seta (unilaterally dilated, nearly disk-like versus elongate fusiform). *Oppia bifidus* has unilaterally dilated, nearly disk-like head of bothridial seta (Bayoumi and Al-Khalifa 1985), therefore, we maintain the initial synonymy (*O. bifidus* with *D. (C.) cylindrica*) of Subías in 2004.

2. Subías (2004, 2022) considered the species *Oppia agricola* Fujikawa, 1982 as a junior synonym of *D. (C.) cylindrica*. However, all morphological traits of *O. agricola* clearly correspond to the generic diagnosis of *Microppia* Balogh, 1983 (Balogh 1983; Subías and Shtanchaeva 2023); for example, this species has one pair of small sclerotized tubercles running from dorsosejugal scissure to basal part of prodorsum (versus absent in *Discoppia*) and differing localization of notogastral seta *lm* to *la* (medially versus mostly posteriorly in *Discoppia*). Therefore, we: 1) do not support the synonymy proposed by Subías (2004), 2) propose a transfer of *O. agricola* in *Microppia: Microppia agricola* (Fujikawa, 1982) **n. comb.**

3. Subías (2004, 2022) included the species *Oppia tenuis* Hammer, 1958 in *D. (Cylindropia)*. However, some important morphological traits of this species clearly do not correspond to the generic diagnosis of *Discoppia*; for example, *O. tenius* has differing localization of notogastral seta *lm* to *la* (medially versus mostly posteriorly in *Discoppia*) and seta *c* (distanced from humeral region versus in humeral region). Therefore, we do not support the placement of *O. tenuis* in *Discoppia*. The problem is that *O. tenuis* was described briefly by Hammer (1958), and its ventral side, containing important diagnostic characters (e.g., epimeral formula; number of genital setae; localization of adanal setae and adanal lyrifissure), is completely unknown, therefore, an objectively correct placement of *O. tenuis* in any known oppiid genus seems almost impossible at the moment.

4. *Discoppia* (*Cylindroppia*) *cylindrica rostroincisa* was described by Subías and Rodríguez (1986). This subspecies differs from *D. (C.) cylindrica cylindrica* only by one morphological trait (rostrum incised medially versus rounded). The morphology of the rostrum is not variable in specimens of the same oppiid species and is widely used in taxonomy of the family at the species and generic levels. Therefore, in our opinion, this difference is sufficient to support the insularity of *Discoppia rostroincisa* on species level, and we propose a species status (instead of the existing subspecies status) for this species: *Discoppia* (*Cylindroppia*) *rostroincisa* Subías and Rodríguez, 1986 **n. stat.**

## Key to known species of *Discoppia*

1. Anterior notogastral margin straight; one pair of interbothridial tubercles present; body length: 228

... Subgenus *Discoppia* (*Discoppia*); single representative: *Discoppia* (*Discoppia*) *limae* (Balogh and Mahunka, 1974)

— Anterior notogastral margin medially clearly convex; interbothridial tubercles absent

2. Subgenus *Discoppia* (*Cylindroppia*)

2. All notogastral setae simple; five pairs of genital setae present; body length: 230

... *Discoppia* (*Cylindroppia*) *pentasetata* Subías and Rodríguez, 1986 (see also Hammer 1979)

— All notogastral setae branched; four pairs of genital setae present

3
species from which only solitary *Micropilia minus* (Paoli, 1908) is the semicosmopolitan polyzonal species. All other mites are either south European/Palaearctic species (*Epilohmannia styriaca* Schuster, 1960; *Lalmoppia krivolutskyi* (Poltavskaya, 1994); *Oxyoppia (Oxyoppiella) minuscula* Hugo-Coetzee, Lotfollahi, Abbasi-Kalo and Movahedzade, 1917) or the broader distributed species that in the Palaearctic inhabit only southern areas (*Sellnickochthonius furcatus* (Weis-Fogh, 1948); *D. cylindrica*; *Graptoppia (Stenoppia) boucheri* Ermilov and Frolov, 2021).

The discovery of two recently described oppiid species are of particular interest. *Oxyoppia (Oxyoppiella) minuscula* till now was known only from Iran (Hugo-Coetzee et al. 2017) where this species resides also open biotopes (wheat fields and grassland). Our finding in the seaboard Dagestan is the first record of *O. minuscula* in Russia and the northernmost record of the semicosmopolitan subgenus *O*. (*Oxyoppiella*) Subías and Rodríguez, 1986 (Hugo-Coetzee et al. 2017). *Graptoppia (Stenoppia) boucheri* was known only from passalid beetles *Erionomus planiceps* Eschscholtz collected in Ghana and Gabon, West Africa (Ermilov and Frolov 2021). So, our finding of this species in sandy soil of maritime Dagestan is its first record in the Palaearctic and one of a few northernmost records of the semicosmopolitan subgenus *G*. (*Stenoppia*) Balogh, 1983 (Subías 2022). Distributional range of the mainly tropical beetle family Passalidae in total is far away from Caucasus (Boucher 2005), that indicates the need for the further investigation of habits and habitats of *G. boucheri*.

Thus, the oribatid species composition revealed in the sandy coastal soil of middle Caspian Sea can be determined as thermophilous consisting mainly of small-size species. The size of most species found is smaller than 300 µm (only the length of *E. styriaca* exceeds 475 µm). The specific habitat of deep sandy soil (where the vertical migration of small arthropod is enhanced) may serve as a stable concentration site of the tropical-subtropical oribatid fauna.

Notably, that the most species (five from seven) and individuals (283 from 285) in studied biotope were parthenogenetic (thelytokous), i.e. reproducing without males (so far presumably for *O. minuscula*). This community feature also argues the stable resource ability that was demonstrated recently in the comprehensive analysis of very different oribatids assemblages (Maraun *et al.* 2019). Possibly, this tendency is common for the free-living mites in general (see Manning and Halliday 1994; Walter and Lindquist 1995).

## Acknowledgements

We are thankful to L.S. Subías and U.Ya. Shtanchaeva for consultations and two anonymous reviewers for valuable comments. This research was partially supported by the grant from the Russian Foundation for Basic Research (No. 20-54-56054 Iran_t).

## ORCID

Sergey G. Ermilov https://orcid.org/0000-0002-0913-131X

Olga L. Makarova https://orcid.org/0000-0002-1820-2511

## References

* Abd-el-Hamid M.E., Hussein M.A., Bayoumi B.M, Al-Assiuty A.I. 1983. The distribution of oribatid mites (Acari: Oribatida) in Gharbia governorate, Egypt. Delta J. Sci., 7(2): 734-755.

* Abdurakhmanov G.M., Grikurova A.A., Subías L.S., Shtanchaeva U.Ya., Kuramagomedov B.M. 2013. Species composition and zoogeographical characteristics of the armored mites (Acariformes, Oribatida) of coast and islands of north-western part of the Caspian Sea. Ecology of animals. The South of Russia: Ecol., Develop., 1: 16-21. https://doi.org/10.18470/1992-1098-2013-1-16-21

* Akrami M.A. 2015. An annotated checklist of oribatid mites (Acari: Oribatida) of Iran. Zootaxa, 3963(4): 451-501. https://doi.org/10.11646/zootaxa.3963.4.1

* Akrami M.A., Behmanesh M. 2015. Oribatid mites (Acari: Oribatida) of Shiraz township, Fars province, Iran. Entomofauna, 36: 377-396.
Akrami M.A., Salehi Sarbizhan A.R. 2020. Oribatid mites (Acari: Oribatida) of Jiroft County, Kerman province, Iran: Introduction of fifty-six species, new records for the fauna of the province. Munis Ent. Zool., 15(1): 73-84.

Akrami M.A., Shahedi A. 2020. Oribatid mites (Acari: Oribatida) of Taft county, Yazd province of Iran, with new records. Pers. J. Acarol., 9(2): 141-160. https://doi.org/10.22073/pja.v9i2.58955

Al-Khalifa M.S., Bayoumi B.M. 1983. New records of soil Acarina and Collembola in Riyadh region. J. Coll. Sci. King Saud Univ., 14(1): 57-61.

Arillo A., Subías L.S. 1996. La subfamilia Medioppiinae en el Sur de España. Descripción de *Medioppia pinsapi* sp. nov. (Oppiidae). Estud. Mus. Cien. Nat. Álava (1995-1996), 10-11: 211-221.

Ay Y., Ayyildiz N. 2019. Investigations on oppiid oribatid mites of Amanos Mountains (Turkey). Plant Prot. Bul., 59(3): 15-23. https://doi.org/10.16955/bitkorb.503754

Balogh J. 1983. A partial revision of the Oppiidae Grandjean, 1954 (Acari: Oribatei). Acta Zool. Acad. Sci. Hung., 29(1-3): 1-79.

Balogh J., Mahunka S. 1974. Oribatid species (Acari) from Malaysian soils. Acta Zool. Acad. Sci. Hung., 20(3-4): 243-264.

Bayoumi B.M., Al-Khalifa M.S. 1985. Three new oribatid mites (Acari, Oribatei) from Saudi Arabia. Bul. Soc. Amis Sci. Let. Poznan, Ser. D., Poznan, 25: 119-126.

Boucher S. 2005. Évolution et phylogénie des Coléoptères Passalidae (Scarabaeoidea). Ann. Soc. Ent. Fr., 41: 239-604. https://doi.org/10.1080/00379271.2005.1069744

Ermilov S.G., Corpuz-Raros L.A. 2017. New data on oribatid mites of the family Oppiidae (Acari: Oribatida) from the Philippines. Acta Zool. Bulg., 69(1): 9-16. https://doi.org/10.11646/zootaxa.4171.1.3

Ermilov S.G., Frolov A.V. 2021. New data on oribatid mites (Acari, Oribatida) phoretic on passalid beetles (Coleoptera, Passalidae) from the Afrotropical and Oriental regions, with descriptions of three new species from Congo, Gabon and Ghana. Syst. Appl. Acarol., 26(4): 769-787. https://doi.org/10.11158/saa.26.4.8

Ermilov S.G., Khaustov A.A. 2018. A contribution to the knowledge of oribatid mites (Acari, Oribatida) of Zanzibar. Acarina, 26(2): 151-159. https://doi.org/10.21684/0132-8077-2018-26-2-151-159

Ermilov S.G., Rybalov L.B. 2022. Taxonomic contribution to the knowledge of the oribatid mite genus *Arcoppia* (Acari, Oribatida, Oppiidae). Int. J. Acarol., 48(7): 588-593. https://doi.org/10.1080/01647954.2022.2149854

Ermilov S.G., Anichkin A.E., Wu D. 2012. Two new species of the genus *Papillacarus* (Acari: Oribatida: Lohmanniidae) from caves of Southern Vietnam. Zootaxa, 3593: 75-88. https://doi.org/10.11646/zootaxa.3593.1.4

Ermilov S.G., Hugo-Coetzee E.A., Khaustov A.A. 2021. Contribution to the knowledge of *Geminoppia* (Acari, Oribatida, Oppiidae), with description of a new species from South Africa. Acta Zool. Acad. Sci. Hung., 67(3): 211-222. https://doi.org/10.17109/AZH.67.3.211.2021

Fujikawa T. 1982. Oribatid fauna from nature farm in Nayoro (5). Edaphologia, 27: 1-4.

Hammer M. 1958. Investigations on the oribatid fauna of the Andes Mountains. I. The Argentine and Bolivia. Biol. Skr. Kong. Dansk. Vidensk. Selsk., 10: 1-129.

Hammer M. 1979. Investigations on the oribatid fauna of Java. Biol. Skr. Kong. Dansk. Vidensk. Selsk., 22 (9): 1-78.

Hugo-Coetzee E.A., Lotfollahi P., Abbasi-Kalo A., Movahedzade E. 1917. A new species of *Oxyoppia* (*Oxyoppiella*) (Acari: Oribatida: Oppiidae) from Iran. Turk. J. Zool., 41: 630-637. https://doi.org/10.3906/zoo-1611-32

Ivan O. 2007. Diversity and distribution of the oribatid mites (Acari, Oribatida) in some lawn ecosystems from Moldavian Plain (Romania). Studii si Comunicari, Complexul Muzeal de Stiințele Naturii "Ion Borcea" Bacau, 21: 236-243.

Kulijev K.A. 1962. Fifteen new representatives of oribatid mites (Acariformes, Oribatei) of the genera *Oppia* and *Ceratozetes*. Trudy Azerbaijan NIVI, 13: 250-268.

Mahunka S., Mahunka-Papp L. 1999. Oribatids from the Szarvas Arboretum (SE Hungary) (Acari: Oribatida). Folia Ent. Hung., 60: 83-107. https://doi.org/10.5962/bhl.part.80144

Mahunka S., Murányi D., Kontschán J. 2013. The role of the Balkan Peninsula in the origin and genesis of the soil fauna of the Carpathian Basin: history, aims and results. Opusc. Zool. Budapest, 44 (suppl. 1): 5-10.

Manning M., Halliday R.B. 1994. Biology and reproduction of some Australian species of Macrochelidae (Acarina). Aus. Ent., 21: 89-94.

Maraun M., Caruso T., Hense J., Lehmitz L., Murvanidze M., Nae I., Schulz J., Seniczak A., Scheu S. 2019. Parthenogenetic vs. sexual reproduction in oribatid mite communities. Ecol. Evol. 9(12): 7324-7332. https://doi.org/10.1002/ece3.5303

Movahedzade E., Lotfollahi P., Abbasi A. 2016. Some oppiid mites (Acari: Oribatida) of East Azerbaijan province, with two new records for the fauna of Iran. Pers. J. Acarol., 5 (2): 153-158.

Murvanidze M., Mumladze L. 2016. Annotated checklist of Georgian oribatid mites. Zootaxa, 4089 (1): 1-81. https://doi.org/10.11646/zootaxa.4089.1.1

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.

Norton R.A., Behan-Pelletier V.M. 2009. Oribatida. Chapter 15. In: Krantz G.W., Walter D.E. (Eds.). A Manual of Acarology. Lubbock: Texas Tech University Press. pp. 430-564.

Noto la Diega R., Caruso N., Bernini F. 2004. Oribatid mites of the Pelagian Islands in the southern Mediterranean Sea: a faunistic and biogeographical study. Phytophaga, XIV: 513-518.

Paoli G. 1908. Monografia del genere Dameosoma Berl. e generi affini. Redia, 5: 31-91.
Pérez-Íñigo C. 1965. Especies españolas del género *Oppia* C.L. Koch (Acari, Oribatei). Bol. Real Soc. Esp. Hist. Nat., 62: 385-416.

Pérez-Íñigo C. 1971. Acaros oribátidos de suelos de España peninsular e islas Baleares (Acari, Oribatei). Parte III. Eos, 46: 263-350.

Poltavskaya M.P. 1994. A new genus and new species of oribatid mites from the virgin steppes of the Palaearctic (Acariformes, Oribatei). Zool. Zh., 73(3): 138-141.

Schatz H. 2006. Catalogue of known oribatid mite species (Acari: Oribatida) from the Central American landbridge (First part). Trop. Zool., 19(2): 209-288.

Schuster R. 1960. Über die Morphologie und Artengliederung der Gattung *Epilohmannia* Berlese 1917 (Acari, Oribatei). Zool. Anz., 165(5/6): 197-213.

Shereef G.M. 1976. *Oppia sitnikovae*, a new species of oribatid mite in Giza region, Egypt. Bul. Zool. Soc. Egypt, 28: 8-9.

Shtanchaeva U.Ya., Subías L.S. 2010. Catalogue of oribatid mites of the Caucasus. Makhachkala: Dagestan Sci. Center, Rus. Acad. Sci. 276 p. [In Russian]

Smelyansky I.E. 2010. Structure of oribatid mite (Acariformes, Oribatida) assemblages along a vertical soil gradient in the steppe habitats of the Trans-Volga Region (Samara Province, Russia). Euroasian Ent. J., 9(2): 206-222.

Stary J. 2006. Oribatid mites (Acari: Oribatida) of some localities in Kokořínsko Protected Landscape Area, Czech Republic. Bohemia Centralis, 27: 143-160.

Subías L.S. 2004. Listado sistemático, sinonímico y biogeográfico de los ácaros oribátidos (Acariformes, Oribatida) del mundo (1758-2002). Graellsia, 60: 3-305. https://doi.org/10.3989/graellsia.2004.v60.iExtra.218

Subías L.S. 2022. Listado sistemático, sinonímico y biogeográfico de los ácaros oribátidos (Acariformes, Oribatida) del mundo (excepto fósiles). Monogr. Electr. S.E.A., 12: 1-539.

Subías L.S., Arillo A. 2001. Acari, Oribatei, Gymnonota II. Oppioidea. In: Ramos A. et al. (Eds.). Fauna Iberica. Vol. 15. Madrid: Museo de Ciencias Naturales, 289 pp.

Subías L.S., Rodríguez P. 1986. *Oppiidae* (Acari, Oribatida) de los Sabinares (*Juniperus thurifera*) de España. IX. *Subiasella* (*Lalmoppia*) n. subgen. y *Discoppia* (*Cylindroppia*) n. subgen. Rev. Biol. Univ., 4: 111-121.

Subías L.S., Shtanchaeva U.Ya. 2023. Claves de familias, géneros y subgéneros de ácaros oribátidos del mundo (Acari, Oribatida). Monogr. Electr. S.E.A., 13: 1-290.

Subías L.S., Shtanchaeva U.Ya., Arillo A. 2013. Oribátidos (Acari, Oribatida) de España peninsular e islas Baleares. Distribución. Monogr. Electr. S.E.A., 5: 1-255.

Subías L.S., Zaballos J.P., Banda E., Fontal-Cazalla F., Nieves-Aldrey J.L. 2004. Ácaros oribátidos (Acari, Oribatei) del Parque Nacional de la isla de Coiba, Panamá. Rev. Biol. Trop., 52(1): 85-95. https://doi.org/10.15517/rbt.v52i1.14755

Travé J., Vachon M. 1975. François Grandjean. 1882-1975 (Notice biographique et bibliographique). Acarologia, 17(1): 1-19.

Vasiliu N., Ivan O. 1996. Oribatids, bioindicators of the pollution of agricultural soils. Lucrarile Stiintifice ale celei de a 2-a conferinte Nat. Pentru Prot. Mediului: 370-376.

Walter D.E., Lindquist E.E. 1995. The distribution of parthenogenetic ascid mites (Acari: Parasitiformes) do not support the biotic uncertainty hypothesis. Exp. Appl. Acarol., 19: 423-442. https://doi.org/10.1007/BF00048262

Weis-Fogh T. 1948. Ecological investigations on mites and Collemboles in the soil (Appendix: Description of some new mites (Acari)). Nat. Jutland., Aarhus, 1: 135-270.