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# Ontogenetic instars of *Ciprusenia troglobionta* sp. nov. (Acari: Astigmata: Canestriniidae) from a cave of Zagros Mountains, Iran

Mojgan Sadat-Shojaei IDª, Ryszard Haitlinger IDᵇ, Saber Sadeghi IDᶜ, Mohammad Ali Akrami IDᵈ

## Original research

### ABSTRACT

A new species of canestriniid mite belonging to the genus *Ciprusenia* Haitlinger, 2022 is described based on the morphological characteristics of male, female, tritonymph, protonymph and larva life stages. It is the sixth species of this genus and was collected from *Blaps bushirensis* Kaszab, 1959 (Coleoptera: Tenebrionidae) on the surface soil of Sahlak Cave (Fars, Iran). The genus *Ciprusenia* is a new record for the Astigmata fauna of Iran and the host species is a new host record for the family Canestriniidae. In this study, descriptions are carried out in detailed line drawings and scanning electron microscopic images. The identification key to the world-known species of the genus is also provided.

**Keywords** subterranean; *Ciprusenia*; *Blaps*; troglobite; SEM; Astigmatic mite; cave-dwelling

**Zoobank** http://zoobank.org/FDCBFA34-07A8-4F25-AB8A-B3AA3AB37918

### Introduction

Astigmatic mites of the family Canestriniidae Berlese, 1884 include 370 species belonging to 101 genera. The species of this family are often reported as associated with 12 families of class Insecta including Blattidae, Carabidae, Cerambycidae, Chrysomelidae, Elateridae, Geotrupidae, Lucanidae, Passandridae, Passalidae, Scarabaeidae, Tenebrionidae and Zoopheridae as well as two families of class Diplopoda including Glomeridae and Odontopygidae. They may be phoretic or parasites (Beron 2021; Haitlinger 2022; Okabe et al. 2017; Trach and Khaustov 2011).

Genus *Ciprusenia* was proposed by Haitlinger in 2022 and he transferred five species of the genus *Percanestrinia* Berlese, 1911 to this new genus. In this study, we describe the sixth species of the genus and list all of them as follows: *Ciprusenia troglobionta* Sadat-Shojaei & Haitlinger sp. nov., *C. carabicola* (Berlese, 1882), *C. izabelae* (Haitlinger, 1992), *C. kamelskyi* (Samšiňák, 1970), *C. sellnicki* (Samšiňák, 1965), *C. viviannae* (Haitlinger, 1992) (Berlese 1911; Haitlinger 1992, 2022; Samšiňák 1965, 1970; Vitzthum 1924).

All of the mite life stages including male, female, tritonymph, protonymph and larva specimens were collected from the *Blaps bushirensis* Kaszab, 1959 (Fig. 1B). Only male and female forms were collected from the cave floor soil near the beetle host and not any stable population of the new species was extracted from the soil. Both males and females of the *Blaps bushirensis* were infected with this new mite species.

Received 23 February 2023
Accepted 13 November 2023
Published 11 January 2024

Corresponding authors
Saber Sadeghi ID:
ssadeghi@shirazu.ac.ir
Mohammad Ali Akrami ID:
akrami@shirazu.ac.ir

Academic editor
Akashi Hernandes, Fabio

https://doi.org/10.24349/owvp-jv03

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

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

Sadat-Shojaei M. et al.

Licensed under
Creative Commons CC-BY 4.0

<footnote>ªDepartment of Biology, Faculty of Sciences, Shiraz University, Shiraz, Iran.</footnote>

<footnote>ᵇInstitute of Environmental Biology, Department of Invertebrate Systematics and Ecology, Wrocław University of Environmental and Life Sciences, Poland.</footnote>

<footnote>ᶜDepartment of Plant Protection, School of Agriculture, Shiraz University, Shiraz, Iran.</footnote>
<figure><img src="image_7.png" /><figcaption>**Figure 1** Type locality of *Ciprusenia troglobionta* sp. nov. **A.** The entrance of Sahlak Cave. **B.** *Blaps bushirensis* Kaszab, 1959 as a host for a population of the new species in para-hypogean to hypogean transition zone in the Sahlak Cave.</figcaption></figure>

Usually, a relatively deep cave is divided into three main zones based on the amount of light penetration: the endogean zone or cave entrance area which receives the most amount of light, the para-hypogean zone or twilight area of the cave which receives less light than the entrance and the hypogean or dark area of the cave where no light penetrates (Lee *et al.* 2012).

The site of soil and host sampling was the transition zone of para-hypogean to hypogean of Sahlak Cave. The soil floor of this cave was mixed with bat guano in many parts. This cave is a natural karstic cave in the Zagros Mountains, with almost 330 meters in length which is located in Iran, Fars province, Darab county, Chah Kondar village, Sahlak canyon (Fig. 1A).

The Zagros Mountains form a vast orogenic range in Iran with approximately 2000 km long and 100 to 200 km wide in front of the Iranian and Turkish plateaus (Mouthereau *et al.* 2007; Ramsey *et al.* 2008). The biospeleology of many caves in the Zagros Mountains, especially the troglofauna of cave-dwelling Acari, has remained unknown.

Until now, *Coleopterophagus baali* Haitlinger, 1990 from *Protaetia speciosa* (Adams, 1817), *Photia polymorpha* Samšiňák, 1971 from *Carabus persianus* Roeschke, 1896 and an unidentified deutonymph of genus *Camirohylla* Haitlinger, 1991 from an unidentified species of *Dila* Fischer von Waldheim, 1844 have been recorded. Therefore, the Iranian fauna of the family Canestriniidae is weakly known (Ardeshir *et al.* 2016; Haitlinger 1990a; Samšiňák 1971). *Ciprusenia troglobionta* sp. nov. is the fourth record of the family and the first record of the genus for Astigmata mite fauna of Iran. The troglomorphic characteristics of this subterranean mite species as adaptations for living in a cave environment are discussed. The host species, *Blaps bushirensis*, is a new host record for the family Canestriniidae. The identification key of the world-known species of the genus *Ciprusenia* is provided. The adult male of *C. viviannae* (Haitlinger, 1992) is not listed in the male key due to lack of access. The holotype of this species is a female.

**Material and methods**

**Specimens and Sampling**

A sampling of the *Blaps bushirensis* and the soil of the cave floor near the beetles from the para-hypogean to the hypogean transition zone of Sahlak Cave was carried out on December 4, 2021 (Fig. 1A). Then in Entomology Research Laboratory (Biology Department), male and female mite specimens of the new species were extracted from the soil using Berlese-Tullgren funnels. Male, female, tritonymph, protonymph and larva specimens were collected from the
beetle host with the help of fine laboratory tools using a ZEISS Stemi Sv11 stereomicroscope (Fig. 1B). To remove surface contamination, all of the specimens were preserved in 70% ethanol.

## Observation and documentation

Since the color and general nature of the mite specimens often change after slide mounting, the male and female specimens were photographed in a cavity slide containing 70% ethanol using an Olympus CH2 Microscope equipped with a 7D Canon digital camera. To increase the quality of the output images, we used Zerene Stacker 2022 software (ver. 1.04, Zerene Systems, see https://zerenesystems.com/cms/home). Detailed character images of male and female specimens were also taken using a VEGA3 TESCAN scanning electron microscope. The male, female, tritonymph, protonymph, and larva specimens were cleared in Nesbitt's fluid and mounted into Hoyer's medium (Walter and Krantz 2009). Detailed characters of the mounted mite slides were photographed using a Nikon Eclipse 80i microscope equipped with a digital camera. Manual line drawings of the mounted mite slides were made using an Olympus BX53 phase contrast microscope equipped with a drawing tube. The initial line drawings were processed in Adobe Photoshop 2023 software (ver. 24.1.0, Adobe, see https://www.adobe.com/au/). All measurements are given in micrometers (µm) and taken using Fiji (ImageJ2) 2022 software (ver. 1.53t, ImageJ, see https://imagej.nih.gov/ij/index.html).

## Abbreviations

The terminology and abbreviations followed by related literature (Grandjean 1939; Griffiths *et al.* 1990; Haitlinger and Šundić 2016; Norton 1998; Trach and Khaustov 2011).

Additional abbreviations: IL, length of idiosoma; IW, width of idiosoma; GL, length of gnathosoma; CL, cheliceral length; MA, male adult; FA, female adult; TN, tritonymph; PN, protonymph; LA, larva; GO, Grandjean's organ.

Abbreviations of dorsal and ventral setae: *vi* and *ve*, internal and external vertical setae; *si* and *se*, internal and external scapular setae; *c*₁, *c*₂, *cp* and *c*₃, first (innermost), second, third and fourth setae in the first row on hysterosoma; *d*₁ and *d*₂, first (innermost) and second setae in the second row on hysterosoma; *e*₁ and *e*₂, first (innermost) and second setae in the third row on hysterosoma; *f*₂, second setae in the fourth row on hysterosoma; *h*₁, *h*₂ and *h*₃, first (innermost), second and third setae in the fifth row on hysterosoma; *scx*, supracoxal setae; *1a*, coxal-sternal setae associated with the bases of legs I; *3a*, coxal-sternal setae associated with the bases of legs III; *4a* and *4b*, first (innermost) and second coxal-sternal setae associated with the bases of legs IV; *g*, genital setae; *ad*₁, *ad*₂ and *ad*₃, first (posteriormost), second and third adanal setae; *ps*₁, *ps*₂ and *ps*₃, first (posteriormost), second and third pseudoadanal setae.

Abbreviations of depositories: ZM-CBSU, Zoological Museum, Collection of Biology Department, Shiraz University, Shiraz, Iran; JAZM, Jalal Afshar Zoological Museum, Department of Plant Protection, University of Tehran, Karaj, Iran; MNHWU, the Museum of Natural History, Wrocław University, Wrocław, Poland.

# Results

## Taxonomy

### Family Canestriniidae Berlese, 1884

### Genus Ciprusenia Haitlinger, 2022

*Ciprusenia* Haitlinger, 2022, p. 24. *Type species:* *Ciprusenia izabelae* (Haitlinger, 1992).
*Percanestrinia izabelae* - Haitlinger, 1992, p. 536.
<figure><img src="image_8.png" /><figcaption>**Figure 2** *Ciprusenia troglobionta* sp. nov. imaged in a cavity slide containing 70% ethanol. **A** – Adult male specimen. **B** – Adult female specimen.</figcaption></figure>

**Ciprusenia troglobionta Sadat-Shojaei & Haitlinger, sp. nov.**

Zoobank: 005F2E72-3105-47AE-B134-E2D84351437A

(Figures 1-21)

**Material examined**

**Holotype** — Male, Iran, Fars prov., Darab county, Chah Kondar vill., Sahlak canyon, Sahlak Cave; 28°32′51″N, 55°08′33″E; ex. *Blaps bushirensis*; 04 Dec 2021; leg. M. Sadat-Shojaei; microscopic slide, (ZM-CBSU Aca.00020).

**Paratypes** — Same collection data as holotype; 1 MA, 1 FA, microscopic slides, (MNHWU CTWP1, CTWP2); 1 MA, 1 FA, 1 TN, 1 LA, microscopic slides, (JAZM ARS-20230109-1b, ARS-20230109-1c, ARS-20230109-1d, ARS-20230109-1e); 2 MA, 2 FA, 3 TN, 3 PN, 3 LA, microscopic slides, (ZM-CBSU Aca.00021, Aca.00022, Aca.00029, Aca.00030, Aca.00036, Aca.00037, Aca.00038, Aca.00039, Aca.00040, Aca.00041, Aca.00042, Aca.00043, Aca.00044); 3 MA, 2 FA, mounted for SEM, (ZM-CBSU Aca.00023, Aca.00024, Aca.00025, Aca.00031, Aca.00032); 3 MA, 3 FA, preserved in ETOH. 70%, (ZM-CBSU Aca.00026, Aca.00027, Aca.00028, Aca.00033, Aca.00034, Aca.00035).

**Etymology**

'Troglobiont' refers to an obligatory cave-dwelling organism that is specialized for living in caves and exhibits troglomorphic characteristics.

**Diagnosis**

Male and female adults: white in color, transparent in some parts and without any pigmentation (Figs. 2A, B); propodosomal plate punctate; wavy sejugal furrow present, two diagonal wavy furrows on both sides of the hysterosoma; Grandjean's organs and setae *scx* well-developed; setae *ve* less than 20; setae *vi* and *si* spinulate; setae *se* and *cp* vascular-shaped; setae *vTr* I clearly pectinate.

Male: IL 250-341; setae *e*₂ flame-shaped, setae *nG* tricarinate.

Female: IL 358 – 396; tarsi IV elongated > 90; fine regular fibers layer beneath the genital valves layer.
<figure><img src="image_9.png" /><figcaption>**Figure 3** *Ciprusenia troglobionta* sp. nov., optical micrographs of all life stages. **A** – Adult male. **B** – Adult female. **C** – Tritonymph. **D** – Protonymph. **E** – Larva.</figcaption></figure>

**Description**

**Male** (holotype and 10 paratypes)

**Measurements** — (Paratypes in parentheses), IL 321 (250-341), IW 355 (303-362); GL 88 (78-91), CL 76 (72-82); leg I 191 (180-198), leg II 201 (178-205), leg III 222 (200-222), leg IV 241 (215-241) (Table 1).

<table><caption>**Table 1** Size changes of *Ciprusemia troglobionta* sp. nov. during ontogeny</caption><thead><tr><th>Character</th><th>LA</th><th>PN</th><th>TN</th><th>FA</th><th>MA</th></tr></thead><tbody><tr><td>IL</td><td>171-219</td><td>216-237</td><td>270-306</td><td>358-396</td><td>250-341</td></tr><tr><td>IW</td><td>133-193</td><td>179-189</td><td>216-273</td><td>313-375</td><td>303-362</td></tr><tr><td>GL</td><td>39-41</td><td>50-67</td><td>60-83</td><td>89-90</td><td>78-91</td></tr><tr><td>CL</td><td>40-43</td><td>51-70</td><td>61-77</td><td>76-87</td><td>72-82</td></tr><tr><td>Leg I</td><td>69-82</td><td>96-104</td><td>125-138</td><td>189-192</td><td>180-198</td></tr><tr><td>Leg II</td><td>63-79</td><td>96-104</td><td>130-145</td><td>180-206</td><td>178-205</td></tr><tr><td>Leg III</td><td>78-93</td><td>113-118</td><td>153-168</td><td>216-240</td><td>200-222</td></tr><tr><td>Leg IV</td><td>-</td><td>114-118</td><td>158-180</td><td>250-264</td><td>215-241</td></tr></tbody></table>

**Dorsum** — Idiosoma convex in middle part, maximum width between leg II and leg III, anterior margin finely wavy (Figs. 3A, 4A, 5A); idiosoma of smaller males with reticulate ornamentation, unlike larger ones and holotype (Figs. 4G, 13A); propodosomal plate punctate, with spinulate bulbiform setae *vi*, short smooth simple setae *ve* just behind the propodosomal
<figure><img src="image_10.png" /><figcaption>**Figure 4** *Ciprusenia troglobionta* sp. nov., SEM micrographs of adult male. **A** – Dorsal view. **B** – Ventral view. **C** – Genital and anal region. **D** – Wavy sejugal and two wavy diagonal furrows. **E** – Ventral view of gnathosoma. **F** – Barbed seta cG and smooth simple seta ve. **G** – Reticulate ornamentation of idiosoma. **H** – Grandjean's organ and vTr I seta. **I** – Tricarinate seta nG of leg III. **J** – Setae of propodosoma. **K** – Spinulate seta vi. **L** – Leaf-shaped bulbiform seta c₂.</figcaption></figure>
<figure><img src="image_11.png" /><figcaption>**Figure 5** *Ciprusenia troglobionta* sp. nov., adult male. A – Dorsal view. B – Seta *c*₁. C – Seta *ve*. D – Seta *d*₂. E – Seta *se*. F – Seta *scx*. G – Seta *si*. H – Seta *vi*.</figcaption></figure>
<figure><img src="image_12.png" /><figcaption>**Figure 6** *Ciprusenia troglobionta* sp. nov., adult male. **A** – Ventral view. **B** – Seta *c*₁. **C** – Seta *l*a. **D** – Seta *cp*.</figcaption></figure>

plate (Figs. 4J, K, 5A, C, H); scapular setae *si* and *se* present, smooth simple setae *scx* anterolateral near legI (Figs. 4J, 5F), setae *si* spinulate bulbiform (Fig. 5G), setae *se* long and vascular-shaped, with cup-like bothridium (Figs. 5E, 11D); mild wavy sejugal furrow present, two diagonal mild wavy furrows symmetrically begin between setae *c*₂ and *d*₂ and continue near seta *e*₁ (Figs. 4A, D, G, 5A); setae *c*₂ and *d*₂ leaf-shaped bulbiform (Figs. 4L, 5D, 11E); setae *c*₁, *d*₁ and *e*₁ smooth simple (Fig. 5B); hysterosoma contracted in opisthosomal border, opisthosoma with many fine wrinkled lines; setae *f*₂, *h*₁, *h*₂ and *h*₃ leaf-shaped bulbiform, all located on two lobes at end corners of body near the posterior margin of opisthosoma (Figs. 5A, 8A, 11L), *h*₁ > *h*₂ > *h*₃ > *f*₂; opisthothal gland, *gla*, conical and slightly folded around opening (Fig. 8B); length of dorsal setae: *vi* 57 (51-62), *ve* 12 (11-18), *scx* 49 (44-50), *si* 47 (41-51), *se* 362 (320-362), *c*₁ 32 (24-32), *c*₂ 75 (56-75), *d*₁ 31 (25-33), *d*₂ 89 (82-89), *e*₁ 35 (30-35), *f*₂ 80 (70-80), *h*₁ 427 (414-427), *h*₂ 320 (282-320), *h*₃ 128 (110-128).

**Venter** — Grandjean's organ well developed at the base of leg I close to gnathosoma, include three or four branches connected to an oval structure with a distinct opening and reticulated surface, longest branch just pectinate usually close to vTr I (Figs. 4E, H, 6A, 8C,
<figure><img src="image_13.png" /><figcaption>**Figure 7** *Ciprusenia troglobionta* sp. nov., genital and anal region of adult male.</figcaption></figure>

11C); coxal- sternal setae *1a, 3a, 4a, 4b* present, setae *1a* near base of leg I and smooth simple, long setae *3a* at base of leg III and leaf-shaped bulbiform, long seta *4a* at base of leg IV and leaf-shaped bulbiform, setae *4b* above genital region and leaf-shaped (Figs. 4B, C, 6A, C, 7); setae *c*₃ leaf-shaped bulbiform near to long vascular-shaped setae *cp* (Figs. , 4B, 6A, B, D, 11F); aedeagus observed beneath genital valves layer (Figs. 4C, 6A, 7, 11O); leaf-shaped genital setae *g*, close to genital papillae (Figs. 6A, 7, 11N, O); anal valve with fine striations, anal suckers well-developed, a layer with fine striations around lateral and posterior parts of anal suckers (Figs. 4C, 6A, 7, 11P); short setae *ps*₃ smooth simple, setae *ps*₂ and *ps*₃ leaf-shaped, *ps*₁ > *ps*₂ > *ps*₃; flame-shaped setae *e*₂ close to setae *ps*₁ all located on two lobes at end corners of body (Figs. 4C, 6A, 7, 11K, 11P); posterior margin of body delicately folded (Figs. 6A, 7, 11P); length of ventral setae: *1a* 47 (40-50), *3a* 129 (100-129), *4a* 95 (90-100), *4b* 41 (33-41), *c*₃ 72 (63-72), *cp* 275 (245-290), *g* 28 (24-29), *ps*₁ 40 (40-45), *ps*₂ 38 (38-41), *ps*₃ 19 (15-22), *e*₂ 15 (15-17).
<figure><img src="image_14.png" /><figcaption>**Figure 8** *Ciprusenia troglobionta* sp. nov., detailed characters of adult male. **A** – Structure of setae *h*₁, *h*₂, *h*₃ and *f*₂. **B** – Opisthonotal gland. **C** – Grandjean's organ. **D** – Chelicerae. **E** – Gnathosoma.</figcaption></figure>

**Gnathosoma** — Conical and mostly covered by idiosoma (in natural form and not in microscopic slide) (Figs. 2A, 4A, B, E); gnathosomal palps two segmented, palpal tarsi solenidion setae ω and pulpal tarsi setae pt present, palpal tibia setae *l* and *d* present, long supracoxal setae *elcp* and long subcapitular setae *m* observed dorsally and ventrally respectively (Figs. 4E, 8E, 11B); chelicerae chelate-dentate with 4-5 teeth, cheliceral setae *chb* filiform, *cha*
<figure><img src="image_15.png" /><figcaption>**Figure 9** *Ciprusenia troglobionta* sp. nov., adult male, dorsal view of legs. **A** – Leg I. **B** – Leg II. **C** – Leg III. **D** – Leg IV.</figcaption></figure>

spiniform, two conical spurs on each chelicera (Figs. 8D, 11A); length of gnathosomal setae: ω 6 (6-7), *pt* 9 (8-11), *l* 11 (7-11), *d* 20 (18-20), *elcp* 36 (30-36), *m* 34 (30-34), *cha* 4 (3-4), *chb* 10 (9-10).

**Legs** — Ambulacral disc, empodial claw and condylophore well-developed in all pretarsi;
<figure><img src="image_16.png" /><figcaption>**Figure 10** *Ciprusenia troglobionta* sp. nov., adult male, ventral view of trochanters and femora A – Leg I. B – Leg II. C – Leg III. D – Leg IV.</figcaption></figure>

last half tarsi I-III with fine transverse striations on dorsolatraly; tarsi IV with fine transverse striations on dorsolaterally in complete; setae vTr I clearly pectinate; setae cG I, cG II and vTr II barbed; setae nG tricarinate; postaxial femur I partly punctate dorsally; trochanter IV partly with fine striations dorsally in junction of femur; (Figs. 4F, I, J, 9A-D, 10A-D, 11G, H, I, M); chaetotaxy of legs (I-1V): trochanters 1-1-1-1, femora 1-1-0-0, genua 2(1σ)-2(1σ)-1-0, tibiae (1φ)-(1φ)-(1φ)(1φ), tarsi 7(3ω,1ε)-7(1ω)-4-5, (Table 2); measurements of leg segments and setae (length of tarsi excluding pretarsi): **leg I** 191 (180-198), Tr I 36 (30-36), vTr 47 (46-48), Fe I 41 (37-41), vF 57 (53-62), Ge I 57 (44-57), cG 51 (45-52), mG 39 (30-41), σ 39 (37-42), Ti I 30 (29-32), φ 102 (97-109), Ta I 58 (52-64), ω₁ 28 (21-28), ω₂ 7 (7-9), ε 6 (6), wa 23 (22-24), la 16 (16-20), d 41 (40-43), ω₃ 30 (26-30), e 21 (20-22), f 19 (15-19), p 10 (10), q 10 (10); **leg II** 201 (178-205), Tr II 31 (30-37), vTr 41 (38-41), Fe II 45 (38-45), vF 88 (80-90), Ge II 52 (50-56), cG 39 (39-49), mG 38 (30-44), σ 13 (13-17), Ti II 31 (30-32), φ 72 (67-73), Ta II 56 (54-57), ω 37 (32-41), wa 27 (26-27), la 18 (14-18), d 65 (62-65), e 17 (15-17), f 23 (20-23), p 7 (6-7), q 8 (7-8); **leg III** 222 (200-222), Tr III 54 (47-55), vTr 35 (30-38), Fe III 31 (29-31), Ge
<figure><img src="image_17.png" /></figure>

**Figure 11** *Ciprusenia troglobionta* sp. nov., optical micrographs of detailed characters. **A** – Chelicerae of adult male. **B** – Gnathosoma of adult male. **C** – Grandjean's organ and seta *vTr* I of adult male. **D** – Seta *cG* and set *se* of adult male. **E** – Setae *mG* and *c*₂ of adult male. **F** – Setae *cp* and *c*₃ of adult male. **G** – Tarsus I of adult male, dorsal view. **H** – Tarsus I of adult male, ventral view. **I** – Ventral bases of trochanter II and femura II of adult male. **J** – Tricarinate seta *nG* of leg III of adult male. **K** – Setae *ps*₁ and *e*₂ of adult male. **L** – Setae *h*₁, *h*₂, *h*₃ and *f*₂ of adult male. **M** – Ventral base of trochanter IV of adult male. **N** – Genital region of adult male. **O** – Aedeagus and genital papillae of adult male. **P** – Anal region and suckers of adult male. **Q** – Anal region of adult female. **R** – fine regular fibers layer beneath the genital valves layer of adult female. **S** – Bursa copulatrix details of adult female. **T** – Claparède organ of larva.

III 54 (50-55), *nG* 14 (12-16), Ti III 33 (31-34), φ 59 (50-59), Ta III 53 (51-58), *w* 22 (22-25), *d* 89 (80-90), *f* 21 (19-21), *p* 6 (6); **leg IV** 241 (215-241), Tr IV 55 (47-55), *vTr* 37 (31-43), Fe IV 45 (40-45), Ge IV 65 (56-65), Ti IV 42 (38-42), φ 69 (60-69), Ta IV 58 (50-58), *r* 11 (9-11), *w* 21 (17-22), *d* 89 (84-90), *f* 21 (17-21), *s* 4 (3-4).

**Female** (9 paratypes)
<figure><img src="image_18.png" /><figcaption>**Figure 12** *Ciprusenia troglobionta* sp. nov., SEM micrographs of adult female. **A** – Dorsal view. **B** – Ventral view. **C** – Genital and anal region. **D** – Reticulate ornamentation of idiosoma. **E** – Setae of propodosoma. **F** – Spinulate seta *vi*.</figcaption></figure>

**Measurements** — IL 358-396, IW 313-375, GL 89-90, CL 76-87; leg I 189-192, leg II 180-206, leg III 216-240, leg IV 250-264 (Table 1).

**Dorsum** — Idiosoma moderately convex in middle part, maximum width between leg II and leg III, anterior margin finely wavy (Figs. 3B, 12A, 14A); idiosoma of smaller females with reticulate ornamentation, unlike larger ones (Figs. 12D, 13B); propodosomal plate punctate, with spinulate bulbiform setae *vi*, short smooth simple setae *ve* just behind the propodosomal plate (Figs. 12A, E, F, 14A, C, H,); scapular setae *si* and *se* present, smooth simple setae *scx* anterolateral near leg I, setae *si* spinulate bulbiform, setae *se* long and vascular-shaped, with cup-like bothridium (Figs. 12A, E, 14A, E, F, G); mild wavy sejugal furrow present, two diagonal mild wavy furrows symmetrically begin between setae *c*₂ and *d*₂ and continue near

<table><caption>**Table 2** Legs chaetotaxy of *Ciprusemia troglobionta* sp. nov. during ontogeny</caption><thead><tr><th>Life stage</th><th>Trochanters</th><th>Femora</th><th>Genua</th><th>Tibiae</th><th>Tarsi</th></tr></thead><tbody><tr><td>LA</td><td>0-0-0</td><td>1-1-0</td><td>2(1σ)-2(1σ)-1</td><td>(1φ)-(1φ)-(1φ)</td><td>7(3ω,1ε)-4(1ω)-3</td></tr><tr><td>PN</td><td>0-0-0-0</td><td>1-1-0-0</td><td>2(1σ)-2(1σ)-1-0</td><td>(1φ)-(1φ)-(1φ)-0</td><td>6(3ω,1ε)-5(1ω)-3-3</td></tr><tr><td>TN</td><td>1-1-1-0</td><td>1-1-0-0</td><td>2(1σ)-2(1σ)-1-0</td><td>(1φ)-(1φ)-(1φ)-(1φ)</td><td>7(3ω,1ε)-5(1ω)-3-5</td></tr><tr><td>FA</td><td>1-1-1-1</td><td>1-1-0-0</td><td>2(1σ)-2(1σ)-1-0</td><td>(1φ)-(1φ)-(1φ)-(1φ)</td><td>7(3ω,1ε)-7(1ω)-4-5</td></tr><tr><td>MA</td><td>1-1-1-1</td><td>1-1-0-0</td><td>2(1σ)-2(1σ)-1-0</td><td>(1φ)-(1φ)-(1φ)-(1φ)</td><td>7(3ω,1ε)-7(1ω)-4-5</td></tr></tbody></table>
<figure><img src="image_19.png" /><figcaption>**Figure 13** *Ciprusenia troglobionta* sp. nov., reticulate ornamentation of idiosoma of smaller specimens. **A** – Small adult male. **B** – Small adult female.</figcaption></figure>

setae *e*₁ (Figs. 12A, D, 14A); setae *c*₂ and *d*₂ leaf-shaped bulbiform (Figs. 14A, D); setae *c*₁, *d*₁ and *e*₁ smooth simple (Figs. 12A, D, 14A, B); terminal margin of opisthosoma with some vertical folding; opisthothal gland, *gla*, oval form (Fig. 14A); length of dorsal setae: *vi* 50-59, *ve* 10-14, *scx* 42-44, *si* 38-45, *se* 332-337, *c*₁ 25-33, *c*₂ 50-61, *d*₁ 24-35, *d*₂ 60-67, *e*₁ 14-22.

**Venter** — Grandjean’s organ well-developed at the base of leg I close to gnathosoma; Grandjean’s organ similar to male (Fig. 15A); coxal- sternal setae *1a, 3a, 4a, 4b* present, setae *1a* near base of leg I and smooth simple, long setae *3a* at base of leg III and leaf-shaped bulbiform, long setae *4a* at base of leg IV and leaf-shaped bulbiform, smooth simple setae *4b* next to genital region (Figs. 12B, C, 15A, C, 16A); setae *c*₃ leaf-shaped bulbiform near to long vascular-shaped setae *cp* (Figs. 15A, B, D); fine regular fibers layer beneath the genital valves layer, leaf-shaped genital setae *g*, close to genital papillae (Figs. 11R, 12C, 15A, 16A); four pairs of smooth simple setae around anal region, shortest one setae *ad*₁, others setae *ad*₂, setae *ps*₂ and setae *ps*₃; long smooth simple setae *ps*₁ little farther toward corner of body, *ps*₁ > *ps*₃ > *ps*₂ (Figs. 11Q, 15A); setae *e*₂ and *f*₂ smooth simple; two pairs long thick setae *h*₁ and *h*₂, leaf-shaped bulbiform setae *h*₃, setae *e*₂ and setae *f*₂ all located on two lobes at end corners of body (Figs. 15A, 16A); bursa copulatrix below anal region and between two lobes, copulatory opening ring-shaped, spermathecal duct thick and twisted, two U-shaped sclerites of oviduct end to spermatheca (Figs. 11S, 16A, B); terminal margin of body wrinkled (Figs. 15A, 16A). Length of ventral setae: *1a* 41-46, *3a* 85-95, *4a* 110-133, *4b* 28-37, *c*₃ 64-70, *cp* 248-270, *g* 22-25, *ps*₁ 46-53, *ps*₂ 20-24, *ps*₃ 22-28, *e*₂ 25-30, *f*₂ 22-31, *h*₁ 374-392, *h*₂ 321-335, *h*₃ 111-128.

**Gnathosoma** — General structure and setae similar to male (Figs. 15E, F); length of gnathosomal setae: *ω* 7, *pt* 12-14, *l* 9-10, *d* 19-23, *elcp* 32-39, *m* 29-34, *cha* 3-4, *chb* 7-8.

**Legs** — Ambulacral disc, empodial claw and condylophore well-developed in all pretarsi; setae *vTr* I clearly pectinate; setae *cG* I, *cG* II, *vTr* II barbed; setae *nG* weakly barbed; transverse lines on all leg solenidiae setae exclude *φ* I and *φ* II; broom-shaped structure includes a cover-like part and a delicate multi-branched appendage below it at each ventral bases of trochanters I, II; one conical structure at each ventral bases of femur I, II and IV, two conical structures at ventral base of femur III; tarsi IV elongated > 90 (Figs. 17A-D, 18A-D); chaetotaxy of legs (I-IV): trochanters 1-1-1-1, femora 1-1-0-0, genua 2(1σ)-2(1σ)-1-0, tibiae (1φ)-(1φ)-(1φ)-(1φ),
<figure><img src="image_20.png" /><figcaption>**Figure 14** *Ciprusenia troglobionta* sp. nov., adult female. **A** – Dorsal view. **B** – Seta *c*₁. **C** – Seta *ve*. **D** – Seta *d*₂. **E** – Seta *se*. **F** – Seta *scx*. **G** – Seta *si*. **H** – Seta *vi*.</figcaption></figure>

tarsi 7(3ω,1ε)-7(1ω)-4-5, (Table 2); measurements of leg segments and setae (length of tarsi excluding pretarsi): **leg I** 189-192, Tr I 26-32, *vTr* 45-51 Fe I 37-44, *vF* 39-45, Ge I 44-55, *cG* 35-47, *mG* 32-40, σ 38-43, Ti I 27-30, φ 106-110, Ta I 57-62, ω₁ 28-33, ω₂ 6-8, ε 5, *wa* 26-28, *la* 14-17, *d* 37-41, ω₃ 29-30, *e* 19-22, *f* 19-22, *p* 9-10, *q* 9-10; **leg II** 180-206, Tr II 28-35, *vTr* 37-44, Fe II 36-41, *vF* 64-83, Ge II 49-54, *cG* 30-33, *mG* 30-36, σ 17-18, Ti II 25-29, φ 68-74,
<figure><img src="image_21.png" /><figcaption>**Figure 15** *Ciprusenia troglobionta* sp. nov., adult female. **A** – Ventral view. **B** – Seta *c*₃. **C** – Seta *la*. **D** – Seta *cp*. **E** – Chelicerae. **F** – Gnathosoma.</figcaption></figure>
<figure><img src="image_22.png" /><figcaption>**Figure 16** *Ciprusenia troglobionta* sp. nov., adult female. **A** – Genital and anal region. **B** – Details of bursa copulatrix.</figcaption></figure>

Ta II 54-65, ω 36-42, wa 25-30, la 14-19, d 51-54, e 20-23, f 20-23, p 7, q 7; **leg III** 216-240, Tr III 37-42, vTr 39-45, Fe III 28-32, Ge III 49-58, nG 42-50, Ti III 38-42, φ 53-59, Ta III 72-74, w 23-25, d 90-99, f 23-26, p 7; **leg IV** 250-264, Tr IV 35-18, vTr 40-50, Fe IV 28-32, Ge IV 60-68, Ti IV 35-40, φ 33-34, Ta IV 92-95, r 11-13, w 23-27, d 102-104, f 20-24, p 7.

**Tritonymph** (4 paratypes)

**Measurements** — IL 270-306, IW 216-273, GL 60-83, CL 61-77; leg I 125-138, leg II
<figure><img src="image_23.png" /><figcaption>**Figure 17** *Ciprusenia troglobionta* sp. nov., adult female, dorsal view of legs. **A** – Leg I. **B** – Leg II. **C** – Leg III. **D** – Leg IV.</figcaption></figure>

130-145, leg III 153-168, leg IV 158-180 (Table 1).

**Dorsum** — Idiosoma oval form, maximum width in middle part between leg II and leg III; propodosomal plate not punctate; anterior margin smooth, lateral and terminal margins wrinkled; anterior propodosoma with wrinkled lines, hysterosoma and posterior propodosoma
<figure><img src="image_24.png" /><figcaption>**Figure 18** *Ciprusenia troglobionta* sp. nov., adult female, ventral view of trochanters and femora A – Leg I. B – Leg II. C – Leg III. D – Leg IV.</figcaption></figure>

with reticulate ornamentation; setae *vi* weakly spinulate bulbiform; short setae *ve* smooth simple; smooth simple setae *scx* anterolateral near leg I; setae *si* weakly spinulate bulbiform; long setae *se* vascular-shaped; mild wavy sejugal furrow and two diagonal mild wavy furrows well-developed; setae *c*₁ and *d*₂ leaf-shaped without transverse veins; all setae *c*₁, *d*₁, *e*₁ and *e*₂ smooth simple; opisthothal gland, *g/a*, unclear (Figs. 3C, 19A); length of dorsal setae: *vi* 41-42, *ve* 7-8, *scx* 33-36, *si* 34-37, *se* 271-285, *c*₁ 26-33, *c*₂ 40-46, *d*₁ 25-31, *d*₂ 46-47, *e*₁ 21-26, *e*₂ 19-26.

**Venter** — Grandjean's organ well-developed at the base of leg I close to gnathosoma; filiform coxal- sternal setae *1a, 3a, 4a, 4b* present, setae *1a* near base of leg I, long setae *3a* at base of leg III, long setae *4a* at base of leg IV, setae *4b* above genital setae *g*; smooth simple setae *c*₃ near to long vascular-shaped setae *cp*; filiform genital setae *g*, close to genital papillae; four pairs of filiform setae around anal region, longest one setae *h*₃, others setae *ps*₁, *ps*₂ and *ps*₃, setae *ps*₁ nearest and setae *ps*₃ furthest to anal region; long vascular-shaped setae *h*₁, *h*₂ and smooth simple setae *f*₂ all located on two lobes at end corners of body (Fig. 19B); length of ventral setae: *1a* 29-34, *3a* 60-72, *4a* 65-85, *4b* 25-28, *c*₃ 36-41, *cp* 203-225, *g* 14-18, *ps*₁ 18-24, *ps*₂ 22-28, *ps*₃ 25-30, *f*₂ 20-22, *h*₁ 320-338, *h*₂ 200-250, *h*₃ 104-113.

**Gnathosoma** — General structure and setae similar to adult form, setae *cha* not observed (Figs. 19G, H); length of gnathosomal setae: ω 5-7, *pt* 9-10, *l* 8-10, *d* 15-19, *elcp* 23-28, *m* 25-30, *chb* 6-7.
<figure><img src="image_25.png" /><figcaption>**Figure 19** *Ciprusenia troglobionta* sp. nov., tritonymph. **A** – Dorsal view. **B** – Ventral view. **C** – Dorsal view of leg I. **D** – Dorsal view of leg II. **E** – Dorsal view of leg III. **F** – Dorsal view of leg IV. **G** – Gnathosoma. **H** – Chelicerae.</figcaption></figure>
<figure><img src="image_26.png" /><figcaption>**Figure 20** *Ciprusenia troglobionta* sp. nov., protonymph. **A** – Dorsal view. **B** – Ventral view. **C** – Dorsal view of leg I. **D** – Dorsal view of leg II. **E** – Dorsal view of leg III. **F** – Dorsal view of leg IV. **G** – Chelicerae. **H** – Gnathosoma.</figcaption></figure>
<figure><img src="image_27.png" /><figcaption>**Figure 21** *Ciprusenia troglobionta* sp. nov., larva. **A** – Dorsal view. **B** – Ventral view. **C** – Dorsal view of leg I. **D** – Dorsal view of leg II. **E** – Dorsal view of leg III. **F** – Chelicerae. **G** – Gnathosoma.</figcaption></figure>
**Legs** — Ambulacral disc, empodial claw and condylophore developed in all pretarsi; setae vTr pectinate; setae cG I, cG II, vTr II barbed; setae nG weakly barbed (Figs. 19B-F); chaetotaxy of legs (I-IV): trochanters 1-1-1-0, femora 1-1-0-0, genua 2(1σ)-2(1σ)-1-0, tibiae (1φ)-(1φ)-(1φ)-(1φ), tarsi 7(3ω,1ε)-5(1ω)-3-5, (Table 2); measurements of leg segments and setae (length of tarsi excluding pretarsi): **leg I** 125-138, Tr I 20-25, vTr 36-40, Fe I 31-34, vF 34-38, Ge I 30-37, cG 27-34, mG 22-27, σ 31-35, Ti I 21-25, φ 72-90, Ta I 41-45, ω₁ 16-27, ω₂ 5-8, ε 5, wa 14-20, la 9-16, d 37-40, ω₃ 22-30, e 12-17, f 9-17, p 9, q 9; **leg II** 130-145, Tr II 21-28, vTr 23-30, Fe II 29-33, vF 47-58, Ge II 30-35, cG 23-28, mG 17-22, σ 12-15, Ti II 18-22, φ 47-60, Ta II 42-44, ω 28-34, wa 20-25, la 11-13, d 41-51, e 17-20, f 16-18; **leg III** 153-168, Tr III 31-35, vTr 32-34, Fe III 23-24, Ge III 34-40, nG 33-36, Ti III 23-27, φ 34-45, Ta III 45-58, w 19-24, d 63-74, f 16-18; **leg IV** 158-180, Tr IV 23-32, Fe IV 24-26, Ge IV 35-37, Ti IV 26-30, φ 14-22, Ta IV 59-70, r 7-9, w 14-22, d 54-84, f 17-21, p 5.

**Protonymph** (3 paratypes)

**Measurements** — IL 216-237, IW 179-189, GL 50-67, CL 51-70; leg I 96-104, leg II 96-104, leg III 113-118, leg IV 114-118 (Table 1).

**Dorsum** — Idiosoma oval form, maximum width in middle part between leg II and leg III; propodosomal plate not punctate; anterior margin smooth, lateral and terminal margins wrinkled; anterior propodosoma with wrinkled lines, hysterosoma and posterior propodosoma with reticulate ornamentation; spinulation setae *vi* and *si* not yet developed; short setae *ve* smooth simple; smooth simple setae *scx* anterolateral near leg I; long setae *se* weakly vascular-shaped; mild wavy sejugal furrow and two diagonal mild wavy furrows well-developed; all setae *c*₁, *c*₂, *d*₁, *d*₃, *e*₁ and *e*₂ smooth simple; opisthothal gland, *gla*, unclear (Figs. 3D, 20A); length of dorsal setae: length of dorsal setae: *vi* 26-30, *ve* 6-7, *scx* 24-28, *si* 22-26, *se* 192-220, *c*₁ 20-23, *c*₂ 26-28, *d*₁ 19-22, *d*₂ 28-32, *e*₁ 17-20, *e*₂ 16-19.

**Venter** — Wrinkled lines on ventral surface of idiosoma; Grandjean's organ developed; filiform coxal-sternal setae *1a, 3a, 4a, 4b* present; filiform setae *c*₃ near to semi-long weakly vascular-shaped setae *cp*; filiform genital setae *g*, close to one pair of genital papillae; four pairs of filiform setae around anal region, longest one setae *h*₃, others setae *ps*₁, *ps*₂ and *ps*₃, setae *ps*₁ nearest and setae *ps*₃ furthest to anal region; long weakly vascular-shaped setae *h*₁, *h*₂ and filiform setae *f*₂ all located on two small lobes at end corners of body (Fig. 20B); length of ventral setae: *1a* 22-24, *3a* 39-40, *4a* 55-56, *4b* 13-14, *c*₃ 26-29, *cp* 143-180, *g* 8-9, *ps*₁ 14-15, *ps*₂ 15-17, *ps*₃ 18-19, *f*₂ 14-18, *h*₁ 230-243, *h*₂ 125-135, *h*₃ 49-50.

**Gnathosoma** — General structure and setae similar to adult form, setae *cha* and *chb* not observed (Figs. 20G, H); length of gnathosomal setae: ω 4-6, *pt* 6-9, *l* 5-7, *d* 11-13, *elcp* 19-21, *m* 15-19.

**Legs** — Ambulacral disc, empodial claw and condylophore developed in all pretarsi; setae cG I and cG II barbed; setae nG weakly barbed (Figs. 20B-F); chaetotaxy of legs (I-IV): trochanters 0-0-0-0, femora 1-1-0-0, genua 2(1σ)-2(1σ)-1-0, tibiae (1φ)-(1φ)-(1φ)-0, tarsi 6(3ω,1ε)-5(1ω)-3-3, (Table 2); measurements of leg segments and setae (length of tarsi excluding pretarsi): **leg I** 96-104, Tr I 17-20, Fe I 24-27, vF 28-34, Ge I 24-26, cG 23-24, mG 17-19, σ 24-26, Ti I 17-20, φ 53-65, Ta I 30-32, ω₁ 15-20, ω₂ 5-6, ε 5, wa 12-14, la 9-10, d 37-40, ω₃ 27-28, e 13-15, f 14-15, p 6-8; **leg II** 96-104, Tr II 14-18, Fe II 23-25, vF 33-34, Ge II 24-27, cG 16-17, mG 13-14, σ 10-13, Ti II 16-17, φ 39-42, Ta II 28-32, ω 17-23, wa 12-16, la 9-10, d 34-40, e 17-19, f 13-16; **leg III** 113-118, Tr III 25-28, Fe III 17-20, Ge III 26-27, nG 27-28. Ti III 20-22, φ 26-30, Ta III 34-38, w 11-15, d 42-50, f 12-14; **leg IV** 114-118, Tr IV 20-23, Fe IV 17-18, Ge IV 23-27, Ti IV 18-22, Ta IV 41-43, r 4-5, w 11-12, d 40-55.

**Larva** (4 paratypes)

**Measurements** — IL 171-219, IW 133-193, GL 39-41, CL 40-43; leg I 69-82, leg II 63-79, leg III 78-93 (Table 1).

**Dorsum** — Idiosoma oval form, maximum width in middle part between leg II and leg III; propodosomal plate not punctate; anterior propodosoma with some wrinkled lines, hysterosoma and posterior propodosoma with uncomplete reticulate ornamentation; smooth simple setae *vi* present; setae *ve* absent; filiform setae *scx* anterolateral near leg I; long smooth setae *se* close
to filiform setae *si*; mild wavy sejugal furrow weakly developed; two diagonal furrows not yet developed; all setae *c*₁, *c*₂, *d*₁, *d*₂, *e*₁ and *e*₂ filiform; opisthonothal gland, *gla*, unclear (Figs. 3E, 21A); *vi* 25-28, *scx* 15-23, *si* 12-13, *se* 129-140, *c*₁ 10-12, *c*₂ 10-14, *d*₁ 9-13, *d*₂ 13-14, *e*₁ 9-11, *e*₂ 9-10.

**Venter** — Grandjean’s organ developed; filiform coaxal-sternal setae *1a, 3a* present, setae *4a, 4b* absent; Claparede organ well-developed; filiform setae *c*₃ near to semi-long smooth setae *cp*; filiform setae *h*₂ below anal region; long weakly vascular-shaped setae *h*₁ located on very small two lobes at end corners of body; setae *h*₃ absent; genital and pseudoanal setae not yet developed (Fig. 21B); length of ventral setae: *1a* 14-17, *3a* 24-36, *c*₃ 18-21, *cp* 90-110, *h*₁ 160-180, *h*₂ 54-65, Claparede organ 10-12.

**Gnathosoma** — General structure and setae similar to adult form, setae *cha* and *chb* not observed (Figs. 21F, G); length of gnathosomal setae: ω 4-5, *pt* 6-7, *l* 4-5, *d* 7-10, *elcp* 14-18, *m* 13-16.

**Legs** — Ambulacral disc, empodial claw and condylophore developed in all pretarsi; no setae yet barbed (Figs. 21B-E); chaetotaxy of legs (I-IV): chaetotaxy of legs (I-III): trochanters 0-0-0, femora 1-1-0, genua 2(1σ)-2(1σ)-1, tibiae (1φ)-(1φ)-(1φ), tarsi 7(3ω,1ε)-4(1ω)-3, (Table 2); measurements of leg segments and setae (length of tarsi excluding pretarsi): **leg I** 69-82, Tr I 13-16, Fe I 12-19, vF 20-21, Ge I 17-20, cG 10-18, mG 11-16, σ 11-13, Ti I 13-16, φ 42-47, Ta I 22-28, ω₁ 10-17, ω₂ 3-5, ε 2, wa 9-12, *la* 4-6, *d* 19-23, *e* 11-12, *f* 8-11, *p* 9-10, *q* 9-10; **leg II** 63-79, Tr II 13-15, Fe II 17-19, vF 26-33, Ge II 17-21, cG 10-15, mG 11-13, σ 4-5, Ti II 11-13, φ 35-36, Ta II 23-25, ω 11-17, wa 8-11, *d* 29-33, *e* 7-10; **leg III** 78-93, Tr III 16-20, Fe III 14-16, Ge III 19-22, *nG* 17-20, Ti III 14-16, φ 22-28, Ta III 32-33, *w* 8-11, *d* 25-40, *f* 12-13;

**Remarks**

The distinguishable characters of *Ciprusenia troglobionta* sp. nov. are mentioned in the diagnosis section. Additional comparisons are as follows: *C. troglobionta* sp. nov. differs from *C. kamelskyi* (Samšiňák, 1970), males, in shorter IL (250-341 vs. 380), *vi* (51-62 vs. 75), *ve* (11-18 vs. 85), *c*₂ (56-75 vs. 85), *e*₂ (15-17 vs. 45), *ps*₂ (38-41 vs. 60), longer *scx* (44-50 vs. 25), *c*₁ (24-32 vs. 10), *d*₁ (25-33 vs. 10), *e*₁ (30-35 vs. 17), *f*₂ (70-80 vs. 17), *ps*₁ (40-45 vs. 35), *h*₃ (110-128 vs. 90), females, shorter *vi* (50-59 vs. 80), *ve* (10-14 vs. 90), *c*₂ (50-61 vs. 85), *d*₂ (60-67 vs. 85), *e*₂ (25-30 vs. 50); *C. troglobionta* sp. nov. differs from *C. sellnicki* (Samšiňák, 1965), males, in shorter IL (250-341 vs. 570), IW (303-362 vs. 380), *scx* (44-50 vs. 60), longer *si* (41-51 vs. 20), *se* (320-362 vs. 300), *c*₁ (24-32 vs. 20), *c*₂ (56-75 vs. 20), *d*₁ (25-33 vs 20), *d*₂ (82-89 vs. 20), *f*₂ (70-80 vs. 20), *h*₁ (414-427 vs. 400), *h*₂ (282-320 vs. 250), *h*₃ (110-128 vs. 100), females, shorter IL (358-396 vs. 880), IW (313-375 vs. 600), *vi* (50-59 vs. 80), *ve* (10-14 vs. 30), *scx* (42-44 vs. 90), *c*₁ (25-33 vs. 45), *d*₁ (24-35 vs. 45), *e*₁ (14-22 vs. 45), *h*₁ (374-392 vs. 550), *h*₂ (321-335 vs. 350), *h*₃ (111-128 vs. 190), *f*₂ (22-31 vs. 45), longer *c*₂ (50-61 vs. 45), *d*₂ (60-67 vs. 45); *C. troglobionta* sp. nov. differs from *C. izabelae* (Haitlinger, 1992), males, in shorter IL (250-341 vs. 480), IW (303-362 vs. 424), *e*₂ (15-17 vs. 46), longer *d*₂ (82-89 vs. 74), *e*₁ (30-35 vs. 20), *f*₂ (70-80 vs. 34), *c*₃ (63-72 vs. 54), *4a* (90-100 vs. 64), females, shorter IL (358-396 vs. 544), IW (313-375 vs. 416), *e*₂ (25-30 vs. 44), longer, *c*₃ (64-70 vs. 48); *C. troglobionta* sp. nov. differs from *C. viviannae* (Haitlinger, 1992), females, in shorter IL (358-396 vs. 424), *f*₂ (22-31 vs. 48), longer *c*₃ (64-70 vs. 44), description of the adult male of *C. viviannae* (Haitlinger, 1992) is not available; *C. troglobionta* sp. nov. differs from *C. carabicola* (Berlese, 1882), males, in shorter IL (250-341 vs. 418-440), longer IW (303-362 vs. 281-291), females, shorter IL (358-396 vs. 438-534).

**Discussion**

One of the most abundant and diverse invertebrates found in caves associated with bat guano are mites, which together with other microarthropods play an effective role in the food chain of caves, so known as subterranean plankton (Palacios-Vargas *et al.* 2011; Thurgate *et al.* 2011).
The most reported mite species richness related to bat guano belongs to the Trombidiformes, Sarcoptiformes and Mesostigmata orders of subclass Acari (Palacios-Vargas et al. 1998; Pellegrini and Ferreira 2013).

Except for a few families, not much data is available about the morphological cave adaptation of many groups of mites, but according to literature, obligatory cavernicoles (troglobionts) expose some troglomorphic characteristics such as loss of eyes, lack of pigmentation, reduction in body size, increase in the number and length of sensory organs, reduction in metabolic rate and increase in lifespan (Culver and Pipan 2013; Romero 2009). These characteristics may vary greatly in some families. The body size of some subterranean mite species is larger than that of soil species (Ducarme 2004).

Lack of eyes is common in all members of the family Canestriniidae. Although Vitzthum, 1924 had reported nut brown color for *Canestrinia sardica* (syn. of *Ciprusenia carabicola*), the lack of pigmentation has been observed in most of the family members including *Ciprusenia troglobionta* sp. nov. Moreover, all life stages of the new species have transparency in most body parts. With comparison in the male holotype of the genus members, *C. troglobionta* sp. nov. has the most reduction in idiosomal length along with an increase in the most of dorsal and ventral setae as the sensory organs. To logically comparison of measurements, we defined setae indices that indicate the percentage ratio of dorsal or ventral setae length to idiosomal length: (seta length/IL) * 100, as shown in Table 3. Due to a lack of access to *C. viviannae* adult males (the holotype is a female), indices have not been calculated for this species. Some measurements required for indices have been calculated based on the available drawings. According to the results of Table 3, among the total number of 24 setae compared, setae *si*, *scx*, *c*₁, *c*₂, *c*₃, *d*₁, *d*₂, *e*₁, *f*₂, *h*₁, *h*₂, *h*₃, *la*, *3a*, *4a*, *ps*₃ equal to 66.67% of all have the most index value in the new species that means the most setae elongation compared to other genus species. This value for other species is as follows: *C. carabicola* 0%, *C. izabelae* 0%, *C. kamelskyi* 29.17%, *C. sellnicki* 4.17%.

<table><caption>**Table 3** Setae indices comparison of *Ciprusenia* spp., adult males (exclude *C. viviannae*)</caption><thead><tr><th>Index</th><th>C. troglobionta sp. nov.</th><th>C. carabicola</th><th>C. izabelae</th><th>C. kamelskyi</th><th>C. sellnicki</th></tr></thead><tbody><tr><td>vi I</td><td>17.76</td><td>11.96</td><td>11.67</td><td>19.74</td><td>8.77</td></tr><tr><td>si I **</td><td>14.64</td><td>1.91</td><td>11.25</td><td>12.37</td><td>3.51</td></tr><tr><td>se I</td><td>112.77</td><td>94.01</td><td>73.96</td><td>118.42</td><td>52.63</td></tr><tr><td>scx I **</td><td>15.26</td><td>11.48</td><td>*</td><td>6.58</td><td>10.53</td></tr><tr><td>c<sub>1</sub> I **</td><td>9.97</td><td>2.39</td><td>6.67</td><td>2.63</td><td>3.51</td></tr><tr><td>c<sub>2</sub> I **</td><td>23.36</td><td>2.15</td><td>14.6</td><td>22.37</td><td>3.51</td></tr><tr><td>d<sub>1</sub> I **</td><td>9.66</td><td>2.63</td><td>4.17</td><td>2.63</td><td>3.51</td></tr><tr><td>d<sub>2</sub> I **</td><td>27.72</td><td>2.15</td><td>15.42</td><td>22.37</td><td>3.51</td></tr><tr><td>e<sub>1</sub> I **</td><td>10.9</td><td>1.91</td><td>4.17</td><td>4.47</td><td>3.51</td></tr><tr><td>e<sub>2</sub> I</td><td>3.51</td><td>7.18</td><td>9.58</td><td>11.84</td><td>5.26</td></tr><tr><td>f<sub>2</sub> I **</td><td>24.92</td><td>1.91</td><td>7.08</td><td>4.47</td><td>3.51</td></tr><tr><td>h<sub>1</sub> I **</td><td>131.46</td><td>107.65</td><td>*</td><td>126.31</td><td>70.17</td></tr><tr><td>h<sub>2</sub> I **</td><td>99.69</td><td>61.72</td><td>*</td><td>85.53</td><td>14.25</td></tr><tr><td>h<sub>3</sub> I **</td><td>39.87</td><td>16.75</td><td>12.5</td><td>23.68</td><td>17.54</td></tr><tr><td>c<sub>3</sub> I **</td><td>22.43</td><td>13.16</td><td>11.25</td><td>14.21</td><td>11.4</td></tr><tr><td>cp I</td><td>85.67</td><td>87.32</td><td>77.71</td><td>101.05</td><td>43.86</td></tr><tr><td>la I **</td><td>14.64</td><td>10.05</td><td>5.83</td><td>*</td><td>14.56</td></tr><tr><td>4b I</td><td>12.77</td><td>5.98</td><td>4.58</td><td>15.79</td><td>7.72</td></tr><tr><td>3a I **</td><td>40.17</td><td>20.81</td><td>*</td><td>19.74</td><td>11.05</td></tr><tr><td>4a I **</td><td>29.59</td><td>14.35</td><td>13.33</td><td>17.89</td><td>7.54</td></tr><tr><td>g I</td><td>8.72</td><td>9.09</td><td>7.08</td><td>11.31</td><td>7.37</td></tr><tr><td>ps<sub>3</sub> I **</td><td>5.92</td><td>4.78</td><td>3.12</td><td>3.68</td><td>3.51</td></tr><tr><td>ps<sub>2</sub> I</td><td>11.84</td><td>9.09</td><td>13.96</td><td>15.79</td><td>5.78</td></tr><tr><td>ps<sub>1</sub> I</td><td>12.46</td><td>12.2</td><td>10.42</td><td>9.21</td><td>13.16</td></tr></tbody></table>

<footnote>* Measurement information is not available.</footnote>

<footnote>** The indices that have the most value for *C. troglobionta* sp. nov.</footnote>
# Key to the world-known species of the genus Ciprusenia Haitlinger, 2022

## Male

<table><tr><td>1. Idiosomal length &lt; 400; setae d<sub>2</sub> &gt; 80</td><td>2</td></tr><tr><td>— Idiosomal length &gt; 400; setae d<sub>2</sub> &lt; 80</td><td>3</td></tr></table>

<table><tr><td>2. Hysterosoma without two symmetric diagonal furrows</td><td>C. kamelskyi (Samšiňák, 1970)</td></tr><tr><td>— Hysterosoma with two symmetric diagonal furrows</td><td>C. troglobionta sp. nov.</td></tr></table>

<table><tr><td>3. Setae c<sub>2</sub> longer than c<sub>1</sub> (at least twice)</td><td>C. izabelae (Haitlinger, 1992)</td></tr><tr><td>— Setae c<sub>2</sub> not longer than c<sub>1</sub></td><td>4</td></tr></table>

<table><tr><td>4. Idiosomal length &gt; 500</td><td>C. sellnicki (Samšiňák, 1965)</td></tr><tr><td>— Idiosomal length &lt; 500</td><td>C. carabicola (Berlese, 1882)</td></tr></table>

## Female

<table><tr><td>1. Idiosomal width &gt; 400</td><td>2</td></tr><tr><td>— Idiosomal width &lt; 400</td><td>3</td></tr></table>

<table><tr><td>2. Idiosomal length &gt; 700; Setae vi &gt; 60</td><td>C. sellnicki (Samšiňák, 1965)</td></tr><tr><td>— Idiosomal length &lt; 700; Setae vi &lt; 60</td><td>C. izabelae (Haitlinger, 1992)</td></tr></table>

<table><tr><td>3. Hysterosoma with two symmetric diagonal furrows</td><td>C. troglobionta sp. nov.</td></tr><tr><td>— Hysterosoma without two symmetric diagonal furrows</td><td>4</td></tr></table>

<table><tr><td>4. Setae si &lt; 25; setae d<sub>2</sub> &lt; 35</td><td>C. carabicola (Berlese, 1882)</td></tr><tr><td>— Setae si &gt; 25; setae d<sub>2</sub> &gt; 35</td><td>5</td></tr></table>

<table><tr><td>5. Setae vi &lt; 70; opisthosomal margin with obvious lobes</td><td>C. viviannae (Haitlinger, 1992)</td></tr><tr><td>— Setae vi &gt; 70; opisthosomal margin without obvious lobe</td><td>C. kamelskyi (Samšiňák, 1970)</td></tr></table>

Note: Lombardini (1950) described two species: *Coleoglyphus rosascostai* and *Grandiella rosascostai*. Haitlinger (1990c) transferred *C. rosascostai* to the genus *Hatohylla* Haitlinger, 1990. In Haitlinger's work (2022) *G. rosascostai* was confused with *H. rosascostai* and excluded from the genus *Grandiella* Lombardini, 1939. It is an error, *G. rosascostai* is a good species (Haitlinger 1990b, 2022; Lombardini 1950).

## Conflicts of Interest

The authors declare that they have no conflicts of interest.

## Acknowledgments

Special thanks to Mohammad Javad Ghasempoor and Dr. Yaser Bakhshi (Shiraz University, Iran) for accompanying the first author in caving and sampling. We are very grateful to Dr. Maxim Nabozhenko (Daghestan Federal Research Centre, Russia) for his kind help in identifying the host beetle. We are grateful to the anonymous reviewers for their valuable comments. The current study has been partly supported by Iran National Science Foundation (INSF) under grant number 95849576.
ORCID

Mojgan Sadat-Shojaei https://orcid.org/0000-0003-4104-407X

Ryszard Haitlinger https://orcid.org/0000-0002-5661-7099

Saber Sadeghi https://orcid.org/0000-0002-3183-8979

Mohammad Ali Akrami https://orcid.org/0000-0002-7561-9508

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