# Revision of the family Dolichocybidae (Acari: Heterostigmata) from the collection of V.D. Sevastianov

Alexander A. Khaustov, Viacheslav A. Trach

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Alexander A. Khaustov, Viacheslav A. Trach. Revision of the family Dolichocybidae (Acari: Heterostigmata) from the collection of V.D. Sevastianov. Acarologia, 2018, 58 (4), pp.897-918. hal-01917048

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# Revision of the family Dolichocybidae (Acari: Heterostigmata) from the collection of V.D. Sevastianov

Alexander A. Khaustovª, Viacheslav A. Trachᵇ,ᶜ

## ABSTRACT

The dolichocybid mites in the collection of V.D. Sevastianov, namely *Pavania riparia* Sevastianov, 1980, *P. tadjikistanica* Sevastianov, 1980, *P. protracta* Sevastianov, 1980, *P. tahanae* Sevastianov and Abo-Korah, 1985 and *Dolichocybe firjusae* Sevastianov and Chydyrov, 1994, are revised and redescribed based on both the type and additional materials. An updated key to species of the genus *Pavania* is provided. The taxonomic position of *Pavania protracta* and *P. tahanae* is discussed.

**Keywords** systematics; redescription; *Pavania*; *Dolichocybe*; dung beetles; soil

**Zoobank** http://zoobank.org/3C207D47-8CE2-41A8-89E4-AFEAF392C621

## Introduction

The family Dolichocybidae is a small group of early-derivative heterostigmatic mites that currently includes 2 subfamilies, 6 genera and 45 species (Hajiqanbar and Khaustov 2010; Rahiminejad et al. 2011; Zhang et al. 2011; Loghmani et al. 2013; Katlav et al. 2014; Bahramian et al. 2015; Mortazavi et al. 2015; Sobhi et al. 2017; Khaustov and Frolov 2017, 2018a, b; Khaustov and Trach 2017; Khaustov 2017). Little is known about the way of life of dolichocybid mites, but all of them are probably fungivorous (Rack 1967; Magowski 1988; Kaliszewski et al. 1995). Some species are important pests of edible mushrooms (Lan et al. 2017).

The identification of many dolichocybid mites is difficult or even impossible due to their incomplete and inadequate descriptions. All species described by V.D. Sevastianov and his co-authors belong to this “problematic” group. The list of dolichocybids described by V.D. Sevastianov and his co-authors includes 6 species, namely *Formicomotes octipes* Sevastianov, 1980, *Pavania riparia* Sevastianov, 1980, *P. tadjikistanica* Sevastianov, 1980, *P. protracta* Sevastianov, 1980, *P. tahanae* Sevastianov and Abo-Korah, 1985, and *Dolichocybe firjusae* Sevastianov and Chydyrov, 1994 (Sevastianov 1980; Sevastianov & Abo-Korah 1985; Sevastianov et al. 1994). Only *Formicomotes octipes* was recently redescribed (Khaustov and Frolov 2018a), while other species remain unstudied since the times of their original descriptions.

The main aim of this article is to provide redescriptions of dolichocybid mites described by V.D. Sevastianov and co-authors based on type and additional material.

## Materials and methods

The type materials of dolichocybid mites were loaned from the collection of the Odessa I. I. Mechnikov National University Museum of Zoology, Odessa, Ukraine. Additional material of

Received 26 September 2018
Accepted 29 October 2018
Published 09 November 2018

Corresponding author
Alexander A. Khaustov:
alkhaustov@mail.ru

Academic editor
Ekaterina Sidorchuk

DOI
10.24349/acarologia/20184295

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

Copyright

Khaustov A. A. and Trach V. A.

Distributed under
Creative Commons CC-BY 4.0

<footnote>ªTyumen State University, Tyumen, Russia.</footnote>

<footnote>ᵇOdessa I.I. Mechnikov National University, Odessa, Ukraine.</footnote>

<footnote>ᶜUkrainian I.I. Mechnikov Anti-Plague Research Institute, Odessa, Ukraine.</footnote>
<figure><img src="image_7.png" /><figcaption>**Figure 1** *Pavania riparia* Sevastianov, 1980, female: A – dorsum of the body, B – venter of the body. Legs omitted.</figcaption></figure>

*Pavania riparia* was collected by junior author on the dung beetle *Copris lunaris* and mounted in Hoyer's medium. One non type material slide of *P. riparia* from the collection of the junior author was remounted and several specimens were removed, dusted with gold and scanned with aid of a JEOL-JSM-6510LV SEM microscope.

Mite morphology was studied using a Carl Zeiss AxioImager A2 compound microscope
with phase contrast and DIC objectives. Photomicrographs were taken with an AxioCam ICc5 digital camera.

The terminology of the idiosoma and legs follows Lindquist (1986); the nomenclature of subcapitular setae follows Grandjean (1944). All measurements are given in micrometers (µm). For leg chaetotaxy the number of solenidia is given in parentheses.

## Taxonomy

**Family Dolichocybidae Mahunka, 1970a**

**Genus Pavania Lombardini, 1949**

**Type species: Pavania fusiformis Lombardini, 1949, by original designation.**

**Pavania riparia Sevastianov, 1980**

**Pavania riparia Sevastianov, 1980, p.1457, Figs 1-3.**

**(Figs 1-4)**

### Redescription

**Female (Figs 1-4)**

Body oval, weakly sclerotized. Length of idiosoma 125–130, width 74–86.

**Gnathosoma (Figs 4A-C)** — Gnathosomal capsule, excluding palps, almost round, its length 22, width 22–24. Dorsally with two pairs of weakly barbed and blunt-ended cheliceral setae (cha, chb). Setae cha 13–14 distinctly longer than chb 9–10. Dorsal median apodeme well developed. Postpalpal setae (pp) short, divided distally (Fig. 4C). Venter of gnathosoma with one pair of smooth, pointed subcapitular setae *m* 12. Palps with smooth setae *dFe* and *dGe* dorsolaterally, setae *dGe* 10–11 pointed, more than two times longer than blunt-ended

<figure><img src="image_8.png" /><figcaption>**Figure 2** *Pavania riparia* Sevastianov, 1980, female: A – right leg I in dorsal view, B – right leg II in dorsal view, C – right leg III in dorsal view, D – right leg IV in dorsal view.</figcaption></figure>
dFe 4. Palps ventrally with two solenidia (sol). Inner solenidion slightly shorter than outer one. Palp tibiotarsus with short, blunt-ended ventrolateral seta (probably tibial l") and distal eupathidium-like tiny seta (eup) (Fig. 4B). Palps terminate with well-developed tibial claw (cl) (Fig. 4A, B). Cheliceral stylets strong, curved. Pharynx thin-walled, elongate, with weak lateral projections.

**Idiosomal dorsum (Figs 1A, 3A)** — All dorsal shields with small sparsely distributed dimples (Fig. 3A). Prodorsal shield with three pairs of setae ($v_1, v_2, sc_2$) and a pair of clavate, barbed trichobothria $sc_1$ with rounded apex. All dorsal setae blunt-ended. Most of dorsal setae smooth or with very small barbs and only setae $h_1$ with distinct barbs. Tips of setae $h_2$ thickened into tiny clubs. Cupules *ia* on tergite D and *ip* on tergite EF small, round; other cupules not evident. Posterior margins of tergites C, D, and EF with several weak projections. Lengths of dorsal setae: $v_1$ 16, $v_2$ 6, $sc_2$ 30–32, $c_1$ 23–27, $c_2$ 18, *d* 17, *e* 10–12, *f* 17–18, $h_1$ 13–14, $h_2$ 54–63. Distances between setae: $v_1-v_1$ 16–17, $v_2-v_2$ 24–26, $sc_2-sc_2$ 28, $c_1-c_1$ 24–28, *d-d* 48–49, *e-e* 40–41, *f-f* 28–29, $h_1-h_1$ 10, $h_1-h_2$ 7–8.

**Idiosomal venter (Figs 1B, 3B, 4D)** — All ventral plates smooth. All ventral setae smooth; setae 2c pointed, other setae blunt-ended. Apodemes 1 (ap1) and apodemes 2 (ap2) well developed; prosternal apodeme not evident; apodemes 3 (ap3) and 4 (ap4) well developed. Coxal fields I-IV each with three pairs of setae. Lengths of ventral setae: 1a 6, 1b 7, 1c 6, 2a 10, 2b 6, 2c 19, 3a 9, 3b 8–9, 3c 10–11, 4a 8, 4b 8–9, 4c 9, *ag* 9–10, $g_1$ 4–5, $g_2$ 4, *ps* 9.

<figure><img src="image_9.png" /><figcaption>**Figure 3** DIC photomicrographs of *Pavania riparia* Sevastianov, 1980, female: A – dorsal view, B – ventral view.</figcaption></figure>
<figure><img src="image_10.png" /><figcaption>**Figure 5** *Pavania tadjikistanica* Sevastianov, 1980, female (holotype): A – dorsum of the body, B – venter of the body. Legs omitted.</figcaption></figure>

**Material examined**

Female holotype, and 10 female paratypes, slide No. D-T-2, Ukraine, vicinity of Odessa, coast of Kuyalnik Liman, on *Copris lunaris* (Coleoptera, Scarabaeidae), 22 May 1960, Sevastianov V.D. leg.; 10 females, Ukraine, Odessa Prov., Razdelnaya District, vicinity of settlement Kolontaevka (46°43' N, 30°18' E), on *C. lunaris*, 20 Sept. 2009, Trach V.A. leg.; 8 females,
Ukraine, Kherson Prov., Kalanchak District., vicinity of settlement Preobrazhenka (46°11' N, 33°36' E), on *C. lunaris*, 2-4 May 2011, Trach V.A. leg.

**Remarks**

The holotype and paratypes available for study are in poor condition and redescription is based on additional material collected by the junior author.

**Pavania tadjikistanica Sevastianov, 1980**

**Pavania tadjikistanica Sevastianov, 1980, p.1457, Figs 4-7.**
(Figs 5-7)

**Redescription**

**Female (Figs 5-7)**

The holotype and two paratypes available for this study are in rather bad condition (Fig. 7) and some tiny structures like cupules, dimples of dorsal plates and weak barbs on setae are not discernible. Length of idiosoma 120, width 80.

**Gnathosoma** — Gnathosomal capsule, excluding palps, almost round, its length 27, width 31. Dorsally with two pairs of blunt-ended cheliceral setae (cha, chb). Setae cha 15 distinctly longer than chb 10. Dorsal median apodeme well developed. Postpalpal setae (pp) short, peg-like. Venter of gnathosoma with one pair of pointed subcapitular setae m 9. Palps with smooth and pointed setae dFe and dGe dorsolaterally, setae dGe 11 almost twice longer than dFe 6. Palps ventrally with two solenidia. Inner solenidion much shorter than outer one. Palp tibiotarsus with short, blunt-ended ventrolateral seta (probably tibial l''). Palps terminated with well-developed tibial claw. Cheliceral stylets strong, falcate. Pharynx not discernible.

**Idiosomal dorsum (Figs 5A, 7A)** — Prodorsal shield with three pairs of setae ($v_1$, $v_2$, $sc_2$) and a pair of broken trichobothria $sc_1$. Setae $sc_2$ pointed, other dorsal setae blunt-ended. Tips of setae $h_2$ thickened into tiny clubs. Posterior margins of tergites without projections. Lengths of

<figure><img src="image_11.png" /><figcaption>**Figure 6** *Pavania tadjikistanica* Sevastianov, 1980, female (holotype): A – right leg I in dorsal view, B – right leg II in dorsal view, C – left leg III in dorsal view, D – left leg IV in dorsal view.</figcaption></figure>
dorsal setae: *v*₁ 14, *v*₂ 6, *sc*₂ 36, *c*₁ 20, *c*₂ 20, *d* 22, *e* 16, *f* 17, *h*₁ 13, *h*₂ 53. Distances between setae: *v*₁-*v*₁ 22, *v*₂-*v*₂ 29, *sc*₂-*sc*₂ 34, *c*₁-*c*₁ 24, *d*-*d* 34, *e*-*e* 43, *f*-*f* 29, *h*₁-*h*₁ 12, *h*₁-*h*₂ 6.

**Idiosomal venter (Figs 5B, 7B)** — All ventral setae blunt-ended. Ap1 and ap2 well developed; prosternal apodeme not evident; sejugal apodeme (apsej) weakly developed laterally; ap3 and ap4 well developed. Coxal fields I-IV each with three pairs of setae. Genital setae (*g*₁, *g*₂) very small, vestigial. Lengths of ventral setae: 1*a* 6, 1*b* 6, 1*c* 5, 2*a* 8, 2*b* 5, 2*c* 8, 3*a* 7, 3*b* 7, 3*c* 9, 4*a* 6, 4*b* 9, 4*c* 8, *ag* 7, *g*₁ 1, *g*₂ 1, *ps* 9.

**Legs (Fig. 6)** — Leg setation as in previous species. Leg I (Fig. 6A). Tarsus with two small claws and semioval empodium. Setae *l*' of femur, *l*' of genu, *k*, *l*' and *v*' of tibia blunt-ended; other leg setae (except eupathidia) pointed. Trochanter dorsally probably without spine-like projections. Lengths of solenidia *ω*₁ 6, *ω*₂ 3, *φ*₁ 6, *φ*₂ 5; solenidia *ω*₁, *φ*₂ and *ω*₂ weakly clavate, solenidion *φ*₁ distinctly clavate. Leg II (Fig. 6B). Tarsal claws simple, hooked; empodium large. Solenidia *ω* 5 and *φ* 3 weakly clavate. Trochanter probably without spine-like projections. All leg setae pointed. Leg III (Fig. 6C). Claws and empodium of same shape as on tarsus II. Seta *d* of femur blunt-ended, other leg setae pointed. Leg IV (Fig. 6D). Claws and empodium of same

<figure><img src="image_12.png" /><figcaption>**Figure 7** DIC photomicrographs of *Pavania tadjikistanica* Sevastianov, 1980, female (holotype): A – dorsal view, B – ventral view.</figcaption></figure>
<figure><img src="image_13.png" /><figcaption>**Figure 8** *Pavania protracta* Sevastianov, 1980, female (paratype): A – dorsum of the body, B – venter of the body. Legs omitted.</figcaption></figure>

shape as on tarsus III. Seta *d* of femur blunt-ended, other leg setae pointed.
**Male unknown.**

**Material examined**

Female holotype, and 2 female paratypes, slide No. T-D-3, Tadjikistan, vicinity of Dushanbe,
under elytra of *Onthophagus* sp. (Coleoptera, Scarabaeidae), 25 Aug. 1969, Ilyasov I.N. leg.
**Pavania protracta** Sevastianov, 1980

**Pavania protracta** Sevastianov, 1980, p.1459, Figs 8-10.
(Figs 8-10)

**Redescription**

**Female (Figs 8-10)**

Length of idiosoma 150, width 75.

**Gnathosoma** — Gnathosomal capsule, excluding palps, almost round, rather small, its length 18, width 19. Dorsally with two pairs of weakly barbed cheliceral setae (cha, chb). Setae cha 9 pointed, distinctly longer than blunt-ended chb 5. Dorsal median apodeme weakly developed. Postpalpal setae (pp) short, peg-like. Venter of gnathosoma with one pair of pointed and smooth subcapitular setae m 11. Palps with smooth setae dFe and dGe dorsolaterally, setae dGe 8 pointed, two times longer than blunt-ended dFe 4. Palps ventrally with two solenidia. Inner solenidion much shorter than outer one. Palp tibiotarsus with short, blunt-ended ventrolateral seta (probably tibial l"). Palps terminated with well-developed tibial claw. Cheliceral stylets relatively small, curved. Pharynx not discernible.

**Idiosomal dorsum (Figs 8A, 10A)** — All dorsal shields smooth. Prodorsal shield with three pairs of setae ($v_1, v_2, sc_2$) and a pair of clavate and distinctly barbed trichobothria $sc_1$. All dorsal setae weakly barbed. Setae $v_2$ blunt-ended, other dorsal setae pointed. Posterior margins of tergites without projections. Cupules not evident. Lengths of dorsal setae: $v_1$ 18, $v_2$ 7, $sc_2$ 28, $c_1$ 20, $c_2$ 26, $d$ 21, $e$ 20, $f$ 21, $h_1$ 16, $h_2$ 36. Distances between setae: $v_1-v_1$ 9, $v_2-v_2$ 18, $sc_2-sc_2$ 20, $c_1-c_1$ 21, $d-d$ 37, $e-e$ 38, $f-f$ 25, $h_1-h_1$ 12, $h_1-h_2$ 8.

**Idiosomal venter (Figs 8B, 10B)** — All ventral plates smooth. Setae 1a, 1b, 1c, 2b, 2c, and $g_1$ blunt-ended; other ventral setae pointed. Setae 1a, 1b, 1c, 2b, and 2c weakly barbed; other ventral setae smooth. Only ap2 and ap3 present, weakly developed; other apodemes not evident. Coxal fields I-IV each with three pairs of setae. Lengths of ventral setae: 1a 5, 1b 3,

<figure><img src="image_14.png" /><figcaption>**Figure 9** *Pavania protracta* Sevastianov, 1980, female (paratype): A – right leg I in dorsal view, B – right leg II in dorsal view, C – right leg III in dorsal view, D – right leg IV in dorsal view.</figcaption></figure>
1c 5, 2a 9, 2b 4, 2c 8, 3a 8, 3b 8, 3c 10, 4a 8, 4b 9, 4c 10, ag 8, g₁ 5, g₂ 9, ps 11.

**Legs (Fig. 9)**— Leg setation as in previous species. Leg I (Fig. 9A). Tarsus with two small claws and very short empodium. Setae v' of genu, k and v' of tibia blunt-ended; other leg setae (except eupathidia) pointed. Setae v'', (l) of femur, k of tibia, pl', (u) and all eupathidia of tarsus smooth; other leg setae weakly barbed. Trochanter dorsally with three spine-like projections; central projection distinctly smaller than lateral ones. Lengths of solenidia ω₁ 9, ω₂ 4, φ₁ 8, φ₂ 5; Solenidia ω₁ and φ₂ finger-shaped; solenidion ω₂ weakly clavate, solenidion φ₁ distinctly clavate. Leg II (Fig. 9B). Tarsal claws simple, hooked; empodium small, semioval. Solenidia ω 5 finger-shaped and φ 3 weakly clavate. Trochanter with one spine-like projection. All leg setae pointed and weakly barbed. Leg III (Fig. 9C). Claws and empodium of same shape as on tarsus II. Seta pv' of tarsus thickened, spiniform, blunt-ended; other leg setae pointed. Setae (pv) and tc'' of tarsus smooth; other leg setae weakly barbed. Leg IV (Fig. 9D). Claws and empodium of same shape as on tarsus III. Seta pv' of tarsus thickened, spiniform, blunt-ended; other leg setae pointed. Setae pv' and tc'' of tarsus smooth; other leg setae weakly barbed.

<figure><img src="image_15.png" /><figcaption>**Figure 10** DIC photomicrographs of *Pavania protracta* Sevastianov, 1980, female (holotype): A – dorsal view, B – ventral view.</figcaption></figure>
<figure><img src="image_16.png" /><figcaption>**Figure 11** *Pavania tahanae* Sevastianov and Abo-Korah, 1985, female (holotype): A – dorsum of the body, B – venter of the body. Legs omitted.</figcaption></figure>

Male unknown.

**Material examined**

Female holotype, and 1 female paratype, slide No. T-D-4, Russia, Tatarstan, vicinity of Kazan,
soil under maize, 27 Aug. 1968, Artemjeva T.I. leg.

**Remarks**

The holotype is in rather bad condition (Fig. 13). One female paratype was remounted and description is based on this paratype specimen.
**Pavania tahanae Sevastianov and Abo-Korah, 1985**

**Pavania tahanae Sevastianov and Abo-Korah, 1985, p.35, Figs 1–4.**
(Figs 11–13)

**Redescription**

**Female (Figs 11–13)**

Length of idiosoma 135, width 80.

**Gnathosoma** — Gnathosomal capsule, excluding palps, almost round, its length 19, width 20. Dorsally with two pairs of smooth and blunt-ended cheliceral setae (cha, chb). Setae cha 14 distinctly longer than chb 11. Dorsal median apodeme indistinct. Postpalpal setae (pp) short, peg-like. Venter of gnathosoma with one pair of pointed and smooth subcapitular setae m 11. Palps with smooth setae dFe and dGe dorsolaterally, setae dGe 12 pointed, almost two times longer than blunt-ended dFe 7. Palps ventrally with two subequal solenidia. Palp tibiotarsus with short, blunt-ended ventrolateral seta (probably tibial l"). Palps terminated with well-developed tibial claw. Cheliceral stylets rather small, curved. Pharynx not discernible.

**Idiosomal dorsum (Figs 11A, 13A)** — All dorsal shields smooth. Prodorsal shield with three pairs of setae ($v_1, v_2, sc_2$) and a pair of clavate and weakly barbed trichobothria $sc_1$. Setae e weakly barbed, other dorsal setae apparently smooth. Setae $v_1$ and $v_2$ blunt-ended, other dorsal setae pointed. Posterior margins of tergites without projections. Cupules not evident. Lengths of dorsal setae: $v_1$ 18, $v_2$ 9, $sc_2$ 32, $c_1$ 22, $c_2$ 29, d 23, e 25, f 28, $h_1$ 18, $h_2$ 35. Distances between setae: $v_1-v_1$ 11, $v_2-v_2$ 20, $sc_2-sc_2$ 28, $c_1-c_1$ 37, d-d 53, e-e 46, f-f 42, $h_1-h_1$ 11, $h_1-h_2$ 9.

**Idiosomal venter (Figs 11B, 13B)** — All ventral plates smooth. All ventral setae smooth. Setae $g_1$ and $g_2$ blunt-ended, other ventral setae pointed. Only ap2 and ap3 clearly discernible; other apodemes not evident. Coxal fields I-V each with three pairs of setae. Lengths of ventral setae: 1a 7, 1b 8, 1c 6, 2a 13, 2b 6, 2c 15, 3a 9, 3b 7, 3c 13, 4a 7, 4b 8, 4c 8, ag 11, $g_1$ 4, $g_2$ 4, ps 10.

<figure><img src="image_17.png" /><figcaption>**Figure 12** *Pavania tahanae* Sevastianov and Abo-Korah, 1985, female (holotype): A – right leg I in dorsal view, B – right leg II in dorsal view, C – right leg III in dorsal view, D – right leg IV in dorsal view.</figcaption></figure>
**Legs (Fig. 12)** — Leg setation as in previous species. Leg I (Fig. 12A). Tarsus with two small claws, empodium not evident. All leg setae smooth. Seta *k* of tibia and all eupathidia of tarsus blunt-ended; other leg setae pointed. Trochanter dorsally without spine-like projections. Lengths of solenidia ω₁ 9, ω₂ 5, φ₁ 8, φ₂ 3; solenidion ω₁ finger-shaped; solenidion ω₂ baculiform, solenidia φ₁ and φ₂ distinctly clavate. Leg II (Fig. 12B). Tarsal claws simple, hooked; empodium very small, semioval. Solenidia ω 8 finger-shaped and φ 2 weakly clavate. Trochanter without spine-like projections. All leg setae pointed and smooth. Leg III (Fig. 12C). Claws and empodium of same shape as on tarsus II. Solenidion φ 2 weakly clavate. All leg setae pointed and smooth. Leg IV (Fig. 12D). Claws and empodium of same shape as on tarsus II. Solenidion φ 2 weakly clavate. All leg setae pointed and smooth.

Male unknown.

<figure><img src="image_18.png" /><figcaption>**Figure 13** DIC photomicrographs of *Pavania tahanae* Sevastianov and Abo-Korah, 1985, female (holotype): A – dorsal view, B – ventral view.</figcaption></figure>
Material examined

Female holotype, slide No. 704, Egypt, vicinity of Shibin El Kom, soil under cotton, 1 Sept. 1980, Abo-Korah leg.

Key to world species of Pavania (based on Khaustov & Frolov 2018b)

<table><tbody><tr><td>1. Setae <i>sc</i><sub>1</sub> absent</td><td>2</td></tr><tr><td>— Setae <i>sc</i><sub>1</sub> present</td><td>5</td></tr><tr><td>2. Setae 1c and 2c present</td><td>3</td></tr><tr><td>— Setae 1c and 2c absent</td><td>P. neotropica Khaustov and Frolov, 2017 (Brazil)</td></tr><tr><td>3. Setae <i>v</i><sub>1</sub> shorter than distance between their bases; setae <i>cha</i> less than three times longer than <i>chb</i>; setae <i>e</i> never longer than <i>f</i>; setae <i>h</i><sub>2</sub> at most seven times longer than <i>h</i><sub>1</sub></td><td>4</td></tr><tr><td>— Setae <i>v</i><sub>1</sub> longer than distance between their bases; setae <i>cha</i> three times longer than <i>chb</i>; setae <i>e</i> longer than <i>f</i>; setae <i>h</i><sub>2</sub> 15 times longer than <i>h</i><sub>1</sub></td><td>P. gymnopleuri Hajiqanbar and Khaustov, 2010 (Iran)</td></tr><tr><td>4. Genu I with one seta (<i>v</i>'); dorsal idiosomal setae smooth; setae <i>c</i><sub>1</sub> longer than <i>c</i><sub>2</sub>; setae <i>c</i><sub>1</sub> and <i>d</i> pointed</td><td>P. sabzevarensis Hajiqanbar and Khaustov, 2010 (Iran)</td></tr><tr><td>— Genu I with two setae (<i>v</i>', <i>l</i>'); dorsal idiosomal setae weakly barbed; setae <i>c</i><sub>2</sub> longer than <i>c</i><sub>1</sub>; setae <i>c</i><sub>1</sub> and <i>d</i> distinctly blunt-ended</td><td>P. onthophagi Hajiqanbar and Khaustov, 2010 (Iran)</td></tr><tr><td>5. Setae <i>sc</i><sub>1</sub> capitate</td><td>6</td></tr><tr><td>— Setae <i>sc</i><sub>1</sub> seta-like</td><td>P. setiformis Loghmani and Hajiqanbar, 2013 (Iran)</td></tr><tr><td>6. Setae (<i>u</i>) and (<i>pv</i>) of tarsus I not lanceolate</td><td>7</td></tr><tr><td>— Setae (<i>u</i>) and (<i>pv</i>) of tarsus I lanceolate</td><td>P. lanceolata Bahramian and Hajiqanbar, 2015 (Iran)</td></tr><tr><td>7. Coxal fields II with 3 pairs of setae</td><td>8</td></tr><tr><td>— Coxal fields II with 2 pairs of setae</td><td>P. equisetosa Mahunka, 1975 (Ghana)</td></tr><tr><td>8. Empodium on tarsi II-IV small, not reaching beyond tips of claws, seta <i>pv</i>' on tarsi III and IV simple</td><td>9</td></tr><tr><td>— Empodium on tarsi II-IV large, reaching beyond tips of claws</td><td>10</td></tr><tr><td>9. Seta <i>pv</i>' on tarsi III and IV thickened, spiniform and blunt-ended, solenidia on tibiae III and IV absent</td><td>P. rotracta Sevastianov, 1980 (Russia)</td></tr><tr><td>— Seta <i>pv</i>' on tarsi III and IV simple, solenidia on tibiae III and IV present</td><td>P. tahanae Sevastianov and Abo-Korah, 1985 (Egypt)</td></tr><tr><td>10. Setae <i>h</i><sub>2</sub> less than 3.5 times longer than <i>h</i><sub>1</sub></td><td>11</td></tr><tr><td>— Setae <i>h</i><sub>2</sub> more than 3.5 times longer than <i>h</i><sub>1</sub></td><td>15</td></tr><tr><td>11. Setae <i>c</i><sub>1</sub> never reaching beyond bases of setae <i>f</i>; setae <i>c</i><sub>1</sub> shorter than <i>h</i><sub>2</sub>; setae <i>d</i> shorter than <i>h</i><sub>2</sub></td><td>12</td></tr><tr><td>— Setae <i>c</i><sub>1</sub> reaching beyond bases of setae <i>f</i>; setae <i>c</i><sub>1</sub> longer than <i>h</i><sub>2</sub>; setae <i>d</i> and <i>h</i><sub>2</sub> subequal</td><td>P. perhirsuta Mahunka, 1973 (Ghana)</td></tr><tr><td>12. Setae <i>sc</i><sub>2</sub> subequal to distance between their bases</td><td>13</td></tr><tr><td>— Setae <i>sc</i><sub>2</sub> distinctly longer than distance between their bases</td><td></td></tr></tbody></table>
<figure><img src="image_19.png" /><figcaption>**Figure 14** *Dolichocybe firjusae* Sevastianov and Chydyrov, 1994, female (holotype): A – dorsum of the body, B – venter of the body. Legs omitted.</figcaption></figure>
<figure><img src="image_20.png" /><figcaption>**Figure 15** *Dolichocybe firjusae* Sevastianov and Chydyrov, 1994, female (holotype): A – left leg I in dorsal view, B – left leg II in dorsal view, C – right leg III in dorsal view, D – right leg IV in dorsal view.</figcaption></figure>

Body elongate, weakly sclerotized. Length of idiosoma 180, width 77.

**Gnathosoma** — Gnathosomal capsule, excluding palps, almost oval, its length 20, width 15. Dorsally with two pairs of smooth and pointed cheliceral setae (cha, chb). Setae cha 12 distinctly longer than chb 6. Dorsal median apodeme indistinct. Postpalpal setae (pp) rod-like. Venter of gnathosoma with one pair of smooth, pointed subcapitular setae m 17. Palps freely articulated to gnathosomal capsule, with smooth and pointed setae dFe and dGe dorsolaterally, setae dGe 12 pointed, more than two times longer than blunt-ended dFe 5. Palps ventrally with two subequal solenidia. Palp tibiotarsus with short ventrolateral seta (probably tibial l") and distal eupathidium-like tiny seta. Palps terminated with well-developed tibial claw. Cheliceral stylets small, indistinct. Pharynx thin-walled, elongate, with weak lateral projections.

**Idiosomal dorsum** (Figs 14A, 16) — All dorsal shields smooth. Prodorsal shield with three pairs of setae (v₁, v₂, sc₂) and a pair of clavate, smooth trichobothria sc₁ with rounded apex; setae v₁ and v₂ located on the same transverse line. Setae v₁, v₂, and c₁ blunt-ended, other dorsal setae pointed. Setae h₁ and h₂ smooth, other dorsal setae weakly barbed. Cupules ia on tergite D and ip on tergite EF small, round; other cupules not evident. Lengths of dorsal setae: v₁ 14, v₂ 23, sc₂ 44, c₁ 13, c₂ 25, d 18, e 19, f 25, h₁ 13, h₂ 76. Distances between setae: v₁–v₁ 10, v₂–v₂ 22, sc₂–sc₂ 20, c₁–c₁ 30, d–d 36, e–e 36, f–f 30, h₁–h₁ 13, h₁–h₂ 5.

**Idiosomal venter** (Figs 14B, 16) — All ventral plates smooth. Setae 3a, 3c, and ag weakly barbed, other ventral setae smooth; setae g₁ and g₂ blunt-ended, other ventral setae pointed. Only ap3 and ap4 weakly developed, other apodemes indistinct. Coxal fields I-IV each with three pairs of setae. Lengths of ventral setae: 1a 13, 1b 11, 1c 10, 2a 17, 2b 11, 2c 23, 3a 12, 3b 13, 3c 15, 4a 9, 4b 10, 4c 12, ag 14, g₁ 3, g₂ 4, ps 7.
All leg setae pointed. Seta *d* of tibia and all tarsal setae smooth, other leg setae weakly barbed.
**Male unknown.**

**Material examined**

Female holotype, slide No. 908, Turkmenistan, Chardzhou Prov., vicinity of settlement Karabekaul, soil on cotton field, 24 Sept. 1982, Chydyrov P.R. leg.

**Discussion**

The main differences between the genera *Dolichocybe* and *Pavania* according to Cross (1965) are the shape of cheliceral stylets (falcate and well developed in *Pavania* and small, indistinct in *Dolichocybe*) and position of palps (palps arising ventrolaterally in *Pavania* and laterally in *Dolichocybe*). In fact, the position of palps is rather uniform in all dolichocybid mites and can not be used as a generic-level character. The shape and size of stylets are also highly variable. In typical *Pavania* (like most species associated with dung beetles), stylets are really large and falcate. In typical *Dolichocybe* (*D. keiferi*, *D. subcorticalis* Khaustov, 2006, *D. sibiriensis* Khaustov, 2017, *D. firjusae*) stylets are small and indistinct. However, at least in *Pavania protracta* and *P. tahanae*, size of the cheliceral stylets is intermediate between typical for *Pavania* and for *Dolichocybe*. Therefore, the shape of cheliceral stylets is a vague character, difficult to use in separation of the genera *Dolichocybe* and *Pavania*. Rahiminejad *et al*. (2011) redefined the genus *Dolichocybe* and used the following characters to separate *Dolichocybe* from *Pavania*: “gnathosoma apparently longer than wide; chelicerae small and indistinct; tarsus I with 10 or 11 setae; tibiae III and IV with one minute solenidion each; with deep constriction between propodosoma and hysterosoma separated by soft and transversely striated cuticle” (in *Dolichocybe*); “gnathosoma almost as long as wide; chelicerae large and distinct; tarsus I with 11 setae; tibiae III and IV with no solenidia; without deep constriction between propodosoma and hysterosoma” (in *Pavania*). Based on our study, the number of setae on tarsus I is the same in *Pavania* and *Dolichocybe* and can not be used in separation of these genera. The presence of solenidia on tibiae III and IV in *Dolichocybe* is also problematic character. In fact, solenidia on tibiae III and IV are apparently absent in *D. keiferi* (type species) (Cross 1965), *D. hippocastani* Rack, 1967, *D. picea* Rack, 1967 (Rack 1967), and absent in *D. subcorticalis*, *D. sibiriensis* (Khaustov 2006, 2017) and *D. firjusae* (present study). Potentially solenidia on tibiae III and IV are present in *Pavania brasiliensis* Mahunka, 1970, at least Mahunka (1970) depicted a tiny solenidion on the tibia III. Therefore, the presence of solenidia on tibiae III and IV also can not be used for separation of these genera. It is also difficult to understand the extent of constriction between propodosomal and hysterosoma.

Two redescribed herein species, *Pavania protracta* and *P. tahanae*, have intermediate characters between *Dolichocybe* and *Pavania*: body shape is similar to that in *Pavania* (not strongly elongate), gnathosoma is rather small (as in *Dolichocybe*), cheliceral stylets are of medium size (intermediate), constriction between propodosomal and hysterosoma is not deep (as in *Pavania*). In addition, both of these species have unusually short — considerably shorter than tarsal claws — empodia on tarsi II-IV. *Pavania tahanae* also has solenidia on tibiae III and IV (like some “*Dolichocybe*”). Both species were collected in soil (not phoretic on insects) and potentially could represent a non-phoretic form of *Pavania/Dolichocybe* complex. At present female dimorphism has been evident only in the genus *Formicomotes* Sevastianov, 1980. Non-phoretic form of *F. heteromorphus* Magowski, 1988 has much shorter empodia than phoretic form (Magowski 1988). In this paper, we retain all species in the genera in which they were originally described. The redefinitions of the genera *Dolichocybe* and *Pavania* are necessary based on redescription of their type species and more discoveries of new species in these two close genera (particularly *Dolichocybe*) are required (phoretic or in soil) to see what characters are more variable.
Acknowledgements

The authors are grateful to Dr. Vladimir A. Lobkov and Yuri V. Suvorov, of the Odessa I. I. Mechnikov National University Museum of Zoology (Odessa, Ukraine) for access to the type material of V.D. Sevastianov.

The study was supported by the Russian Foundation for Basic Research (RFBR), research project No. 18-04-01092A.

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