# Taxonomic comments on some species of the genus Eustigmaeus Berlese (Trombidiformes: Stigmaeidae), with a revised checklist of Eustigmaeus and descriptions of two new species from Türkiye

Salih Doğan, Sibel Doğan

► To cite this version:

Salih Doğan, Sibel Doğan. Taxonomic comments on some species of the genus Eustigmaeus Berlese (Trombidiformes: Stigmaeidae), with a revised checklist of Eustigmaeus and descriptions of two new species from Türkiye. Acarologia, 2024, 64 (3), pp.711-732. 10.24349/e3fo-3y3r. hal-04625569

HAL Id: hal-04625569

https://hal.science/hal-04625569v1

Submitted on 26 Jun 2024

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# Taxonomic comments on some species of the genus *Eustigmaeus* Berlese (Trombidiformes: Stigmaeidae), with a revised checklist of *Eustigmaeus* and descriptions of two new species from Türkiye

Salih Doğan IDª, Sibel Doğan IDᵇ

## Original research

### ABSTRACT

Two new species of *Eustigmaeus* Berlese (Acari: Stigmaeidae), *E. incisus* n. sp. and *E. paraincisus* n. sp. are described and illustrated from Türkiye. This paper presents compile checklist of the valid *Eustigmaeus* species. *Eustigmaeus sphagneticolus* Cooreman stat. nov., originally described as a subspecies of *E. ottavii* by Cooreman in 1943, is regarded as a separate species. *Eustigmaeus aminiae* (Nazari and Khanjani) n. comb.. and *E. indiscretus* (Dönel and Doğan) n. comb.. (from Ledermuelleriopsis Willmann) have a partially detached hysterosomal shield. Currently, we consider that these species belong more comfortably in *Eustigmaeus* than in *Ledermuelleriopsis*, where they are considered transitional species between the genera *Eustigmaeus* and *Ledermuelleriopsis*.

**Keywords** Acari; description; new combination; new rank; taxonomy; species list

**Zoobank** http://zoobank.org/4C4816C3-0090-46EE-ABB4-09628294D8F2

## Introduction

The family Stigmaeidae currently comprises over 640 species from 33 recognised genera (Fan et al. 2016, 2019; Beron, 2022; Khaustov et al. 2023; Doğan et al. 2024b). Of the Stigmaeidae, *Eustigmaeus* Berlese is one of the largest genera. There are 140 species (Supplementary Table 1) of this genus worldwide (Fan 2005; Fan et al. 2016, 2019; Beron 2020, 2022; Bizarro et al. 2020; Khaustov 2021a,b Khaustov et al. 2023; Bagheri and Mohammad-Doustaresharaf 2024), of which 33 (including two species described here) have been documented from Türkiye (Doğan, 2007, 2019; Doğan and Doğan 2020; Pekağırbaş et al. 2023; Doğan et al. 2024b).

The majority of *Eustigmaeus* are free-living predators; however, six-identified and one-unidentified species of *Eustigmaeus* are parasitic on sand flies as evidenced by the feeding scars on the hosts' bodies (*Pekağırbaş et al. 2023*). There is no obvious indication of host haemolymph in the digestive tracts of these mites, and it is unknown how they affect the host flies (*Mortazavi et al. 2018*).

We depict and describe two new species of *Eustigmaeus* from Türkiye: *E. incisus* n. sp. and *E. paraincisus* n. sp. A compendium of the valid species of *Eustigmaeus* is included in this paper. *Eustigmaeus sphagneticolus* Cooreman n. stat. described as a subspecies, *E. ottavii* sphagneticola, by Cooreman (1943) has been elevated to species level.

Received 15 March 2024

Accepted 08 May 2024

Published 27 May 2024

Corresponding author

Salih Doğan ID:

salihdogan@erzincan.edu.tr

Academic editor

Faraji, Farid

https://doi.org/10.24349/e3fo-3y3r

ISSN 0044-586X (print)

ISSN 2107-7207 (electronic)

<figure><img src="image_2.png" /></figure>

Doğan S. and Doğan S.

Licensed under

Creative Commons CC-BY 4.0

<footnote>ªDepartment of Biology, Faculty of Arts and Sciences, Erzincan Binali Yıldırım University, Erzincan, Türkiye.</footnote>

<footnote>ᵇVocational School of Health Services, Erzincan Binali Yıldırım University, Erzincan, Türkiye.</footnote>
**Material and methods**

The *Eustigmaeus* species list (Supplementary Table 1) has been created by combining from the catalogues and checklists (Fan 2005; Fan *et al.* 2016, 2019; Stathakis *et al.* 2016; Beron 2020, 2022) and updated from recent publications (Bizarro *et al.* 2020; Khaustov 2021a,b; Khaustov *et al.* 2023; Bagheri and Mohammad-Doustaresharaf 2024). Each species' current name, converted in accordance with the masculine gender of the genus, with its authorship has been listed together with a bibliographic reference that contains the original description of the species along with its original name.

Using Berlese-Tullgren funnels, the mite specimens were extracted, cleared in 60% lactic acid, and mounted in Hoyer's medium on microscope slides (Fan and Zhang 2005; Walter and Krantz 2009). A Leica DM 4000B phase contrast equipped with a drawing tube was used to make the illustrations. The nomenclature of the idiosomal setation follows that proposed by Grandjean (1939) as adapted for Prostigmata by Kethley (1990). The terminologies of subcapitular, palp and the leg setations follow those proposed by Grandjean (1944, 1946). All measurements and the scale bars in the figures were taken in micrometers (µm) with the aid of the Leica Application Suite (LAS) Software Version 4.8. The holotype measurements are followed by the range of paratypes in parenthesis. The mite materials are deposited in EBYU (Acarology Laboratory of Erzincan Binali Yıldırım University, Erzincan, Türkiye).

**Results**

**Family Stigmaeidae Oudemans, 1931**

**Genus *Eustigmaeus* Berlese, 1910**

*Type species* — *Stigmaeus kermesinus* Koch, 1841 by original designation.

***Eustigmaeus incisus*** *n.* sp.

Zoobank: 614493EF-BA61-42A1-9AC2-46E28568B69B

(Figures 1–4, 5A, C and 6–8)

**Diagnosis**

**Female** Dorsal shields ornamented with pits, 10–28 vacuoles present in pits; dorsal setae homeomorphic, pudgy and resembling thorny bush; humeral setae $c_2$ similar in shape to dorsal body setae; lateral incision of hysterosomal shield; a pair of almost rounded patterned areas located anterolaterad aggenital shield.

**Description**

**Female** (n=16 examined, n=6 measured) (Figs 1–4, 5A)

Length of idiosoma 319 (289–317), width 230 (209–220).

**Dorsum** — (Fig. 1). Dorsal shields ornamented with pits, 10–28 vacuoles present in pits. Fossettes on dorsal shields as illustrated. Propodosomal shield with four pairs of setae: *vi*, *ve*, *sci* and *sce*, and a pair of ocelli, 7 (7–11) in diameter. Lateral margins of hysterosomal shield incised, and this shield with six pairs of setae: $c_1$, $d_1$, $d_2$, $e_1$, $e_2$ and $f_1$. Suranal shield with two pairs of setae: $h_1$ and $h_2$. Dorsal setae pudgy and bushy. Lengths and distances of dorsal setae as follows: *vi* 18 (16–18), *ve* 18 (16–19), *sci* 18 (13–17), *sce* 17 (16–17), $c_1$ 19 (15–18), $d_1$ 16 (15–17), $d_2$ 16 (15–17), $e_1$ 19 (18–19), $e_2$ 16 (15–18), $f_1$ 27 (24–26), $h_1$ 21 (21–23), $h_2$ 20 (21–23), *vi–vi* 33 (31–40), *ve–ve* 85 (72–88), *vi–ve* 41 (36–42), *sci–sci* 137 (122–135), *ve–sci* 30 (26–30), *sce–sce* 181 (152–177), *sci–sce* 40 (28–39), $c_1-c_1$ 76 (58–77), $d_2-d_2$ 186 (164–189), $c_1-d_1$ 57 (50–56), $c_1-d_2$ 58 (51–56), $d_1-d_1$ 68 (60–67), $d_2-d_1$ 66 (52–67), $e_2-e_2$ 143 (129–142), $e_1-e_1$ 98 (53–99), $e_2-e_1$ 42 (36–43), $f_1-f_1$ 57 (52–59), $e_1-f_1$ 41 (39–41), $e_2-f_1$ 78 (72–83), $h_1-h_1$ 31 (26–38), $h_2-h_2$ 72 (62–76), and $h_1-h_2$ 17 (19–21).
<figure><img src="image_3.png" /><figcaption>**Figure 1** *Eustigmaeus incisus* n. sp. (female) – Dorsal view. Scale bar: 100.</figcaption></figure>

**Venter** — (Fig. 2). Coxisternal shields undivided, and with three pairs of smooth intercoxal setae: 1a, 3a and 4a. Lengths and distances of these setae: 1a 9 (9–12), 3a 11 (9–15), 4a 8 (8–11), 1a–1a 25 (21–25), 3a–3a 37 (32–35), and 4a–4a 24 (22–25). Aggenital shield with three pairs of setae: *ag*₁, *ag*₂ and *ag*₃. Anogenital shields with three pairs of pseudanal setae: *ps*₁, *ps*₂ and *ps*₃. Lengths of these setae: *ag*₁ 7 (6–8), *ag*₂ 8 (7–8), *ag*₃ 10 (8–10), *ps*₁ 13 (12–16), *ps*₂ 12 (11–14), and *ps*₃ 11 (10–12). Aggenital and pseudanal setae slightly barbed, *ps*₁ thicker than other pseudanal setae, and distinctly barbed. Patterns on humeral shields similar to those of dorsal shields. Patterns on coxisternal, aggenital, pseudanal and suranal shields almost similar in shape as in Figure 2. Ovoid patterned area located between coxae III and IV and a pair of almost rounded patterned areas located anterolaterally on aggenital shield. Indentations and folds on anterior part of humeral shields present, but indistinguishable in some paratypes, setae c₂ 17 (17–19), similar in shape to dorsal setae.

**Legs** — (Figs 3 and 4). Faintly punctuated. Setal formulae of legs I–IV: coxae 2–2–2–2, trochanters 1–1–2–1, femora 6–4–3–2, genua 3(+1κ)–3(+1κ)–1–1, tibiae 5(+1φ+1φρ)–5(+1φρ)–5(+1φρ)–5(+1φρ), tarsi 13(+1ω)–9(+1ω)–7(+1ω)–7.

Leg I (Fig. 3A). 150 (147–171). Leg setation: Co 2 (1b, 1c) and needle-like leg supracoxal setae (el), Tr 1 (v'), Fe 6 (d, l', l'', v', v'', bv''), Ge 3 (d, l', l'') + 1 κ, Ti 5 (d, l', l'', v', v'') + 1 solenidion φ + 1 solenidion φρ, Ta 13 (p'ξ, p''ξ, tc'ξ, tc''ξ, ft'ξ, ft''ξ, u', u'', a', a'', pl', pl'', vs) +
<figure><img src="image_4.png" /><figcaption>**Figure 2** *Eustigmaeus incisus* n. sp. (female) – Ventral view. Scale bar: 100.</figcaption></figure>

1 solenidion ω, 17 (16–18). Setae *d* and *l''* on femur, *d* on genu, *d* on tibia distinctly barbed; setae *l', v', v''* on femur, *l'* on genu, *l''* on tibia pointed and smooth; setae *tc'ξ, tc"ξ, ft'ξ, ft"ξ, p'ξ, p"ξ* on tarsus smooth and weakly blunt-tipped; other setae on leg I pointed and barbed.

Leg II (Fig. 3B). 117 (113–131). Leg setation: Co 2 (2b, 2c), Tr 1 (*v*'), Fe 4 (*d*, *l'', l''*, *bv''*), Ge 3 (*d*, *l'', l''*) + 1 κ, Ti 5 (*d*, *l'', l''*, *v'', v''*) + 1 solenidion φρ, Ta 9 (*p'ξ*, *tc'ξ*, *tc''*, *u'*, *u''*, *a'*, *a''*, *pl'*, *vs*) + 1 solenidion ω, 12 (8–13). Setae *d* and *l''* on femur, *d* on genu, *d* on tibia distinctly barbed; setae *l'* on femur and genu, *l''* on tibia pointed and smooth; setae *p'ξ* and *tc'ξ* on tarsus smooth and weakly blunt-tipped; other setae on leg II pointed and barbed.

Leg III (Fig. 4A). 114 (108–130). Leg setation: Co 2 (3b, 3c), Tr 2 (*l'', v*'), Fe 3 (*d*, *l'', ev*'), Ge 1 (*d*), Ti 5 (*d*, *l'', l''*, *v'', v''*) + 1 solenidion φρ, Ta 7 (*tc'*, *tc''*, *u'*, *u''*, *a'*, *a''*, *vs*) + 1 solenidion ω, 3 (3–3). Seta *l'* on femur, setae *d* on genu and tibia distinctly barbed; *ev'* on femur, *l''* on tibia pointed and smooth; other setae on leg III pointed and barbed.

Leg IV (Fig. 4B). 146 (134–148). Leg setation: Co 2 (4b, 4c), Tr 1 (*v*'), Fe 2 (*d*, *ev*'), Ge 1 (*d*), Ti 5 (*d*, *l'', l''*, *v'', v''*) + 1 solenidion φρ, Ta 7 (*tc'*, *tc''*, *u'*, *u''*, *a'*, *a''*, *vs*), lacking solenidion ω. Setae *d* on femur, genu and tibia strongly serrate; other setae pointed and barbed.
<figure><img src="image_5.png" /><figcaption>**Figure 3** *Eustigmaeus incisus* n. sp. (female) – A – Leg I, B – Leg II. Scale bar: 40.</figcaption></figure>

**Gnathosoma** — (Fig. 5C). Length of subcapitulum 71 (63–71). Subcapitulum faintly punctuated. Length of chelicerae 92 (78–88). Palp faintly punctuated, length of palp 95 (85–93). Lengths and distances between subcapitular setae: *m* 11 (11–13), *n* 10 (9–11), *m–m* 20 (18–20), *n–n* 23 (19–26), and *m–n* 13 (10–12). Number of setae on palp segments: Tr 0, Fe 3 (*d*, *l'*, *v''*), Ge 2 (*d*, *l''*), Ti 3 (*d*, *l'*, *l''*) +1 claw, Ta 8 (fused eupathidia *ul'ξ*, *ul''ξ* and *sulξ*, eupathidion *acmξ*, *ba*, *bp*, *lp*, *va*) + 1 solenidion ω. All setae on palp femur barbed, other palp setae smooth. Palp supracoxal setae (*ep*) needle-like. Palp tibial seta *l'* paw-like, not seta-like.

**Male** (n=2) (Figs 6–8)

Idiosoma oval, much smaller than female. Length of idiosoma 261–269, width 168–173.

**Dorsum** — (Fig. 6A). Dorsal ornamentations as in female. Fossettes on dorsal shields as illustrated. Eyes 8 in diameter. Hysterosomal shield completely divided into two shields; anterior shield with three pairs of setae *c*₁, *d*₁, and *d*₂; posterior shield with three pairs of setae *e*₁, *e*₂, and *f*₁. Suranal shield located dorsally. Dorsal setae as in female except setae *e*₁ nearly smooth, setae *f*₁ barbed towards the tip, and setae *e*₂, *h*₁ and *h*₂ weakly barbed. Lengths and
<figure><img src="image_6.png" /><figcaption>**Figure 4** *Eustigmaeus incisus* n. sp. (female) – A – Leg III, B – Leg IV. Scale bar: 40.</figcaption></figure>

distances of dorsal setae as follows: *vi* 14, *ve* 13, *sci* 10–11, *sce* 11–12, *c*₁ 9–11, *d*₁ 11, *d*₂ 11–12, *e*₁ 10, *e*₂ 10–11, *f*₁ 31–32, *h*₁ 18–20, *h*₂ 21–26, *vi*-*vi* 21–26, *ve*-*ve* 63–64, *vi*-*ve* 33, *sci*-*sci* 102–106, *ve*-*sci* 23, *sce*-*sce* 129–133, *sci*-*sce* 28–33, *c*₁-*c*₁ 52–54, *c*₂-*c*₂ 146–153, *d*₂-*d*₂ 133–136, *c*₁-*d*₁ 45–46, *c*₁-*d*₂ 43–47, *d*₁-*d*₁ 44–45, *d*₂-*d*₁ 49–50, *e*₂-*e*₂ 98–99, *e*₁-*e*₁ 58–64, *e*₂-*e*₁ 21–26, *f*₁-*f*₁ 43–50, *e*₁-*f*₁ 20–23, *e*₂-*f*₁ 44–49, *h*₁-*h*₁ 20–23, *h*₂-*h*₂ 39–42, and *h*₁-*h*₂ 7–8.

**Venter** — (Fig. 6B). Coxisternal shields fused. Lengths and distances of intercoxal setae: *1a* 8–11, *3a* 10–14, *4a* 9–12, *1a*–*1a* ?–21, *3a*–*3a* 24–26, and *4a*–*4a* 15–16. Lengths of aggenital and pseudanal setae: *ag*₁ 8–11, *ag*₂ 10–14, *ps*₁ 1–2, *ps*₂ 3–4 and *ps*₃ 7. Pseudanal setae *ps*₁ and *ps*₂ spine-like. Aggenital and pseudanal setae smooth. Setae *c*₂ barbed and 15 long. Patterns on coxisternal, aggenital and humeral shields similar to those of dorsal shields. Aedeagus prominent and partly protruding.

**Legs** — (Figs 7 and 8). Length of legs, from leg I to leg IV respectively: 163–167, 136–139, 127–131, and 148–152. Counts of setae and solenidia on legs I–IV as in female, except enlarged solenidion (ω♂) on all tarsi present. Setae *d* on femora of legs I, II, and IV and setae *d* on genu and tibia of leg IV barbed; all other setae smooth.

*Gnathosoma*. As in female. Subcapitulum 57–60 long. Length of chelicerae 74–76, length of palp 79–81. Lengths and distances between subcapitular setae: *m* 11–12, *n* 9–10, *m–m* 16–17, *n–n* 18–19, and *m–n* 8–9.

**Type material**

Holotype: ♀ from moss, Acemoğlu Gorge, the Karasu Valley, TÜRKİYE, 39°36'33" N 39°9'13" E, 1140 m a.s.l., 29 October 2022, col. Salih Doğan. Paratypes: 1♀ and 1♂, same data
<figure><img src="image_7.png" /><figcaption>**Figure 5** A,B – Dorsal setae *c*₂ and *vi* in females; C,D – Venter of gnathosoma in females – A,C – *Eustigmaeus incisus* *n. sp*., B,D – *Eustigmaeus paraincisus* *n. sp*. Scale bar: 40.</figcaption></figure>

as holotype; 5 ♀♀ from oak litter, 39°36′32″ N 39°09′05″ E, 1120 m a.s.l., 27 January 2023; 3 ♀♀ from soil and 1 ♀ from oak litter, 39°36′16″ N 39°9′21″ E, 1195 m a.s.l., 25 February 2023; 5 ♀♀ and 1 ♂ from litter and soil under the hawthorn, 39°35′25″ N 39°01′00″ E, 1093 m a.s.l., 24 November 2023—the Karasu Valley, TÜRKİYE.

## Etymology

The species name “*incisus*” is derived from the Latin noun “*incisio*” which means cut and incision and refers to the presence of incision on lateral margins of hysterosomal shield in both sexes.

## Differential diagnosis

This new species resembles *Eustigmaeus anauniensis* (Canestrini, 1889), *E. hashmii* (Chaudhri, 1968), which was re-described by Stathakis *et al*. (2016) based on specimens from Greece, and *E. kentingensis* Tseng, 1982, by dorsum decorated with polygonal reticulate, dorsal setae thorny bush-like and setae *c*₂ similar in shape to other dorsal setae. However, it differs from them in that the hysterosomal shield is incised laterally vs hysterosomal shield is undivided in *E. anauniensis*, *E. hashmii* and *E. kentingensis*.

This new species is also similar to *Eustigmaeus aminiae* (Nazari and Khanjani, 2017) *n. comb.*, *E. bisetalis* (Doğan, 2004), *E. indiscretus* (Dönel and Doğan, 2011) *n. comb.*, by the hysterosomal shield incised laterally; however, it differs from them in the dorsal and ventral ornamentations. This new species can easily be distinguished from *E. bisetalis* by having
<figure><img src="image_8.png" /><figcaption>**Figure 6** *Eustigmaeus incisus* n. sp. (male) – A – Dorsal view, B – Ventral view. Scale bar: 100.</figcaption></figure>

homeomorphic dorsal body setae (heteromorphic in *E. bisetalis*). It differs from *E. indiscretus* in that the coxisternal shields are granulate (reticulate in *E. indiscretus*), and setae *c*₂ are similar in shape to dorsal body setae (not similar in shape to dorsal body setae in *E. indiscretus*). This new species can be distinguished from *E. aminiae* in that dorsal shields are prominently decorated with pits along the entire surface, palp tibial seta *l*′ is robust, paw-like (not seta-like), and coxisternal shields are covered with small grains (granulate) vs dorsal idiosoma has punctuations and its anterior and lateral margins bears pits, palp tibial seta *l*′ is seta-like, and coxisternal shields are marked with small points (punctuate) in *E. aminiae*. Additionally, a pair of rounded patterned areas in anterolateral parts of aggenital shield are present in the new species vs the patterned areas are absent in *E. aminiae*.
<figure><img src="image_9.png" /><figcaption>**Figure 7** *Eustigmaeus incisus* n. sp. (male) – A – Leg I, B – Leg II. Scale bar: 40.</figcaption></figure>

**Eustigmaeus paraincisus** n. sp.

Zoobank: 6939A1A2-2E77-4E9D-A367-0F5E2D5E6D01

(Figures 5B, D and 9–15)

**Diagnosis**

**Female** — Dorsal shields ornamented with pits, 9–33 vacuoles present on pits; dorsal setae homeomorphic, resembling a bush; humeral setae $c_2$ dissimilar to dorsal body setae; lateral incisions of hysterosomal shield present; a pair of almost rounded patterned areas located anterolaterally on aggenital shield.

**Description**

**Female** (n=6) (Figs 9–12)

Length of idiosoma 312 (312–335), width 234 (219–246).

**Dorsum** — (Fig. 9). Dorsal shields ornamented with pits, 9–33 vacuoles present on pits. Fossettes on dorsal shields as illustrated. Propodosomal shield with four pairs of setae: *vi*, *ve*, *sci* and *sce*, and a pair of ocelli, 10 (10–11) in diameter. Hysterosomal shield with incisions on its lateral margins, with six pairs of setae $c_1$, $d_1$, $d_2$, $e_1$, $e_2$ and $f_1$. Suranal shield with two pairs of setae: $h_1$ and $h_2$. Dorsal setae bushy. Setae $c_2$ longer and narrower than dorsal setae. Lengths and distances of dorsal setae as follows: *vi* 24 (21–22), *ve* 24 (22–24), *sci* 21 (16–21), *sce* 24 (21–23), $c_1$ 23 (20–22), $d_1$ 19 (18–21), $d_2$ 23 (19–22), $e_1$ 25 (24–25), $e_2$ 22 (19–23), $f_1$ 30 (29–32), $h_1$ 31 (28–30), $h_2$ 27 (24–26), *vi*-*vi* 29 (30–33), *ve*-*ve* 87 (80–84), *vi*-*ve* 39 (40–42), *sci*-*sci* 139 (130–135), *ve*-*sci* 33 (29–35), *sce*-*sce* 186 (173–182), *sci*-*sce* 39 (36–43), $c_1-c_1$ 76 (71–77), $d_2-d_2$ 197 (183–196), $c_1-d_1$ 60 (64–66), $c_1-d_2$ 66 (61–67), $d_1-d_1$ 76 (68–73), $d_2-d_1$ 67 (64–70), $e_2-e_2$ 157 (142–150), $e_1-e_1$ 106 (96–104), $e_2-e_1$ 50 (39–44),
<figure><img src="image_10.png" /><figcaption>**Figure 8** *Eustigmaeus incisus* n. sp. (male) - A - Leg III, B - Leg IV. Scale bar: 40.</figcaption></figure>

$f_1-f_1$ 60 (56–63), $e_1-f_1$ 39 (42–47), $e_2-f_1$ 88 (86–91), $h_1-h_1$ 41 (34–37), $h_2-h_2$ 62 (65–74), and $h_1-h_2$ 14 (16–18).

**Venter** — (Fig. 10). Coxisternal shields fused and with three pairs of intercoxal setae: 1a, 3a and 4a. Lengths and distances of these setae: 1a 13 (12–16), 3a 15 (13–14), 4a 12 (11–12), 1a–1a 27 (22–26), 3a–3a 36 (34–36), and 4a–4a 25 (22–26). Intercoxal setae smooth. Aggenital shield with three pairs of setae: *ag*₁, *ag*₂ and *ag*₃. Anogenital shields with three pairs of pseudanal setae: *ps*₁, *ps*₂ and *ps*₃. Lengths of these setae; *ag*₁ 8 (8–9), *ag*₂ 9 (8–10), *ag*₃ 12 (12–13), *ps*₁ 18 (16–19), *ps*₂ 17 (16–17) and *ps*₃ 15 (14–15).
<figure><img src="image_11.png" /><figcaption>**Figure 9** *Eustigmaeus paraincisus* n. sp. (female) – Dorsal view. Scale bar: 100.</figcaption></figure>

setae slightly barbed, ps₁ thicker than other pseudanal setae, and distinctly barbed. Patterns on humeral shields similar to those of dorsal shields. Patterns on coxisternal, aggenital, pseudanal and suranal shields as in Figure 10. Oval patterned area located between coxae III and IV and a pair of rounded patterned areas located anterolaterally on aggenital shield. Indentations and folds on anterior part of humeral shields present in a few paratypes, setae c₂ 34 (29–34), longer and thinner than dorsal body setae, pointed and barbed.

**Legs** — (Figs 11 and 12). Faintly punctuated. Setal formulae of legs I–IV: coxae 2–2–2–2, trochanters 1–1–2–1, femora 6–4–3–2, genua 3(+1κ)–3(+1κ)–1–1, tibiae 5(+1φ+1φρ)–5(+1φρ)–5(+1φρ)–5(+1φρ), tarsi 13(+1ω)–9(+1ω)–7(+1ω)–7.

Leg I (Fig. 11A). 172 (144–173). Leg setation: Co 2 (1b, 1c) and needle-like leg supracoxal setae (el), Tr 1 (v'), Femur 6 (d, l', l'', v', v'', bv''), Ge 3 (d, l', l'') + 1 κ, Ti 5 (d, l', l'', v', v'') + 1 solenidion φ + 1 solenidion φρ, Ta 13 (p'ξ, p''ξ, tc'ξ, tc''ξ, ft'ξ, ft''ξ, u', u'', a', a'', pl', pl'', vs) + 1 solenidion ω, 22 (21–23). Setae d on femur and tibia distinctly barbed and pointed; l', v', v'' on femur, l' on genu, l', v', v'' on tibia, pl', pl'', vs on tarsus pointed and smooth; setae tc'ξ, tc''ξ, ft'ξ, ft''ξ, p'ξ, p''ξ on tarsus smooth and weakly blunt-tipped; other setae on leg pointed and weakly barbed.

Leg II (Fig. 11B). 142 (129–147). Leg setation: Co 2 (2b, 2c), Tr 1 (v'), Fe 4 (d, l', l'', bv''), Ge 3 (d, l', l'') + 1 κ, Ti 5 (d, l', l'', v', v'') + 1 solenidion φρ, Ta 9 (p'ξ, tc'ξ, tc'', u', u'', a', a'', pl', vs) + 1 solenidion ω, 14 (13–14). Setae d on femur and tibia distinctly barbed and pointed; l' on femur, v'' on tibia pointed and smooth; setae tc'ξ, p'ξ on tarsus smooth and weakly blunt-tipped;
<figure><img src="image_12.png" /><figcaption>**Figure 10** *Eustigmaeus paraincisus* n. sp. (female) – Ventral view. Scale bar: 100.</figcaption></figure>

other setae on leg pointed and weakly barbed.

Leg III (Fig. 12A). 149 (134–145). Leg setation: Co 2 (3b, 3c), Tr 2 (*l', v*'), Fe 3 (*d, l', ev*'), Ge 1 (*d*), Ti 5 (*d, l', l'', v', v''*) + 1 solenidion φρ, Ta 7 (*tc', tc'', u', u'', a', a'', vs*) + 1 solenidion ω, 4 (3–4). Setae *d* on femur, genu and tibia distinctly barbed and pointed; vs, *tc', tc''* on tarsus smooth; other setae on leg pointed and weakly barbed.

Leg IV (Fig. 12B). 168 (147–170). Leg setation: Co 2 (4b, 4c), Tr 1 (*v*'), Fe 2 (*d, ev*'), Ge 1 (*d*), Ti 5 (*d, l', l'', v', v''*) + 1 solenidion φρ, Ta 7 (*tc', tc'', u', u'', a', a'', vs), lacking solenidion ω. Setae *d* on femur, genu and tibia, *l*' on tibia distinctly barbed and pointed; *l'', v', v''* on tibia, *tc', tc''* on tarsus smooth; other setae on leg pointed and weakly barbed.

**Gnathosoma** — (Fig. 5D). Subcapitulum 68 (66–69) long, faintly punctuated. Length of chelicerae 82 (81–83). Palp faintly punctuated, length of palp 87 (82–86). Lengths and distances between subcapitular setae: *m* 12 (11–12), *n* 9 (9–11), *m–m* 18 (18–19), *n–n* 22 (22–24), and *m–n* 11 (10–12). Number of setae on palpal segments: Tr 0, Fe 3 (*d, l', v''*), Ge 2 (*d, l''*), Ti 3 (*d, l', l''*) +1 claw, Ta 8 (fused eupathidia *ul'ξ*, *ul"ξ* and *sulξ*, eupathidion *acmξ*, *ba*, *bp*, *lp*, *va*) + 1 solenidion ω. All setae on palp femur barbed, other palp setae smooth. Palpal supracoxal setae (*ep*) needle-like. Palp tibial seta *l'* paw-like, not seta-like.

**Male** — (n=1) (Figs 13–15)

Idiosoma oval, much smaller than female. Length of idiosoma 268, width 174.

**Dorsum** — (Fig. 13A). Dorsal ornamentations as in female. Fossettes on dorsal shields as illustrated. Ocelli 8 in diameter. Hysterosomal shield completely divided into two shields;
<figure><img src="image_13.png" /><figcaption>**Figure 11** *Eustigmaeus paraincisus* n. sp. (female) – A – Leg I, B – Leg II. Scale bar: 40.</figcaption></figure>

anterior shield with three pairs of setae *c*₁, *d*₁, and *d*₂; posterior shield with three pairs of setae *e*₁, *e*₂, and *f*₁. Suranal shield located dorsally. Dorsal setae *vi*, *ve*, *sci* and *sce* weakly barbed, other setae smooth. Lengths and distances of dorsal setae as follows: *vi* 18, *ve* 14, *sci* 11, *sce* 14, *c*₁ 12, *d*₁ 9, *d*₂ 11, *e*₁ 8, *e*₂ 7, *f*₁ 32, *h*₁ 18, *h*₂ 23, *vi*-vi 27, *ve*-ve 62, *vi*-ve 33, *sci*-sci 102, *ve*-sci 24, *sce*-sce 133, *sci*-sce 30, *c*₁-*c*₁ 53, *c*₂-*c*₂ 154, *d*₂-*d*₂ 138, *c*₁-*d*₁ 50, *c*₁-*d*₂ 48, *d*₁-*d*₁ 48, *d*₂-*d*₁ 48, *e*₂-*e*₂ 99, *e*₁-*e*₁ 56, *e*₂-*e*₁ 26, *f*₁-*f*₁ 49, *e*₁-*f*₁ 20, *e*₂-*f*₁ 44, *h*₁-*h*₁ 28, *h*₂-*h*₂ 44, and *h*₁-*h*₂ 6.

**Venter** — (Fig. 13B). Coxisternal shields fused. Lengths and distances of intercoxal setae: *1a* 11, *3a* 12, *4a* 9, *1a*–*1a* 28, *3a*–*3a* 28, and *4a*–*4a* 19. Lengths of aggenital and pseudanal setae: *ag*₁ 9, *ag*₂ 12, *ps*₁ 2, *ps*₂ 4, and *ps*₃ 7. Pseudanal setae *ps*₁ and *ps*₂ spine-like. Aggenital and pseudanal setae smooth. Setae *c*₂ as in female, but not distinctly barbed, 15 long. Patterns on coxisternal, aggenital and humeral shields similar to those of dorsal shields.

**Legs** — (Figs 14 and 15). Length of legs, from leg I to leg IV respectively: 166, 131, 134, and 137. Counts of setae and solenidia on legs I–IV as in female except enlarged male
<figure><img src="image_14.png" /><figcaption>**Figure 12** *Eustigmaeus paraincisus* n. sp. (female) – A – Leg III, B – Leg IV. Scale bar: 40.</figcaption></figure>

solenidion (ω♂) on all tarsi present. All leg setae smooth.

*Gnathosoma*. As in female. Subcapitulum 57 long. Length of chelicerae 70, length of palp 77. Lengths and distances between subcapitular setae: *m* 12, *n* 6, *m–m* 16, *n–n* 17, and *m–n* 7.

**Type material**

Holotype: ♀ from oak litter, the Karasu Valley, TÜRKİYE, 39°12'14" N 38°34'29" E, 1065 m a.s.l., 28 October 2022, col. Salih Doğan. Paratypes: ♀ from oak litter, 39°12'12" N 38°34'25" E, 1085 m a.s.l., 27 November 2022; ♀ from soil, 39°12'35" N 38°34'31" E, 990 m a.s.l., 27 November 2022; ♀ from oak litter, 39°12'12" N 38°34'29" E, 1080 m a.s.l., 25 February 2023; ♀ from oak litter, 39°12'15" N 38°34'32" E, 1073 m a.s.l., 25 March 2023; 1♂ and 1♂ from moss, 39°13'33" N 38°34'12" E, 1180 m a.s.l., 26 October 2023—the Karasu Valley, TÜRKİYE.

**Differential diagnosis**

This new species is very similar to *Eustigmaeus incisus* n. sp. in having similar ornamentations on dorsal and ventral shields, lateral incisions of hysterosomal shield and dorsal body setae bush-like. However, it differs from *E. incisus* n. sp. in that setae *c*₂ are more slender, not
<figure><img src="image_15.png" /><figcaption>**Figure 13** *Eustigmaeus paraincisus* n. sp. (male) – A – Dorsal view, B – Ventral view. Scale bar: 100.</figcaption></figure>

similar in shape to other dorsal body setae vs c₂ are robust, similar in shape to others in *E. incisus* n. sp., dorsal body setae of female are relatively thinner and longer than those of *E. incisus* n. sp., and dorsal body setae of male are slenderer than those of *E. incisus* n. sp. For the differences between other closely related species, see the key to *Eustigmaeus* species with lateral incisions of hysterosomal shield and the “differential diagnosis” of *E. incisus* n. sp.

## Etymology

The species name "paraincisus" is a combination of the Greek prefix *para-* which means resembling, close to or similar to, and species name *incisus*, indicating that this new species resembles *Eustigmaeus incisus* n. sp.
<figure><img src="image_16.png" /><figcaption>Figure 14 *Eustigmaeus paraincisus* n. sp. (male) – A – Leg I, B – Leg II. Scale bar: 40.</figcaption></figure>

A key to *Eustigmaeus* species (female) with lateral incisions of hysterosomal shield

1. Dorsal body setae homeomorphic .................................................... 2
— Dorsal body setae heteromorphic .................................................. *E. bisetalis* (Doğan)

2. Coxisternal shields not reticulate .................................................... 3
— Coxisternal shields reticulate .................................................. *E. indiscretus* (Dönel and Doğan)
3. A pair of rounded patterned areas present anterolaterally on the aggenital shield, palp tibial seta l' paw-like (not seta-like) .................................................................................... 4
— Patterned areas absent anterolaterally on the aggenital shield, palp seta l' not modified ... E. aminiae (Nazari and Khanjani)

4. Setae $c_2$ robust, similar in shape to dorsal body setae.................................................. *E. incisus* n. sp.
— Setae $c_2$ more slender, dissimilar in shape to dorsal body setae.................. *E. paraincisus* n. sp.

Discussion

*Eustigmaeus sphagneticolus* Cooreman **n. stat.** originally described as a subspecies, *E. ottavii sphagneticola*, by Cooreman (1943) has been elevated to species level due to the following characteristics: the dorsal body setae are relatively shorter and the dorsal propodosomal setae do not extend to the base of the following setae, and two pairs of aggenital setae (*ag*₁ and

<figure><img src="image_17.png" /><figcaption>**Figure 15** *Eustigmaeus paraincisus* n. sp. (male) – A – Leg III, B – Leg IV. Scale bar: 40.</figcaption></figure>
$ag_2$) based on the description and illustrations by Cooreman (1943, pp. 4–6, and figs 6–7) vs dorsal body setae (especially *ve*) are longer and the dorsal propodosomal setae extend to the base of the following setae, and three pairs of aggenital setae in *E. ottavii* (Berlese) according to re-descriptions and characteristic features of the species by Summers and Price (1961), Wood (1973), Stathakis *et al.* (2016) and Khaustov (2019). Additionally, the humeral callosities were not mentioned in the original description and illustration, and it is clear that Cooreman’s populations do not belong to *ottavii*. Cooreman’s *sphagneticolus* specimens seem to be very close to the genus *Villersia* Oudemans by the presence of three pairs of setae on the propodosomal shield, and setae $d_2$ are located on hysterosomal shield as in *Villersia jamaliensis* (Khaustov, 2014). In the type specimens of *Paravillersia grata* Kuznetsov, synonymised with *Eustigmaues ottavii* by Khaustov (2019), setae *sce* are also located on separate shields or only on the thin striated incisions of the propodosomal shield (on propodosomal shield in other species of *Eustigmaeus*), and other characters of Cooreman’s *sphagneticolus* specimens are typical for *Eustigmaeus*, so it is more appropriate to keep Cooreman’s species in *Eustigmaeus* as originally given.

Khaustov (2019) suggested that *E. isfahaniensis* Khanjani, Najaf-Abadi and Khanjani, a species described from Iran, potentially could be a synonym of *E. ottavii* (Berlese). Later, Bagheri and Mohammad-Doustaresharaf (2024) considered *E. isfahaniensis* as a synonym of *E. ottavii*.

Another species from Iran, *Eustigmaeus nasrinae* Khanjani and Ueckermann is probably conspecific with *E. anauniensis* (Fan *et al.* 2016). On the other hand, the original description of *Eustigmaeus hooriaae* Khan, Afzal and Bashir from Pakistan is inadequate and the counts of its anogenital, palp and leg setae are inaccurate. It is therefore necessary to examine the type materials of the above-mentioned species.

*Raphignathus curtipilus* Berlese, 1888 was moved to the genus *Eustigmaeus* in Stigmaeidae by Doğan *et al.* (2024a).

*Liostigmaeus pulchellus* Thor, 1930, previously included in *Eustigmaeus*, is most likely belongs to the genus *Cheylostigmaeus* Willmann in Stigmaeidae (with recommendation of A.A. Khaustov, 2024). Therefore, it has now been moved to *Cheylostigmaeus*.

The hysterosomal shield is partially detached in *Eustigmaeus aminiae* (Nazari and Khanjani) **n. comb.**, and *E. indiscretus* (Dönel and Doğan) **n. comb.** (from Ledermuelleriopsis Willmann). We consider them intermediate species between the genera *Eustigmaeus* and *Ledermuelleriopsis*, and currently regard them as better placed in *Eustigmaeus*.

## Acknowledgements

The mite materials were collected from a project (121Z986) that was supported by the Scientific and Technological Research Council of Türkiye (TÜBİTAK). We appreciate the financial assistance of TÜBİTAK. We are grateful to TR Ministry of Agriculture and Forestry for granting the legal approvals (E-21264211-288.04-4255389 and E-50411936-903.07.02-4372543) for the field surveys. Dr A.A. Khaustov (Tyumen State University, Tyumen, Russia) was kind enough to call to the authors’ attention to the transfer of the species *Liostigmaeus pulchellus* Thor, 1930 to the genus *Cheylostigmaeus*. We are also grateful for the valuable contribution of the other reviewer to the manuscript.

## ORCID

Salih Doğan https://orcid.org/0000-0001-5030-0544

Sibel Doğan https://orcid.org/0000-0002-0644-0280
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