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

HAL
open science

# New species of Plateremaeoidea (Acari, Oribatida) from Mexico

Sergey G Ermilov, Andrey A Yurtaev

► To cite this version:

Sergey G Ermilov, Andrey A Yurtaev. New species of Plateremaeoidea (Acari, Oribatida) from Mexico. Acarologia, 2023, 63 (2), pp.390-410. 10.24349/0qv6-5yy2. hal-04055931

HAL Id: hal-04055931

https://hal.science/hal-04055931

Submitted on 3 Apr 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_2.png" /></figure>

Distributed under a Creative Commons Attribution 4.0 International License
<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
# New species of Plateremaeoidea (Acari, Oribatida) from Mexico

Sergey G. Ermilov IDª, Andrey A. Yurtaev IDª

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

## Original research

### ABSTRACT

Two new species of plateremaeoid oribatid mites from the genera *Jacotella* and *Plateremaeus*, *J. alexandrovskiyi* n. sp. and *P. sedovi* n. sp., are described from leaf litter of secondary semi-evergreen tropical forest in Mexico. The description of *J. alexandrovskiyi* is based on adult and tritonymphal instars, *P. sedovi* on adult and larval instars. Identification keys to known species of *Jacotella* and *Plateremaeus* are provided.

**Keywords** oribatid mites; *Jacotella*; *Plateremaeus*; taxonomy; morphology; identification key; Neotropical region

**Zoobank** http://zoobank.org/0FE5E694-9682-44E8-B35E-077F3859898D

## Introduction

During taxonomic identification of oribatid mites (Acari, Oribatida) of the superfamily Plateremaeoidea (system according to Norton and Behan-Pelletier 2009; Schatz *et al.* 2011) collected from Mexico, we found two new species belonging the genera *Jacotella* Banks, 1947 (Gymnodamaeidae) and *Plateremaeus* Berlese, 1908 (Plateremaeidae). The primary goal of the paper is to describe these two new species under the names *Jacotella alexandrovskiyi* n. sp. and *Plateremaeus sedovi* n. sp.

The description of *J. alexandrovskiyi* is based on adult and tritonymphal instars. At present, the juvenile instars of *Jacotella* are known only in two species (Seniczak and Seniczak 2011; Ermilov and Liao 2021): *J. neonominata* Subías, 2004 (all instars are studied) and *J. puyuma* Ermilov and Liao, 2021 (tritonymph). The morphological comparison for juveniles of Gymnodamaeidae was provided by Bayartogtokh and Ermilov (2013), with subsequent additions in Ermilov and Salavatulin (2022). The description of *P. sedovi* is based on adult and larval instars. At present, the juvenile instars of *Plateremaeus* are unknown; hence, data on the larva of *P. sedovi* is the first information on juveniles in this genus. The juvenile instars in Plateremaeidae are known only in one species (Seniczak *et al.* 2020): *Lopheremaeus mirabilis* (Csiszár, 1962) (all instars are studied). The additional goals of the paper are to compare known tritonymphs within *Jacotella* and larvae within Plateremaeidae, and to present identification keys to adults of known species of *Jacotella* and *Plateremaeus*.

*Jacotella* was proposed by Banks (1947), with *Gymnodamaeus quadricaudiculus* Jacot, 1937 as type species. The genus comprises 10 known species, which have collectively a semicosmopolitan distribution (Subías 2022). The main generic traits have been summarized by Paschoal (1983) and Walter (2009). *Plateremaeus* was proposed by Berlese (1908), with *Damaeus ornatissimus* Berlese, 1888 as type species. The genus comprises 10 known species (see the Discussion section below), which are distributed collectively in the Afrotropical, Australasian, Neotropical, and Oriental regions (Subías 2022). The main generic traits have been summarized by Paschoal (1987).

Presently, two species of *Jacotella* and three species of *Plateremaeus* were registered in the Mexican fauna (e.g., Palacios-Vargas and Iglesias 2004; Vázquez-González *et al.* 2016; Subías

Received 13 February 2023
Accepted 27 March 023
Published 30 March 2023

Corresponding author
Sergey G. Ermilov ID:
ermilovacari@yandex.ru

Academic editor
Pfingstl, Tobias

https://doi.org/10.24349/0qv6-5yy2

ISSN 0044-586X (print)
ISSN 2107-7207 (electronic)

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

Ermilov S. G. and Yurtaev A. A.

Licensed under
Creative Commons CC-BY 4.0
2022): *J. ornata* (Balogh and Csiszár, 1963), *J. quadricaudicula* (Jacot, 1937), *P. berlesei* Balogh and Mahunka, 1978, *P. costulatus* Balogh and Mahunka, 1978, and *P. ornatissimus* (Berlese, 1888).

# Material and methods

## Specimens

Our work is based on materials, which belong to collections of the Tyumen State University Museum of Zoology, Tyumen, Russia (Gashev *et al.* 2005).

## 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 (in juveniles: gastronotic region). Notogastral (in juveniles: gastronotic) width refers to the maximum width of the notogaster (in juveniles: gastronotic region) 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". SEM micrographs were made with the aid of a JEOL-JSM-6510LV SEM microscope.

## Terminology

Morphological terminology used in this paper mostly 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:* *r* = ridge; *P* = propodolateral apophysis; *ro, le, in, bs, ex* = rostral, lamellar, interlamellar, bothridial, and exobothridial setae, respectively. *Notogaster/gastronotic region:* *LaE, PnE, DnE* = larval, protonymphal and deutonymphal exuvial scalp, respectively; *c, da, dm, dp, lp, h, p* = setae; *ia, im, ip, ih, ips* = lyrifissures/cupules; *gla* = opisthonotal gland opening. *Gnathosoma:* *a, m, h* = subcapitular setae; *or* = adoral seta; *a.s.* = axillary saccule; *d, l, sup, inf, cm, acm, ul, su, vt, lt* = palp setae; *ω* = palp solenidion; *cha, chb* = cheliceral setae; *Tg* = Trägårdh's organ. *Epimeral and lateral podosomal regions:* *Cl* = Claparède's organ; *1a–1h, 2a–2f, 3a–3f, 4a–4j* = epimeral setae; *Sp* = posterior tubercle of parastigmatic enantiophysis; *dis* = discidium. *Anogenital region:* *g, ag, an, ad* = genital, aggenital, anal, and adanal setae, respectively; *iad* = adanal lyrifissure/cupule. *Legs:* *Tr, Fe, Ge, Ti, Ta* = trochanter, femur, genu, tibia, and tarsus, respectively; *trp* = trochanteral process; *sac* = saccule; *d, l, v, bv, ev, ft, tc, it, p, u, a, s, pv, pl* = setae; *ω, σ, φ* = solenidia. *Instar:* *La* = larva; *Tn* = tritonymph; *Ad* = adult.

# Taxonomy

## *Jacotella alexandrovskiyi* n. sp.

Zoobank: F59E346B-A1F0-4CDD-9398-E8F5E18FBEE1
(Figures 1–5)
<figure><img src="image_8.png" /><figcaption>**Figure 1** *Jacotella alexandrovskiyi* n. sp., adult: a – dorsal view (legs omitted); b – ventral view (gnathosoma and legs omitted); c – anterior part of prodorsum, lateral view; d – notogaster, lateral view. Scale bar 100 µm.</figcaption></figure>
<figure><img src="image_9.png" /><figcaption>**Figure 2** *Jacotella alexandrovskiyi n. sp.*, adult: a – subcapitulum, ventral view; b – palp, right, antiaxial view; c – chelicera, left, paraxial view; d – leg I, without trochanter, left, paraxial view; e – leg II, without tarsus and trochanter, left, ventral view; f – leg III, without tarsus, left, ventral view; g – leg IV, left, ventral view. Scale bars 50 µm (d–g), 20 µm (a–c).</figcaption></figure>
<figure><img src="image_10.png" /><figcaption>**Figure 3** *Jacotella alexandrovskiyi n. sp.*, adult, SEM micrographs: a – dorsal view; b – ventral view; c – left ventrolateral view; d – left lateral view. Scale bar 200 µm.</figcaption></figure>
<figure><img src="image_11.png" /><figcaption>**Figure 4** *Jacotella alexandrovskiyi n. sp.*, adult, SEM micrographs: a – dorsoanterior view; b – anal setae; c – notogastral cerotegument; d – anterior part of leg IV, left, antiaxial view. Scale bars 100 µm (a), 20 µm (b), 10 µm (c, d).</figcaption></figure>

**Diagnosis**

**Adult**

Body length: 570–645. Body surface microsculpturing tuberculate, covered by tubercular cerotegument. Bothridial seta very long, subflagellate, densely barbed. Central part of notogaster depressed. Four pairs of setae present: *h*₁ and *p*₁ long, subflagellate, densely barbed; *p*₂ and *p*₃ medium-sized, slightly thickened, pointed apically, densely barbed; *h*₁ longest, *p*₂,
<figure><img src="image_12.png" /><figcaption>**Figure 5** *Jacotella alexandrovskiyi n. sp.*, tritonymph (a, b) and its exuvial scalps (c, d): a – dorsal view (legs omitted); b – ventral view (legs omitted); c – larval exuvial scalp; d – proto- and deutonymphal exuvial scalps. Scale bars 100 µm (a, b), 50 µm (c, d).</figcaption></figure>
$p_3$ shortest. Epimeral, aggenital and adanal setae slightly thickened, pointed apically, densely barbed; anal seta setiform, roughened. Discidium developed. Two pairs of adanal setae present. All leg femora with saccule.

## Tritonymph

Bothridial seta and gastronotic seta $h_1$ very long, subflagellate, with short, dense cilia; other gastronotic ($c_1, c_2, c_3, lp, h_2, h_3, p_1, p_2, p_3$) and exuvial (except broken $h_1$) setae medium-sized or short, slightly thickened, pointed apically, densely barbed. Exobothridial seta comparatively long. Two pairs of adanal setae present.

## Description of adult

**Measurements** — Body length: 630 (holotype, female), 570–645 (10 paratypes: five males and five females); body width: 360 (holotype), 330–375 (10 paratypes). No differences between males and females in body size.

**Integument** — (Figs 3a–d, 4a–d) — Body color brown. Body and legs covered by thick layer of gel-like cerotegument comprising tubercular structures (diameter up to 7); band-like and reticulate cerotegumental ornamentations absent on notogaster. Body surface densely microtuberculate. Setae often with cerotegument.

**Prodorsum** — (Figs 1a, 1c, 3a, 3c, 3d, 4a) – Rostrum rounded. Interbothridial region with broad inverse V-shaped ridge-like structure; two diagonal ridges present, each located between interbothridial ridge-like structure and insertion of lamellar seta; transverse ridge located between insertions of lamellar setae (Figs 1a, 3a). Enantiophysis absent. Rostral and lamellar setae (90–94) slightly thickened, pointed apically, roughened; interlamellar seta (11) spiniform, smooth; bothridial seta (206–213) subflagellate, densely barbed; exobothridial seta (22–26) setiform, roughened.

**Notogaster** — (Figs 1a, 1d, 3a, 3d, 4a) – Dorsal side flat, with depressed central part (Fig. 3a). Exuvial scalps absent in all specimens. Humeral process not developed. Four pairs of setae present: $h_1$ (161–180) and $p_1$ (75–82) subflagellate, densely barbed, inserted on distinct tubercles; $p_2$ and $p_3$ (both 49–52) slightly thickened, pointed apically, densely barbed. Opisthonotal gland opening and all lyrifissures distinct.

**Gnathosoma** — (Figs 2a–c) – Subcapitulum size: 135–146 × 97–109; subcapitular setae (a: 37–41; m, h: 45–49) setiform, barbed; adoral seta (15–19) setiform, roughened. Axillary saccule absent. Palp length: 86–94; setation: 0–2–1–3–9(+ω); postpalpal seta (7) spiniform, roughened. Chelicera length: 127–135; cheliceral seta *cha* (34) setiform, shortly ciliate dorsally; *chb* (30) slightly thickened, rounded apically, densely barbed.

Epimeral and lateral podosomal regions (Figs 1b, 1c, 3b–d) – Epimeral setal formula: 3–1–3–3; setae (*1a, 2a, 3a, 4a*: 22–30; *1b, 1c, 3b*: 30–34; *4b*: 26–34; *3c, 4c*: 52–60) slightly thickened, pointed apically, densely barbed. Tubercle *Sa* of parastigmatic enantiophysis not observed; *Sp* slightly developed. Discidium triangular, with rounded top.

Anogenital region (Figs 1b, 3b, 3c, 4b) – Seven pairs of genital setae present, all (22–26) setiform, densely barbed; aggenital (26–30) and adanal (34–37) setae slightly thickened, pointed apically, densely barbed; anal seta (22–26) setiform, roughened; two pairs of adanal setae present. Anal lyrifissure slightly visible; adanal lyrifissure not observed.

**Legs** — (Figs 2d–g, 3a–d, 4a, 4d) – Heterotridactylous; claws slightly barbed on dorsal side. Tibia I with dorsodistal cylindrical apophysis bearing two solenidia and seta. Femora I–IV with saccule (instead porose area). Formulas of leg setation and solenidia: I (1–5–4–5–20) [1–2–2], II (1–5–4–5–16) [1–1–2], III (2–3–3–4–15) [1–1–0], IV (1–2–3–4–12) [0–1–0]; homology of setae and solenidia indicated in Table 1. Famulus on tarsus I sunken in cup-like cuticular deepening; seta *s* on tarsus I setiform, barbed (not eupathidial). Solenidion $\phi_1$ on tibia I subflagellate; other solenidia rod-like.
## Description of tritonymph

**Measurements** — Body length: 525–555 (three tritonymphs); gastronotum width: 300–330 (three tritonymphs).

**Integument** — (Figs 5a–c) — Body color grey yellowish. Body and legs covered by thick layer of gel-like cerotegument comprising tubercular structures (diameter up to 4). Gastronotum (except dorsocentral part), ventral side of body, lateral and basal parts of prodorsum folded. Setae with or without cerotegument. Exuvial scalps with reticulate ornamentation.

**Prodorsum** — (Fig. 5a) — Rostrum rounded. Dorsal side with two diagonal ridges in central part and one transverse ridge anteriorly to insertions of lamellar setae. Rostral (71–75), lamellar (71–75) and exobothridial (60–64) setae slightly thickened, pointed apically, ro and le roughened, ex densely barbed; interlamellar seta (4) spiniform, smooth; bothridial seta (206–225) subflagellate, with short, dense cilia.

**Gastronotic region** — (Figs 5a, 5b) — Gastronotum carrying exuvial scalps of previous instars; nymphal scalps with two pairs of strong posterior apophyses bearing setae $h_1$ and $p_1$; larval exuvial scalp with nine, proto- and deutonymphal scalps with seven pairs of setae, of them, $h_1$ broken, others medium-sized or short, thick (da, dm, dp in larval scalp) or slightly thickened, pointed apically, densely barbed. Ten pairs of gastronotic setae present: seta $h_1$ (225–236) subflagellate, with short, dense cilia; others ($c_1, c_3$: 52–60; $c_2$: 22–26; lp: 37–49; $h_2, h_3$: 41–56; $p_1$: 60–64; $p_2, p_3$: 26–30) slightly thickened, pointed apically, densely barbed. Opisthonotal gland opening and all cupules well visible.

**Gnathosoma** — (Fig. 5b) — Subcapitulum size: 120–124 × 94–97; subcapitular setae (a, m, h: 26–30) setiform, barbed; adoral seta (11) setiform, roughened. Axillary saccule absent. Palp length: 79; setation: 0–2–1–3–9(+ω); postpalpal seta (7) spiniform, smooth. Chelicera length: 116–120; cheliceral seta cha (30) setiform, shortly ciliate dorsally; chb (26) setiform, barbed.

**Epimeral and lateral podosomal regions** — (Fig. 5b) — Epimeral setal formula: 3–1–3–3; setae (1a, 2a, 3a: 15–19; others: 26–30) setiform, densely barbed.

**Anogenital region** — (Fig. 5b) — Genital and setae (15–19) setae setiform, roughened; aggenital (15–19) and adanal (26–34) setae setiform, densely barbed. Adanal cupule distinct.

**Legs**— Single claw of each leg strong, slightly roughened on dorsal side. Saccule and porose area on segments not observed. Formulas of leg setation and solenidia: I (1–5–4–5–18) [1–2–2], II (1–5–4–5–15) [1–1–2], III (2–3–3–4–15) [1–1–0], IV (1–2–3–4–12) [0–1–0]; homology of setae and solenidia indicated in Table 1. Famulus on tarsus I emergent.

## Material examined

Holotype (female), 10 paratypes (five males and five females) and three tritonymphs: Mexico, 20°33'N, 87°13'W, Quintana Roo, Municipio de Solidaridad, vicinities of Playa del Carmen,

<table><caption>**Table 1** Leg setation and solenidia of tritonymph and adult *Jacotella alexandrovskiyi* n. sp.</caption><thead><tr><th></th><th>Tr</th><th>Fe</th><th>Ge</th><th>Ti</th><th>Ta</th></tr></thead><tbody><tr><td colspan="6"><strong>Leg I</strong></td></tr><tr><td>Tn</td><td>v'</td><td>d, (l), bv'', v''</td><td>d, (l), v', σ</td><td>(l), (v), d φ₁, φ₂</td><td>(ft), (tc), (it), (p), (u), (a), s, (pv), (pl), ε, ω₁, ω₂</td></tr><tr><td>Ad</td><td>v'</td><td>d, (l), bv'', v''</td><td>d, (l), v', σ</td><td>(l), (v), d φ₁, φ₂</td><td>(ft), (tc), (it), (p), (u), (a), s, (pv), v', (pl), l'', ε, ω₁, ω₂</td></tr><tr><td colspan="6"><strong>Leg II</strong></td></tr><tr><td>Tn</td><td>v'</td><td>d, (l), bv'', v''</td><td>d, (l), v', σ</td><td>d, (l), (v), φ</td><td>(ft), (tc), (it), (p), (u), (a), s, (pv), l'', ω₁, ω₂</td></tr><tr><td>Ad</td><td>v'</td><td>d, (l), bv'', v''</td><td>d, (l), v', σ</td><td>d, (l), (v), φ</td><td>(ft), (tc), (it), (p), (u), (a), s, (pv), l'', ω₁, ω₂</td></tr><tr><td colspan="6"><strong>Leg III</strong></td></tr><tr><td>Tn</td><td>l', v'</td><td>d, l', ev'</td><td>d, l', v', σ</td><td>d, l', (v), φ</td><td>(ft), (tc), (it), (p), (u), (a), s, (pv)</td></tr><tr><td>Ad</td><td>l', v'</td><td>d, l', ev'</td><td>d, l', v', σ</td><td>d, l', (v), φ</td><td>(ft), (tc), (it), (p), (u), (a), s, (pv)</td></tr></tbody></table>
leaf litter under trees and bushes in secondary semi-evergreen tropical forest (data and collector unknown; collection of the Tyumen State University Museum of Zoology, Tyumen, Russia; Gashev *et al.* 2005).

**Type deposition**

The holotype is deposited in the collection of the Senckenberg Museum of Natural History, Görlitz, Germany; 10 paratypes and three tritonymphs 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 species is named in honor of our friend and colleague, well-known soil scientist Dr. Alexander L. Alexandrovskiy (Institute of Geography, Russian Academy of Sciences, Moscow, Russia).

**Remarks**

The adult of *Jacotella alexandrovskiyi n. sp.* differs from all representatives of the genus by the presence of very long (versus distinctly shorter), subflagellate (versus with slightly or well developed head) bothridial seta. *Jacotella ornata* (Balogh and Csiszár, 1963), which is known from the Neotropical region and India, and *J. puyuma* Ermilov and Liao, 2021, which is known from Taiwan, have almost setiform bothridial seta with slightly developed elongate head due to cerotegument, however, the new species differs from both by the presence of two pairs (versus three) of adanal setae. Also, *J. ornata* has specifically band-like cerotegument (versus absent) on the notogaster and five (versus seven) pairs of genital setae; *J. puyuma* is smaller (length: 365–415 versus 570–645) and has (versus absent) specific ridge-like structures in the central part of notogaster and deep depression in anterior part of the prodorsum.

The tritonymphal instar in *Jacotella* is known for *J. neonominata* (see Seniczak and Seniczak 2011) and *J. puyuma* (see Ermilov and Liao 2021). The tritonymph of *J. alexandrovskiyi* differs from those in both species by the length and morphology of bothridial (very long, subflagellate, with short, dense cilia versus distinctly shorter, fusiform, barbed), exobothridial (long, thickened versus minute, spiniform) and gastronotic ($h_1$ very long, subflagellate, others medium-sized or short, slightly thickened versus $c_3$ very long, subflagellate, others medium-sized or short, phylliform except long seta $h_1$ in *J. puyuma*), and exuvial (*da*, *dm*, *dp* medium-sized, thick $c_1$, $c_2$, *la*, *lm*, *lp*, $h_2$, $h_3$, $p_1$ shorter, slightly thickened versus *da*, *dm*, *dp*, $c_1$, $c_2$, *la*, *lm*, *lp*, $h_2$, $h_3$, $p_1$ mostly short, phylliform) setae, and the number (two pairs versus three pairs) of adanal setae.

**Platermaeus sedovi n. sp.**

Zoobank: AB0C5C94-471C-4DF6-967A-D5CCC5C673CD

(Figures 6–10)

**Diagnosis**

**Adult**

Body length: 870–975. Body surface microsculpturing tuberculate, covered by tubercular and shortly filamentous cerotegument; subcapitular mentum, dorsal part of prodorsum, lateral part of epimere I, and leg femora I–IV and trochanters III, IV partially foveolate. Rostrum rounded, without lateral teeth. Enantiophysis absent. Bothridial seta long, thickened, slightly barbed, distally bent having unilateral short cilia creating illusion of a slightly developed head. Five pairs of medium-sized, setiform, barbed notogastral setae (*lp*, $h_1$, $h_2$, $p_2$, $p_3$) present; $h_1$ longest. Epimeral setal formula: 8–6–6–10. Epimeral and adanal setae slightly thickened, pointed apically, roughened; genital, aggenital and anal setae setiform, slightly roughened. Seven
<figure><img src="image_13.png" /><figcaption>**Figure 6** *Plateremaeus sedovi n. sp.*, adult: a – dorsal view (legs omitted); b – ventral view (gnathosoma and legs omitted); c – anterior part of prodorsum, lateral view; d – notogaster, lateral view. Scale bar 200 µm.</figcaption></figure>

pairs of genital setae present. Adanal setae *ad*₂ and *ad*₃ located close to anal aperture. Leg trochanters III, IV with dorsodistal process.

Larva

Subcapitular mentum and partially prodorsum reticulate; gastronotum, epimeral and anogenital regions folded. Bothridial seta with large, fusiform, barbed head. Eight pairs of gastronomic setae (*c*₁, *c*₂, *dm*, *dp*, *lp*, *h*₁, *h*₂, *h*₃) present: *h*₃ represented by small alveolus; *lp*, *h*₁ long,
<figure><img src="image_14.png" /><figcaption>**Figure 7** *Plateremaeus sedovi n. sp.*, adult: a – subcapitulum, ventral view; b – palp, right, antiaxial view; c – chelicera, left, paraxial view; d – leg I, without trochanter, right, antiaxial view; e – leg II, without tarsus, right, antiaxial view; f – leg III, without tarsus, right, paraxial view; g – leg IV, left, antiaxial view. Scale bars 100 µm (d–g), 50 µm (a, c), 20 µm (b).</figcaption></figure>
<figure><img src="image_15.png" /><figcaption>**Figure 8** *Platermaeus sedovi n. sp.*, adult, SEM micrographs: a – dorsal view; b – ventral view; c – left ventrolateral view; d – left lateral view. Scale bar 200 µm.</figcaption></figure>
<figure><img src="image_16.png" /><figcaption>**Figure 9** *Platermaeus sedovi n. sp.*, adult, SEM micrographs: a – dorsoanterior view; b – cerotegument and seta 2e on epimere II; c – anal setae; d – anterior part of leg IV, left, antiaxial view; e – notogastral cerotegument. Scale bars 200 µm (a), 50 µm (d), 20 µm (c), 10 µm (b), 1 µm (e).</figcaption></figure>

subflagellate, barbed; others medium-sized, setiform, barbed.

**Description of adult**

**Measurements** — Body length: 900 (holotype: male), 870–975 (eight paratypes: four males and four females); body width: 675 (holotype), 660–675 (eight paratypes). No differences
<figure><img src="image_17.png" /><figcaption>**Figure 10** *Platermaeus sedovi n. sp.*, larva: a – dorsal view (legs omitted); b – ventral view (legs omitted); c – larval exuvial scalp; d – proto-and deutonymphal exuvial scalps. Scale bar 100 µm.</figcaption></figure>

between males and females in body size.

**Integument** — (Figs 8a–d, 9a, 9b, 9d, 9e) – Body color brown to dark brown. Body and legs covered by thick layer of gel-like cerotegument comprising microgranular and shortly filamentous structures; reticulate cerotegumental pattern absent. Body surface densely microtuberculate; subcapitular mentum, dorsal part of prodorsum, lateral part of epimere I, and leg femora I–IV and trochanters III, IV partially foveolate (diameter up to 7). Setae with or without cerotegument.

**Prodorsum** — (Figs 6a, 6c, 8a, 8c, 8d, 9a) – Rostrum rounded, without lateral teeth. Centrodorsal part of prodorsum slightly depressed (Fig. 8a). Enantiophysis absent. Propodolateral apophysis large, thorn-like. Rostral and lamellar setae (135–150) setiform, roughened; interlamellar seta (9–11) spiniform, smooth; bothridial seta (195–202) thickened, slightly barbed, distally bent having unilateral short cilia creating illusion of a slightly developed head; exobothridial seta absent.
**Notogaster** — (Figs 6a, 6d, 8a, 8d, 9a) – Dorsal side flat. Exuvial scalps absent in all specimens. Humeral process not developed. Five pairs of setae present, all (lp: 82–90; h₁: 176–187; h₂, p₂, p₃: 60–67) setiform, barbed. Opisthonotal gland opening and all lyri fissures distinct.

**Gnathosoma** — (Figs 7a–c) – Subcapitulum size: 165–169 × 142–150; subcapitular rutellum mediodistal tooth; subcapitular mentum with broad transverse ridge; subcapitular (a, m: 45–49; h: 79–86) and adoral (19) setae setiform, roughened. Axillary saccule present. Palp length: 101–105; setation: 0–2–1–3–8(+ω); postpalpal seta (7) spiniform, roughened. Chelicera length: 161–169; cheliceral setae (cha: 75–82; chb: 30–32) setiform, barbed.

**Epimeral and lateral podosomal regions** — (Figs 6b, 6c, 8b–d) – Epimeral setal formula: 8–6–6–10; setae (1b: 52–60; 1c: 64–67; 1d: 97–106; others: 41–45) slightly thickened, pointed apically, roughened. Tubercles of parastigmatic enantiophysis and discidium absent.

**Anogenital region** — (Figs 6b, 8b, 8c, 9c) – Genital, aggenital and anal setae (22–30) setiform, slightly roughened; adanal seta (64–67) slightly thickened, pointed apically, roughened; seven pairs of genital setae present; adanal setae *ad*₂ and *ad*₃ located close to anal aperture. Anal and adanal lyri fissures slightly visible.

**Legs** — (Figs 7d–g, 8a–d, 9a, 9d) – Heterotridactylous; claws slightly barbed on dorsal side. Tibia I with dorsodistal cylindrical apophysis bearing two solenidia. Trochanter III with small tooth-like dorsodistal triangular process; trochanter IV with large triangular dorsodistal triangular process. Femora I–IV and trochanters III, IV without porose area and saccule. Formulas of leg setation and solenidia: I (1–8–4–4–20) [1–2–2], II (1–7–4–5–16) [1–1–2], III (3–5–3–4–15) [1–1–0], IV (3–5–3–4–12) [0–1–0]; homology of setae and solenidia indicated in Table 2. Famulus on tarsus I sunken in cup-like cuticular deepening; seta *s* on tarsus I setiform, roughened (not eupathidial). Solenidion φ₁ on tibia I subflagellate; other solenidia rod-like.

## Description of larva

**Measurements** — Body length: 435–450 (three larvae); gastronotum width: 270–300 (three larvae).

**Integument** — (Figs 10a, 10b) – Body color grey yellowish. Body and legs covered by thick layer of gel-like cerotegument comprising amorphous, granular and bacillar structures; bacillar structures often additionally covered by small cerotegumental scales. Prodorsum partially reticulate; gastronotum and ventral side of body folded. Setae with or without cerotegument.

**Prodorsum** — (Fig. 10a) – Rostrum rounded. Rostral and lamellar setae (49–60) setiform,

<table><caption>**Table 2** Leg setation and solenidia of larva and adult *Plateremaeus sedovi* n. sp.</caption><thead><tr><th></th><th>Tr</th><th>Fe</th><th>Ge</th><th>Ti</th><th>Ta</th></tr></thead><tbody><tr><td>Leg I</td><td></td><td></td><td></td><td></td><td></td></tr><tr><td>La</td><td>—</td><td>d, bv"</td><td>(l), σ</td><td>(l), v', φ₁</td><td>(ft), (tc), (p), (u), (a), s, (pv), (pl), ε, ω₁</td></tr><tr><td>Ad</td><td>v'</td><td>d, (l₁), (l₂), bv", (v)</td><td>(l), (v), σ</td><td>(l), (v), φ₁, φ₂</td><td>(ft), (tc), (it), (p), (u), (a), s, (pv), v', (pl), l", ε, ω₁, ω₂</td></tr><tr><td>Leg II</td><td></td><td></td><td></td><td></td><td></td></tr><tr><td>La</td><td>—</td><td>d, bv"</td><td>(l), σ</td><td>d, l', v', φ</td><td>(ft), (tc), (p), (u), (a), s, (pv), ω₁</td></tr><tr><td>Ad</td><td>v'</td><td>d, (l₁), (l₂), bv", v"</td><td>(l), (v), σ</td><td>d, (l), (v), φ</td><td>(ft), (tc), (it), (p), (u), (a), s, (pv), l", ω₁, ω₂</td></tr><tr><td>Leg III</td><td></td><td></td><td></td><td></td><td></td></tr><tr><td>La</td><td>—</td><td>d, ev'</td><td>d, l', σ</td><td>d, v', φ</td><td>(ft), (tc), (p), (u), (a), s, (pv)</td></tr><tr><td>Ad</td><td>l', v₁', v₂'</td><td>d, l₁', l₂', l₃', ev'</td><td>d, l', v', σ</td><td>d, l', (v), φ</td><td>(ft), (tc), (it), (p), (u), (a), s, (pv)</td></tr><tr><td>Leg IV</td><td></td><td></td><td></td><td></td><td></td></tr><tr><td>Ad</td><td>v₁', v₂', v₃'</td><td>d, l', ev', (v)</td><td>d, l', v'</td><td>d, l', (v), φ</td><td>ft", (tc), (p), (u), (a), s, (pv)</td></tr></tbody></table>
slightly barbed; interlamellar (30–41) and exobothridial (22–30) setae setiform, roughened,
*ro* and *le* thicker than *in* and *ex*; bothridial seta (131–139) fusiform, barbed.

**Gastronotic region** — (Fig. 10a, 10b) — Eight pairs of gastronotic setae present: *h*₃ represented by small alveolus; *lp* (94–105) and *h*₁ (157–176) long, subflagellate, barbed; others (*c*₁, *c*₃: 30–41; *dm*, *dp*: 41–52; *h*₂: 37–49) setiform, barbed. Opisthonotal gland opening and cupules (*ia*, *im*, *ip*, *ih*) well visible.

**Gnathosoma** — (Fig. 10b) — Subcapitulum size: 90–94 × 84–86; subcapitular setae (*a*, *h*: 17–19; *m*: 9–11) setiform, roughened; adoral seta (7) setiform, smooth. Axillary saccule present. Palp length: 45; setation: 0–2–1–3–8(+ω); postpalpal seta (5) spiniform, smooth. Chelicera length: 94–101; cheliceral setae (*cha*: 34; *chb*: 15) setiform, barbed.

**Epimeral and lateral podosomal regions** — (Fig. 10b) — Epimeral setal formula: 3–1–2 (third seta of first epimere forms a protective scale over respective Claparède's organ); setae (*la*, *2a*, *3a*: 11; *1b*: 15; *3b*: 22–26) setiform, roughened.

**Anogenital region** — (Fig. 10b) — Without structures and setae.

**Legs** — Single claw of each leg strong, slightly roughened on dorsal side. Saccule and porose area on segments not observed. Formulas of leg setation and solenidia: I (0–2–2–3–16) [1–1–1], II (0–2–2–3–13) [1–1–1], III (0–2–2–2–13) [1–1–0]; homology of setae and solenidia indicated in Table 2. Famulus on tarsus I emergent.

**Material examined**

Holotype (male), eight paratypes (four males and four females) and three larvae: Mexico, 20°33′N, 87°13′W, Quintana Roo, Municipio de Solidaridad, vicinities of Playa del Carmen, leaf litter under trees and bushes in secondary semi-evergreen tropical forest (data and collector unknown; collection of the Tyumen State University Museum of Zoology, Tyumen, Russia; Gashev *et al.* 2005).

**Type deposition**

The holotype is deposited in the collection of the Senckenberg Museum of Natural History, Görlitz, Germany; eight paratypes and three larvae 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 species is named in honor of our friend and colleague, well-known soil scientist Dr. Sergey N. Sedov (National Autonomous University of Mexico, Mexico City, Mexico).

**Remarks**

In the presence of foveolae on prodorsum and in the absence of prodorsal enantiophysis, the
adult of *Plateremaeus sedovi* n. sp. is similar to *Plateremaeus coralgablensis* (Paschoal, 1987)
(see the Discussion section below), which is known from the U.S.A. (Florida), however, the
new species differs from the latter by: larger body size (length: 870–975 versus 667–742); the
density of the arrangement of prodorsal foveolae (sparse versus dense); the presence (versus
absence) of foveolae on subcapitular mentum and in lateral part of epimere I; the absence
(versus presence) of lateral cerotegumental reticulation in lateral part of the notogaster; the
presence of five pairs (versus six) of anal setae; number of epimeral setae on epimeres II–IV
(II with six pairs of setae, III+IV with 16 pairs of setae versus with seven and 17, respectively).

The larval instar in Plateremaeidae is known for *Lopheremaeus mirabilis* (see Seniczak *et al.* 2020). The larva of *P. sedovi* differs from that in the latter by longer interlamellar and exobothridial setae, the absence (versus presence) of gastronotic seta *da*, and the length and morphology of gastronotic setae *lp* and *h*₁ (long, subflagellate versus medium-sized, setiform).
Discussion

*Allodamaeus* was proposed by Banks (1947), with *Allodamaeus ewingi* Banks, 1947 as type species. Taxonomic history of the genus and main generic traits were presented by Paschoal (1987). The validity of the genus *Allodamaeus* is supported by some authors (e.g. Subías 2022). The genus comprises two species, which are distributed collectively in the Polarctic and Neotropical regions (Paschoal 1987). However, *Allodamaeus* and *Plateremaeus* are similar in general, differing (according to Paschoal 1987; Balogh and Balogh 1992) mainly only in the number of neotrichous anal setae (seven pairs versus six pairs). The problem is that some species of *Plateremaeus* have a variation in the number of anal setae (5–7 pairs) or less than seven pairs (e.g., see Balogh and Mahunka 1978; Ermilov & Starý 2021), therefore, the division of *Allodamaeus* and *Plateremaeus*, which is based on the number of anal setae, seems problematic. Balogh and Balogh (2002) included *Allodamaeus coralgablensis* Paschoal, 1987 in *Plateremaeus*, automatically synonymizing *Allodamaeus* to *Plateremaeus*. We agree with the opinion of Balogh and Balogh (2002) and consider both known species of *Allodamaeus* (*A. coralgablensis*, *A. ewingi*) within *Plateremaeus*.

Key to known species of Jacotella

1. Bothridial seta setiform or with slightly developed, elongate head 2
— Bothridial seta with large, compact head 5

2. Notogaster without system of cuticular or cerotegumental ridges in dorsocentral part 3
— Notogaster with system of cuticular or cerotegumental ridges in dorsocentral part 4

3. Bothridial seta subflagellate; epimeral, aggenital and adanal setae slightly thickened; central part of notogaster depressed; body length: 570–645. Jacotella alexandrovskiyi n. sp.
Distribution: Mexico.

— Bothridial seta with slightly developed elongate head due cerotegument; epimeral, aggenital and adanal setae slightly setiform; central part of notogaster not depressed; body length: 350–451. Jacotella quadricaudicula (Jacot, 1937). Distribution: Nearctic, Mexico.

4. Prodorsum and notogaster with dense microreticulate cerotegument; anterior part of prodorsum with depression; body length: 365–415. Jacotella puyuma Ermilov and Liao, 2021. Distribution: Taiwan.

— Prodorsum and notogaster without microreticulate cerotegument; anterior part of prodorsum without depression; body length: 560. Jacotella ornata (Balogh and Csiszár, 1963). Distribution: Neotropical, India.

5. Notogaster without system of cuticular or cerotegumental ridges in dorsocentral part 6
— Notogaster with system of cuticular or cerotegumental ridges in dorsocentral part 8

6. Notogaster with macroreticulate cerotegument; body length: 450–462. Jacotella reticulata Ruiz, Kahwash and Subías, 1990. Distribution: western Mediterranean.
— Notogaster without macroreticulate cerotegument 7

7. Notogastral seta $h_1$ long, others ($p_1, p_2, p_3$) comparatively short, similar in length; body length: 450. Jacotella austriaca (Willmann, 1935). Distribution: Palaearctic.

— Notogastral setae $h_1$ long, $p_1$ medium-sized, both distinctly longer than short $p_2$ and $p_3$; body length: 366–520. Jacotella glabra (Mihelčič, 1957). Distribution: southern Palaearctic, India.

8. Notogaster (partially) and anogenital region with macroreticulate cerotegument; body length: 332–472. Jacotella frondeus (Kulijev, 1979). Distribution: southern Palaearctic, Ethiopia.
— Notogaster and anogenital region without macroreticulate cerotegument; body length: 353–545. *Jacotella neonominata* Subías, 2004. Distribution: Mediterranean, Ethiopia.

## Key to known species of *Platermaeus*

1. Seven pairs of genital setae .................................................................................... 2
— Nine pairs of genital setae .................................................................................... 6

2. Prodorsum partially foveolate.................................................................................... 3
— Prodorsum not foveolate .................................................................................... 4

3. Prodorsal foveolae located sparsely; subcapitular mentum and lateral part of epimere I partially foveolate; notogaster without reticulate cerotegument; five pairs of anal setae; epimere II with six pairs of setae, III+IV with 16 pairs of setae; body length: 870–975. .................................................. *Platermaeus sedovi* n. sp. Distribution: Mexico.

— Prodorsal foveolae located densely; subcapitular mentum and lateral part of epimere I not foveolate; notogaster partially with reticulate cerotegument; six pairs of anal setae; epimere II with seven pairs of setae, III+IV with 17 pairs of setae; body length: 667–742. ................................. *Platermaeus coralgablensis* (Paschoal, 1987). Distribution: U.S.A. (Florida).

4. Prodorsal enantiophysis present; rostrum with two lateral thorn-like structures; notogastral seta *lp* located posterolaterally on notogaster; seven pairs of anal setae; body length: 818–1000. .................................................. *Platermaeus ornatissimus* (Berlese, 1888). Distribution: Neotropical.

— Prodorsal enantiophysis absent; rostrum without lateral thorn-like structures; notogastral seta *lp* located dorsally on notogaster; six pairs of anal setae .................................................... 5

5. Bothridial seta mediodistally with dense cerotegument forming an illusory head; notogastral setae *lp* and *h*₁ slightly differ in length; body length: 686–701. .................................................. *Platermaeus ewingi* (Banks, 1947). Distribution: Holarctic.

— Bothridial seta subflagellate; notogastral seta *lp* distinctly longer than *h*₁; body length: 517–619. .................................................. *Platermaeus latus* Balogh, 1988. Distribution: Sri Lanka.

6. Anterior part of notogaster without setae .................................................................................... 7
— Anterior part of notogaster with setae .................................................................................... 8

7. Prodorsum and notogaster with reticulate cerotegument; bothridial seta clavate; five pairs of anal setae; adanal seta *ad*₃ removed from anal aperture; body length: 597–630. ................................. *Platermaeus yurtaevi* Ermilov and Starý, 2021. Distribution: Bolivia.

— Prodorsum and notogaster without reticulate cerotegument; bothridial seta mediodistally with dense cerotegument forming an illusory head; six pairs of anal setae; adanal seta *ad*₃ located close to anal aperture; body length: 784. .................................................. *Platermaeus novemsetosus* Balogh and Balogh, 1983. Distribution: Australasian.

8. Anterior part of notogaster with one pair of setae; bothridial seta mediodistally with dense cerotegument forming an illusory head; body length: 550–610. .................................................. *Platermaeus anteriosetosus* Woas, 1992. Distribution: Peru.

— Anterior part of notogaster with two pairs of setae; bothridial seta subflagellate ................................. 9

9. Prodorsal enantiophysis present; notogaster partially foveolate; notogastral setae in anterior part of notogaster minute; exobothridial seta short; body length: 737–786. ................................. *Platermaeus costulatus* Balogh and Mahunka, 1978. Distribution: Neotropical.

— Prodorsal enantiophysis absent; notogaster not foveolate; notogastral setae in anterior part
of notogaster medium-sized; exobothridial seta medium-sized; body length: 632–689.........
*Plateremaeus berlesei* Balogh and Mahunka, 1978. Distribution: Neotropical.

## Acknowledgements

We thank two anonymous reviewers for valuable comments and Nikita Shulaev (Tyumen State University, Tyumen, Russia) for SEM micrographs. This research was partially supported by the cooperative agreement No. FEWZ-2021-0004 from the Russian Ministry of Science and Higher Education.

## ORCID

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

Andrey A. Yurtaev https://orcid.org/0000-0003-1780-2598

## References

Ayyildiz N., Toluk A. 2016. Contributions to the Turkish oribatid mite fauna (Acari: Oribatida). Turk. J. Ent., 40(1): 73-85. https://doi.org/10.16970/ted.93525

Banks N. 1947. On some Acarina from North Carolina. Psyche, Cambridge Mass., 54: 110-141.
https://doi.org/10.1155/1947/70181

Balogh J., Balogh P. 1983. New oribatid mites from Australia (Acari: Oribatei). Acta Zool. Acad. Sci. Hung., 29(1-3): 81-105.

Balogh J., Balogh P. 1992. The oribatid mite genera of the World. Vol. 1. Budapest: Hungarian National Museum Press. 263 p.

Balogh J., Balogh P. 2002. Identification keys to the oribatid mites of the Extra-Holarctic regions. Vol. 1. Miskolc: Well-Press Publishing Limited. 453 p.

Balogh J., Csiszár J. 1963. The zoological results of Gy. Topál's collectings in South Argentina. 5. Oribatei (Acarina). Ann. Hist. Nat. Mus. Nat. Hung., 55: 463-485.

Balogh J., Mahunka S. 1978. New data to the knowledge of the oribatid fauna of the Neogea (Acari). III. Acta Zool. Acad. Sci. Hung., 24(3-4): 269-299.

Balogh P. 1988. Oribatid mites (Acari) from Sri Lanka. Acta Zool. Hung., 34(2-3): 171-189.

Bayartogtokh B., Ermilov S.G. (2013) Ontogenetic stages of *Gymnodamaeus irregularis*, with remarks on morphology of the juveniles of Gymnodamaeidae (Acari: Oribatida: Plateremaeoidea). Int. J. Acarol., 39(1): 7-25. https://doi.org/10.1080/01647954.2012.739640

Bayartogtokh B., Smelyansky I.E. 2004. Oribatid mites of the superfamilies Gymnodamaeoidea and Plateremaeoidea (Acari: Oribatida) from Steppe of Russia. Mong. J. Biol. Sci., 2(1): 3-17.
https://doi.org/10.22353/mjbs.2004.02.01

Berlese A. 1888. Acari Austro-Americani quos collegit Aloysius Balzan et illustravit Antonius Berlese. Manipulus primus. Species novas circiter quinquaginta complectens. Bol. Soc. Ent. Ital., 20: 1-52.

Berlese A. 1908. Elenco di generi e specie nuove di Acari. Redia, 5: 1-15.

Csiszár J., Jeleva M. 1962. Oribatid mites (Acari) from Bulgarian soils. Acta Zool. Acad. Sci. Hung., 8(3-4): 273-301.

Ermilov S.G., Liao J.-R. 2021. New faunistical and taxonomic data on oribatid mites (Acari: Oribatida) of Taiwan. Acarologia, 61(2): 297-320. https://doi.org/10.24349/acarologia/20214432

Ermilov S.G., Salavatulin V.M. 2022. Ontogenetic instars of *Gymnodamaeus uslaminorum* sp. nov. (Acari, Oribatida, Gymnodamaeidae) from Russia. Int. J. Acarol., 48(2): 98-105. https://doi.org/10.1080/01647954.2022.2033319

Ermilov S.G., Starý J. 2021. New faunistical and taxonomic data on oribatid mites of the superfamily Plateremaeoidea (Acari, Oribatida) of Bolivia. Syst. Appl. Acarol., 26(7): 1361-1373. https://doi.org/10.11158/saa.26.7.14

Gashev S.N., Elifanov A.V., Tolstikov A.V. 2005. Zoological Museum of the Tyumen State University as the oldest center for studying biodiversity in the Tyumen District. Bul. Tyumen State Univ., 3: 177-189.

Jacot A.P. 1937. Journal of North-American moss-mites. J. New York Ent. Soc., 45(3-4): 353-375.

Kulijev K.A. 1979. New species of oribatid mites of the genera *Plesiodamaeus*, *Allodamaeus*, *Carabodes*, *Hermanniella*. Dokl. Acad. Nauk Azerbaijan SSR, 35(12): 78-83.

Kunst M. 1959. Bulgarische Oribatiden (Acarina) III. Acta Univ. Carolinae Biol., 6(1): 51-74.

Mahunka S. 1979. Neue und interessante Milben aus dem Genfer Museum. XLI. Vierter Beitrag zur Kenntnis der Oribatidenfauna Griechenlands (Acari: Oribatida). Rev. suisse Zool., 86(2): 541-571.
https://doi.org/10.5962/bhl.part.82319

Mahunka S., Mahunka-Papp L. 1995. The oribatid species described by Berlese (Acari). Budapest: Hung. Nat. Hist. Mus. 325 p.
Mihelčič F. 1957. Oribatiden Südeuropas VII. Zool. Anz., 159: 44-68.

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. In: Krantz G.W., Walter D.E. (Eds). A Manual of Acarology (TX). Lubbock: Texas University Press. Chapter 15. pp. 430-564.

Palacios-Vargas J.G., Iglesias R. 2004. Oribatei (Acari). In: Bousquets J.L., Morrone J.J., Ordoñez O.Y., Fernández I.V. (Eds.). Biodiversidad, Taxonomía y Biogeografía de Artrópodos de México: Hacia una síntesis de su conocimiento. Facultad de Ciencias, Universidad Nacional Autónoma de México, México, D.F., vol. IV. pp. 431-468.

Paschoal A.D. 1983. A revision of the genus *Jacotella* (Acari, Oribatei, Gymnodamaeidae) in North America with description of two new species. Rev. Bras. Entomol., 27(2): 127-135.

Paschoal A.D. 1987. A revision of the Plateremaeidae (Acari: Oribatei). Rev. Bras. Zool., 3(6): 327-356.
https://doi.org/10.1590/S0101-81751986000200001

Pérez-Inigo C. 1970. Acaros oribátidos de suelos de España peninsular e islas Baleares (Acari, Oribatei). Parte II. Eos, Rev. Esp. Ent., 45: 241-317.

Pérez-Inigo C. 1972. Acaros oribátidos de la Isla de Tenerife. (Primera parte). Bol. R. Soc. Esp. Hist. Nat. (Biol.), 70: 185-206.

Perez-Inigo C. 1997. Acari. Oribatei. Gymnonota I. In: Ramos A. et al. (Eds.). Fauna Iberica. Mus. Cien. Nat., Madrid, vol. 9. 373 p.

Ruiz E., Kahwash M.A.M. & Subias L.S. 1990. Cuatro nuevos Gymnodamaeoideos del Sur de España. (Acari, Oribatida, Gymnodamaeoidea). Bol. R. Soc. Esp. Hist. Nat. (Sec. Biol.), 85(1-4): 39-49.

Schatz H., Behan-Pelletier V.M., OConnor B.M., Norton R.A. 2011. Suborder Oribatida van der Hammen, 1968. Zootaxa, 3148: 141-148. https://doi.org/10.11646/zootaxa.3148.1.26

Seniczak S., Seniczak A. 2011. Ontogenetic studies of three species of Gymnodamaeidae (Acari: Oribatida) with a focus on regressions of hysterosomal setae. J. Nat. Hist., 45(5-6): 361-391.
https://doi.org/10.1080/00222933.2010.534188

Seniczak S., Ivan O., Kaczmarek S., Seniczak A. 2020. Morphological ontogeny of *Lopheremaeus mirabilis* (Acari: Oribatida: Plateremaeidae), and comments on *Lopheremaeus Paschoal*. Syst. Appl. Acarol., 25(12): 2147-2164. https://doi.org/10.11158/saa.25.12.1

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

Subías L.S. 2022. Listado sistemático, sinonímico y biogeográfico de los ácaros oribátidos (Acariformes: Oribatida) del mundo (excepto fósiles). Monogr. Electrón. S.E.A., 12: 1-538.

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

Vázquez-González M.M., May Uicab D.A., Alamilla-Pastrana E.B. 2016. Riqueza específica y biodiversidad de Cozumel, Quintana Roo, México. Teoría y Praxis, 19: 137-171. https://doi.org/10.22403/UQRDOMX/TYP19/07

Walter D.E. 2009. Genera of Gymnodamaeidae (Acari: Oribatida: Plateremaeoidea) of Canada, with notes on some nomenclatorial problems. Zootaxa, 2206: 23-44. https://doi.org/10.11646/zootaxa.2206.1.2

Warburton C. 1912. The Percy Sladen Trust Expedition to the Indian Ocean in 1905. XX. The Acarina of the Seychelles. Transl. Linn. Soc. London, 2nd ser., Zool., 15: 349-360. https://doi.org/10.1111/j.1096-3642.1912.tb00105.x

Willmann C. 1935. IV. Die Milbenfauna - Oribatei. Zool. Jahrb. Syst., 66: 331-344.

Woas S. 1992. Beitrag zur Revision der Gymnodamaeidae Grandjean, 1954 (Acari, Oribatei). Andrias, 9: 121-161.