# Four new species of mites in Ologamasiphis and Holzmannia genera, and a new Juvaria subgenus defined (Parasitiformes: Parasitidae)

Wojciech Witaliński, Dariusz J Gwiazdowicz

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Wojciech Witaliński, Dariusz J Gwiazdowicz. Four new species of mites in Ologamasiphis and Holzmannia genera, and a new Juvaria subgenus defined (Parasitiformes: Parasitidae). Acarologia, 2023, Acarologia, 63 (3), pp.867-905. 10.24349/0v36-z6a2. hal-04171382

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# Four new species of mites in *Ologamasiphis* and *Holzmannia* genera, and a new *Juvaria* subgenus defined (Parasitiformes: Parasitidae)

Wojciech Witaliński<sup>D a</sup>, Dariusz J. Gwiazdowicz<sup>D b</sup>

## Original research

### ABSTRACT

The present study proposes the restructuring of *Ologamasiphis* genus (family Parasitidae) comprising two subgenera *Ologamasiphis* s. s. and *Holzmannia* Juvara-Bałş, 2002, to have the respective subgenera elevated to the genus level, as well as to have the brand-new *Juvaria* subgenus created within the *Holzmannia* genus. The proposed restructuring is based on two newly described species belonging to the genus *Holzmannia* and subgenus *Juvaria* n. subgen., to wit *Holzmannia* (*Juvaria*) *pilosa* n. sp. and *Holzmannia* (*Juvaria*) *silvestris* n. sp. Furthermore, two new species belonging to *Ologamasiphis* genus are described: *Ologamasiphis setosus* n. sp. and *Ologamasiphis zelkovae* n. sp. Pertinent modification of the existing key to the Parasitidae genera by Hrúzová & Fendá (2018) is also supplied.

**Keywords** mite taxonomy; new species; new subgenus; rare mite fauna; Romania; Greece; Austria; Karawanks Mts

**Zoobank** http://zoobank.org/AA59D10C-2546-4012-B221-94C29784F723

## Introduction

The Parasitidae family comprises two subfamilies, *Parasitinae* and *Pergamasinae*, as well as *Erithosoma*, a genus not assigned to any subfamily. In recent years, the concept of Parasitidae subfamilies was a subject to several changes (Hrúzová & Fendá, 2018; *Juvara-Bałş*, 2019; Makarova, 2019; Yao *et al.*, 2020; Yao *et al.*, 2022) leading to 47 genera in *Parasitidae*: 22 genera in *Parasitinae*, 24 genera in *Pergamasinae*, and *Erithosoma*.

The genus *Ologamasiphis* Athias-Henriot, 1971 belongs to *Pergamasinae* subfamily and comprises scarcely encountered mites. It contains two subgenera, *Ologamasiphis* s. s. and *Holzmannia* Juvara-Bałş, 2002. The main feature of *Ologamasiphis* females is a globular idiosoma and presternal plates (presternal sclerites) fused to the sternal shield, whereas the males are featured with femur II without axillary process and conspicuous structure of genital orifice located within the anterior part of the sternum instead of in front of sternal margin. For both sexes a characteristic feature is an absence of gland pores *gv1*. If present, they are located in the females on sternum close to its posterior margin, either behind *st3* setae or closer to sternal axial line, whereas in the males at the level between pores *iv2* and setae *st3* (see e.g. Witaliński, 2022). This genus has been divided into two subgenera by *Juvara-Bałş* (2002) who created *Holzmannia* subgenus. Both subgenera might be identified owing to several characteristics, the most evident of which are the following ones – in females peritrematal

Received 01 March 2023
Accepted 05 July 2023
Published 12 July 2023

Corresponding author
Wojciech Witaliński<sup>D</sup>:
w.witalinski@gmail.com

Academic editor
Faraji, Farid

https://doi.org/10.24349/0v36-z6a2

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

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

Witaliński W. and Gwiazdowicz D. J.

Licensed under
Creative Commons CC-BY 4.0

OPEN ACCESS

<footnote><sup>a</sup> Department of Comparative Anatomy, Institute of Zoology and Biomedical Research, Jagiellonian University, Gronostajowa 9, 30-387 Kraków, Poland.</footnote>

<footnote><sup>b</sup> Faculty of Forestry and Wood Technology, Poznań University of Life Sciences, Wojska Polskiego 71c, 60-625 Poznań, Poland.</footnote>
shield is posteriorly free and sternal gland pores *gv1* absent (subgenus *Ologamasiphis* s. s.), or in females peritrematal shield is fused with dorsal shield and gland pores *gv1* present (subgenus *Holzmannia* Juvara-Balş, 2002). However, within the recently studied material we encountered the specimens with different combination of characteristics, prompting us to have the *Ologamasiphis* genus restructured. Instead of genus *Ologamasiphis* with two subgenera *Ologamasiphis* and *Holzmannia*, we propose to have *Ologamasiphis* s. s. and *Holzmannia* Juvara-Balş, 2002 elevated to a generic level, as well as have the latter divided into the two subgenera: *Holzmannia* s. s. and *Juvaria n. subgen.* This concept effectively resolves the problem with imprecise Step 27 in the key to Parasitidae genera by Hrúzová & Fendá, 2018.

## Methods

Mites were routinely extracted from forest litter or rotten wood (*Ologamasiphis zelkovae* n. sp.) with the aid of Berlese funnels, into 70% ethanol, mounted in Hoyer's medium on glass slides, cured for several days in an oven (60 °C). All measurements are expressed in micrometres, taken as follows: idiosoma measurements were acquired along the sagittal line (length), and in the widest section (width) of holodorsal shield in the female, and at the margins of idiosoma in the male; setal lengths were measured from the alveolus up to the apex of the seta. The peritreme length was measured inclusive of the stigma. The drawings were made with the aid of Corel Draw X8 and a Wacom Intuos Graphic Tablet. The system of dorsal, ventral, palpal, and leg setal notations was based on Evans and Till (1979), whereas poroidotaxy and adenotaxy on Moraza and Peña (2005), with some necessary adjustments for Parasitidae. Legs I, II and IV in the females are showing some modified setae and are illustrated in ventral perspective with anterolateral side on the left. Leg III, as well as the tarsus I were not considered in detail as not informative enough, much like in the species under study and without modified setae. Furthermore, Ta I carries a lot of setae (more than 30) which are not identifiable. Ta IV was measured without apothele. Setae on the legs are numbered consecutively in a row from distal to proximal end of the article.

In all species both sexes were available for study and a female was chosen as the holotype, since the females in ventral view happen to be the more suitable for species identification than males. Developmental instars were not encountered.

## Results

Taxonomy and diagnoses of the taxa under study are proposed as follows:

### Genus *Ologamasiphis* Athias-Henriot, 1971

Type species: *Pergamasus epigynialis* Willmann, 1940: 211

### Diagnosis

Based on Juvara-Balş, 2002 and Hrúzová & Fendá, 2018.

**Female and male** — Idiosoma usually globular (in *O. zelkovae* n. sp. elliptical in dorsal perspective); podonotal region with 18–20 pairs of setae (*Ologamasiphis setosus* n. sp. 18; *O. zelkovae* n. sp. 18), opisthonotum with 19–22 pairs of setae (*O. setosus* n. sp. 19; *O. zelkovae* n. sp. 19); opisthogaster bears seven pairs of setae; adenotaxy with four pairs of podonotal and three opisthonotal gland pores; on ventral idiosoma *gv1* gland pores absent; setae *av1* and *pv1* on Ta IV can be different.

**Female** — Female with lateral presternal platelets, presternal plates either fused with the sternal shield completely, or more or less discernible as the separate elements in the anterior area of the sternal shield (in *O. zelkovae* n. sp.); epigynial shield irregularly pentagonal with
arcuate thickenings in midregion of lateral margins; peritrematal shield anteriorly fused with the margin of the podonotal shield and posteriorly free.

**Male** — Movable digit of chelicera with two teeth; genital opening located just behind the anterior sternal margin, containing a genital lamina; instead of separate subgenital sclerite, a cuticular ribbon joining the left and right parts of the sternum is continuous and located in front of the genital opening dorsally to genital lamina; Fe II main spur long and narrow, axillary process absent and additional spur located in the article distal region present.

## Species comprising

According to Athias-Henriot, 1967.

* *Ologamasiphis bulgatulus* (Athias-Henriot, 1967)
  *Pergamasus bulgatulus* Athias-Henriot, 1967: 134

* *Ologamasiphis epigynialis* (Willmann, 1940)
  *Pergamasus epigynialis* Willmann, 1940: 211

* *Ologamasiphis judaeortus* (Athias-Henriot, 1967)
  *Pergamasus judaeortus* Athias-Henriot, 1967: 135

* *Ologamasiphis parnethortus* (Athias-Henriot, 1967)
  *Pergamasus parnethortus* Athias-Henriot, 1967: 133

* *Ologamasiphis turdetanus* (Athias-Henriot, 1967)
  *Pergamasus turdetanus* Athias-Henriot, 1967: 134

The newly described species:

* *Ologamasiphis setosus* n. sp.
  female, male; Greece (Peloponnesus)

* *Ologamasiphis zelkovae* n. sp.
  female, male; Greece (Crete)

## Description of the new species

### *Ologamasiphis setosus* n. sp.

Zoobank: D624C028-1560-4802-8AC8-CA6ED6C4F8F9

(Figures 1–7)

## Diagnosis

**Female and male** — Idiosoma globular and well sclerotised; podonotum with 18 pairs of long setae, opisthonotum with 19 regular pairs of long setae, but one supplementary seta (Zx) may also be encountered; Co I with three anterolateral teeth and two anteroventral tubercles, Co III bears a thickening behind the posteroventral seta, Co IV with small posterolateral tubercle located distally; setae *av1* and *pv1* on Ta IV different.

**Female** — Gnathotectum with similar prongs, the central one somewhat longer, the lateral ones divergently directed; the sternal shield anterior margin with a shallow concavity flanked by the incorporated presternal plates, lateral presternal platelets free; paragynial shields with arcuate metagynial sclerites; epigynium anterior part not reticulated, the anterior prong without the axial thickening; dependent on orientation, endogynium oval (ventrally), or cup-shaped (laterally); cheliceral fixed digit with tooth antiaxially and three or four denticles followed by one or two larger teeth ahead of pilus dentilis, and three larger teeth behind it.

**Male** — Gnathotectum with the central prong larger than the lateral ones; anterior margin of the sternal plate convex; genital lamina with subrectangular anterior part, anterior corners rounded and the anterior margin concaved; cheliceral fixed digit features two teeth ahead and two behind pilus dentilis, the proximal-most the largest; leg II ventrally: femoral main spur narrow and slightly arcuate, conical additional spur located at the distal article margin, genual and tibial spurs conical, the genual one at distal article margin, whereas the tibial one located more proximally; leg II laterally: femoral main spur with slightly sinuous external margin,
<figure><img src="image_8.png" /><figcaption>**Figure 1** *Ologamasiphis setosus* n. sp., female (holotype): idiosoma dorsal side. Abbreviations: regular podosomal (*j*, *z*, *s*, *r*) and opisthosomal setae (*J*, *Z*, *S*, *R*); podosomal (*idj*, *idz*, *ids*) and opisthosomal pores (*idJ*, *idZ*, *idS*); gland openings on podonotum (*gdj*, *gdz*, *gds*) and opisthonotum (*gdZ*, *gdS*). Note the supplementary seta *Zx*.</figcaption></figure>

additional spur low, spurs on the genu and tibia somewhat arcuate and pointed, the tibial spur larger.

**Description**

**Female** (Figures 1–5, 7C–E)

**Idiosoma** (Fig. 1) — Globular, elliptic in dorso-ventral perspective, well sclerotised, 640 x 420 (length x width, n=1, holotype). Body incised at the fourth coxae level, opisthosomal part somewhat wider than the podonotal one. **Podonotum** – 18 pairs of long setae (setae *z3*, *s1*, *s2*, *s3*, *r1* and *r4* absent), setae length 46 (*j1*), 44 (*j2*), 76 (*j3*), 95 (*j4*), 75 (*j5*), 78 (*j6*), 133 (*r3*). **Opisthonotum** – with 19 pairs of long setae (in axial series setae *J1*, *J3* and *Z2* are absent, whereas in the more marginal series *S2* and *R2* are not encountered either). The number of setae on the left side is 19, since one *S6* seta is missing, but an additional seta in *Z*
<figure><img src="image_9.png" /><figcaption>**Figure 2** *Ologamasiphis setosus* n. sp., female (holotype): idiosoma ventral side. Abbreviations: I-IV the openings for coxae I to IV; *r2*, *r3* podonotal setae; *st1*–*st3* sternal setae; *st4*, *st5* paragynial and epigynial setae, respectively; *JV* and *ZV* opisthogastral setae series; *Aa* adanal setae; *Pa* postanal seta; *iv1*–*iv5*, *ivo1*–*ivo3*, *ivp* pore openings; *gv2*, *gv3* gland openings.</figcaption></figure>

series (Zx) is encountered. Setae length range from ca. 40 (R1) up to 96 (J5, Z6). Dorsal setae simple, podonotum not reticulated, opisthonotum with a scale-like reticulation. **Adenotaxy** – podonotum with four pairs of glands (*gdj2*, *gdz5*, *gdz6*, *gds4*) and opisthonotum with three pairs of glands (*gdZ4*, *gdZ6*, *gdS2*). **Peritreme** length 252, ending anteriorly in region of Co I, just behind *idz1* pore level.

**Ventral idiosoma** (Fig. 2) — Setae length in holotype: 51 (*st1*), 52 (*st2*), 52 (*st3*), 43 (*st4*), 42 (*st5*), 52 (*JV1*), 64 (*JV2*), 61 (*JV3*), 52 (*JV4*), 10 (*ZV1*), 55 (*ZV2*), 47 (*ZV3*), 29 (*Aa*), 13 (*Pa*). Setae *JV5* and *ZV4* located out of the opisthogastral shield on a soft cuticle. Ventral setae simple, reticulation of the opisthogaster scale-like. The sternal shield anterior margin with shallow concavity flanked by the elongated presternal plates tightly incorporated, lateral presternal platelets free. Gland pores *gv1* absent. Paragynial shields (Figs 2, 3A) with arcuate metagynial sclerites, epigynial shield (Figs 2, 3B) irregularly pentagonal in outline, anterior margins featuring the arcuate thickenings in the margin midregion, the posterior epigynial margin the longest and straight. Epigynial anterior part not reticulated. Endogynium (Figs 2, 3C, D) oval in the ventral perspective, but cup-shaped in the lateral (posterior) view (Fig. 3D). Pores *iv5* on soft cuticle behind epigynium, gland openings *gv2* singular, setae *ZV1* short and
<figure><img src="image_10.png" /><figcaption>**Figure 3** *Ologamasiphis setosus* n. sp., female (holotype): A – paragynium; B – epigynium; C and D – endogynium ventrally and caudally, respectively; E – gnathotectum; F – movable digit of chelicera in the ventral perspective; G – both fixed digits of chelicerae in the ventral view.
Abbreviations as in Figure 2.</figcaption></figure>

located at the level of the opisthogastral anterior margin. Pores *ivo2* on opisthogaster at *JV1* setae level, pores *ivo1*, *ivo3* and *ivp* out of opisthogaster on a soft cuticle. Gland pores *gv3* well discernible.

**Gnathosoma** — Gnathotectum (Fig. 3E) prongs similar, the central one longer and lateral ones divergently directed. Corniculi conical, hypostome with visible ca. eight rows of denticles, out of those the posterior most being semi-circular. Hypostomatic setae simple and similarly long, palpcoxal setae longer and barbed. Palptrochanter *v1* seta simple, *v2* barbed. Chelicera available only in dorsoventral perspective (Fig. 3F, G) – movable digit with three teeth (Fig. 3F), fixed digit (Fig. 3G) ahead of the pilus dentilis shows one tooth located antiaxially and three or four denticles adaxially, which are followed by one or two somewhat larger teeth. Behind the pilus dentilis there are three larger teeth.
<figure><img src="image_11.png" /><figcaption>**Figure 4** *Ologamasiphis setosus* n. sp., female (holotype), legs I and II in the ventral view except Ge I which is obliquely oriented: A – Co I; B – Co I and Tr I; C – Fe I; D – Ge I; E – Ti I; F – Co II, Tr II Fe II, Ge II and Ti II; G – Ta II. The arrow is on the anterolateral side. For abbreviations regarding the setae – see Methods.</figcaption></figure>
<figure><img src="image_12.png" /><figcaption>**Figure 5** *Ologamasiphis setosus* n. sp., female (holotype), leg IV ventrally: A – Co, Tr, Fe and Ge; B – Ta. For abbreviations regarding the setae – see Methods.</figcaption></figure>

**Legs** — Coxa I (Figs 4A, B, 7C) with three teeth on anterolateral surface and two tubercles located anteroventrally, out of those the proximal one weakly pronounced. Coxa III (Fig. 7D) with a thickening behind the posteroventral seta. Coxa IV with small posterolateral tubercle located distally (Fig. 7E). Leg setation – leg I (Figs 4A–E, 7C): seta *all* on Tr I and *pd3* on Fe I somewhat shorter and thicker, the latter one terminally barbed, seta *av1* on Fe I barbed. Leg II (Fig. 4F, G): setae *all* on Tr II and Fe II, as well as both anterolateral setae on Ge II and Ti II barbed. Some setae on Ta II barbed (Fig. 4G). Leg IV (Figs 5, 7E): seta *pd2* on the Fe IV (Fig. 5A) shorter and terminally barbed, seta *ad1* slightly longer than others (80–86), Ta IV (Fig. 5B) 251–255 long, tactile seta (*pd1*) very long (180–195), seta *av1* normal (short and thick),
<figure><img src="image_13.png" /><figcaption>**Figure 6** *Ologamasiphis setosus* n. sp., male: A – sternal shield, genital lamina absent; B – fine thickenings of sternal cuticle at the *iv3* pores level; C, D – anterior sternal margin with genital lamina, two aspects; E – genital lamina; F – anterior part of genital lamina; G – gnathotectum, two aspects; H – hypostomatic groove ventrally; I – chelicera, adaxially. Abbreviations: I–IV the openings for coxae I to IV; *st1–st5* sternal setae; *iv1–iv5* pore openings; *gv2* gland opening. Asterisk marks a genital opening in the anterior part of the sternum.</figcaption></figure>

whereas *pv1* thin and long. Other aspects of legs I–IV unremarkable.

**Male** (Figures 6, 7A, B, F–H)

**Idiosoma** — Globular and well sclerotised, 600–620 x 315–332 (length x width, n=5), body oval, slightly wider in posterior half. **Podonotum** — 18 pairs of long setae, setae length: 45–52 (*j1*), 52–54 (*j2*), 72–75 (*j3*), 86–92 (*j4*), 65–75 (*j5*), 73–78 (*j6*), 110–125 (*r3*). **Opisthonotum** — with long setae, setae length ranging from ca. 50 on anterolateral up to 78 on the posterior
<figure><img src="image_14.png" /><figcaption>**Figure 7** *Ologamasiphis setosus* n. sp., male (A, B, F, G, H) and female (C, D, E): A, B – Fe, Ge and Ti of leg II ventrally (A) and anterolaterally (B); C, F – Co I ventrally; D, G – Co III ventrally; E, H – Co IV ventrally. Some setae are marked. Arrows on the anterolateral side. C–E – holotype.</figcaption></figure>

part. Dorsal setae simple. Reticulation of the podonotum non-discernible, opisthonotum with a scale-like reticulation. **Peritreme** length 235–241, ending anteriorly as in the females.

**Ventral idiosoma** — Setae length: 35–44 (st1), 38–46 (st2), 38–42 (st3), 37–41 (st4), 31–34 (st5), 43–50 (JV1), 10–12 (ZV1), 24–29 (Aa), 14–17 (Pa), other opisthogastral setae ca. 44–58. **Ventral setae simple. Sternal region** (Fig. 6A–D) – presternal plates not visible separately, lateral presternal platelets free. Sternal shield convex anteriorly, with semilunar opening for the genital lamina. In the anterior part of the opening a sclerotised cuticular band
running dorsally to the genital lamina joins left and right parts of the sternum (Fig. 6A, C, D). The anterior part of genital lamina is sub-rectangular, with the anterior corners rounded and the anterior margin more or less concaved (Fig. 6C–F). Sternal gland pores *gv1* absent. In between setae *st3* and *st4* two pairs of fine thickenings of the sternal cuticle are encountered (Fig. 6A, B). Pore *gv2* with one or two openings, pores *iv5* relatively close to *ZV1* setae, at the same level as *gv2* (Fig. 6A). Pores *ivo1-ivo3*, *ivp* and *gv3* well discernible.

**Gnathosoma** — Gnathotectum (Fig. 6G) with central prong larger than the lateral ones, the latter occasionally curved. Corniculi conical, hypostome with ca. eight to nine rows of denticles, out of those the posterior rows are semicircular and the sixth row is made up of larger denticles (Fig. 6H). Hypostomal and palpcoxal setae as in the female. Palptrochanter *v1* seta simple, *v2* larger and one-sidedly barbed. Chelicera (Fig. 6I) – movable digit with two teeth followed by an arcuate edge proximally, fixed digit with two teeth ahead and two teeth behind the pilus dentilis, out of those the proximal one the largest and triangular.

**Legs** — Coxa I (Fig. 7F), Co III and Co IV (Fig. 7G, H) as in the female. Leg II (Fig. 7A, B) spurred as follows: when viewed from the ventral side (Fig. 7A), the femoral main spur is narrow and slightly arcuate, axillary process absent, additional conical spur is located at the distal article margin in line with the genual and tibial spurs. Genual and tibial spurs conical, the genual one is smaller and located closer to the distal article margin than the tibial one. In the lateral perspective (Fig. 7B), femoral main spur has the external margin slightly sinuous, an additional spur is low, spurs on the genu and tibia somewhat arcuate and pointed, the tibial one larger. Leg II setation: femoral anterolateral setae somewhat thickened and terminally barbed, seta *ad3* shorter and thickened, seta *pd2* barbed terminally. On Ge II and Ti II anterolateral setae barbed. Seta *ad1* on Fe IV as in the female (80–84), tactile seta (*pd1*) on Ta IV very long (180–185), setae *av1* and *pv1* different, as in the female. Other aspects of legs I–IV unremarkable.

**Material examined**

**Holotype** — Female (slide no. 2431 A), Peloponnese, Greece, 37.2898°N, 22.5107°E, alt. ca. 842 m a.s.l., 23 Oct. 2012, wet litter in the deciduous forest (chestnut, oak, maple and plane trees). Leg. Dr. M. Pabijan, Jagiellonian University, Kraków, Poland.

**Paratypes** — 6 males (slides no. 2431 B-E), ibid.

**Type deposition** — Types are deposited in the Zoological Division of the Nature Education Centre, Jagiellonian University, Kraków, Poland.

**Etymology**

The specific name *setosus* refers to the relatively long dorsal setae.

**Differential taxonomy**

As far as the length of Ta IV in the females (251-255) is concerned, *Ologamasiphis setosus* n. sp., as well as *Ologamasiphis zelkovae* n. sp. should be compared with the two species characterised by a similar Ta IV length (200-240), i.e. *Ologamasiphis parnethortus* (Athias-Henriot, 1967) and *Ologamasiphis turdetanus* (Athias-Henriot, 1967), rather than the species with much shorter Ta IV (160-175): *Ologamasiphis bulgatulus* (Athias-Henriot, 1967) and *Ologamasiphis judaeortus* (Athias-Henriot, 1967). *Ologamasiphis parnethortus* is characterised by long dorsal setae, as also encountered in *O. setosus* n. sp., and a similar reticulation of the sternum. Both species differ, however, in a distinct sclerotisation of the sternal anterior margin (strong in *O. setosus* n. sp., but weak in *O. parnethortus*), the shape of epigynial central prong (more slender in *O. setosus* n. sp.), a different shape of metagynial sclerites of the paragynia, and the shape of endogynium in the lateral perspective (roundish in *O. parnethortus* rather than a cuboidal one). Males differ mainly in leg II armature: Fe II main spur is laterally narrow and straight in *O. setosus* n. sp., but thicker and curved in *O. parnethortus*, Ge II and Ti II spurs are apically rounded in *O. parnethortus* and pointed in *O. setosus* n. sp.
In *O. turdetanus* only a female has been described; its dorsal setae are short, so they differ from those in *O. setosus n. sp*. Certain other differences are as follows: in *O. turdetanus* many lines of sternal and epigynial reticulations are wavy, whereas in *O. setosus n. sp*. are straight; central epigynial prong is much more slender in *O. setosus n. sp.*, and the shape of endogynium in the lateral perspective is different in both species.

**Ologamasiphis zelkovae n. sp.**

Zoobank: 38A4A1EO-C564-4B03-9358-EE7BD6D5B035

(Figures 8–14)

**Diagnosis**

**Female and male** — Idiosoma elliptical and weakly sclerotised; podonotum and opisthonotum with 18 and 19 pairs of moderately long setae, respectively; Co I with two distinct anterolateral teeth followed by the third smaller, tubercular tooth proximally; Co III with tubercular thickening directed postero-ventrally; setae *av1* and *pv1* on Ta IV different.

**Female** — Gnathotectum with similar narrow prongs, the central one longer; the sternal shield anterior margin with the V-shaped concavity, elongated presternal plates separate, but closely adhering to the sternal margin, lateral presternal platelets free; paragynial shields with arcuate metagynial sclerites; epigynial anterior prong with the axial thickening, the anterior part of epigynium not reticulated; endogynium circular or triangular in outline; cheliceral fixed digit with two teeth in front and two behind the pilus dentilis.

**Male** — Gnathotectum central prong larger and different in shape than the lateral ones; anterior margin of the sternal plate convex; genital lamina with rectangular anterior part, rounded corners and finely concaved margins; cheliceral fixed digit with two teeth ahead and two behind the pilus dentilis, second and third teeth are located close one to the other and separate by pilus dentilis, the fourth (proximal) one more distant and the largest; leg II laterally: femoral main spur with slightly sinuous external margin, additional spur low and located at the distal article margin, genual and tibial spurs conical and straight, genual one smaller than the tibial one and located closer to the distal article margin.

**Description**

**Female** (Figures 8–12, 14C, D)

**Idiosoma** — Elliptical and weakly sclerotised, 680–712 x 360–400 (length x width, n=5), holotype 683 x 429. **Podonotum** (Fig. 8) — 18 pairs of setae (setae *z3*, *s1*, *s2*, *s3*, *r1* and *r4* absent), setae length in paratypes: 39–48 (*j1*), 55–64 (*j2*), 70–82 (*j3*), 84–107 (*j4*), 59–67 (*j5*), 64–69 (*j6*), 118–122 (*r3*), in holotype 38 (*j1*), 65 (*j2*), 82 (*j3*), *j4–j6* not available, 115 (*r3*). **Opisthonotum** (Fig. 8) — with 19 pairs of setae, in axial series setae *J1*, *J3* and *Z2* are absent, whereas in the more marginal series *S2* and *R2* are not encountered either. Setae length range from ca. 57–62 (*R1*, *J2*) up to 94 (*Z6*). Dorsal setae simple, podonotum not reticulated, opisthonotum with a reticulation better visible in the lateral and posterior areas, between the podonotum and the opisthonotum a grove in some specimens may be observed. **Adenotaxy** — podonotum with four pairs of glands (*gdj2*, *gdz5*, *gdz6*, *gds4*) and the opisthonotum with three pairs of glands (*gdZ4*, *gdZ6*, *gdS2*). **Peritreme** length 275–285, in holotype 280, ending anteriorly in the region of Co I, just behind *idz1* pore level.

**Ventral idiosoma** (Fig. 9) — Setae length in paratypes: 42–48 (*st1*), 40–47 (*st2*), 43–50 (*st3*), 32–39 (*st4*), 30–37 (*st5*), 37–43 (*JV1*), 41–55 (*JV2*), 44–52 (*JV3*), 49–56 (*JV4*), 10–12 (*ZV1*), 35–44 (*ZV2*), 47–51 (*ZV3*), 20–23 (*Aa*), 17–20 (*Pa*). in holotype 55 (*st1*), 50 (*st2*), 47 (*st3*), 39 (*st4*), 35 (*st5*), 41(*JV1*), 43 (*JV2*), 51 (*JV3*), 55 (*JV4*), 12 (*ZV1*), 43 (*ZV2*), 47 (*ZV3*), 23 (*Aa*), 17 (*Pa*). Setae *JV5* and *ZV4* located out of the opisthogaster shield on a soft cuticle. Ventral setae simple, reticulation of the opisthogaster scale-like. The anterior margin of sternum with V-shaped concavity, in which the elongated presternal plates are separate, but closely adhering to the sternal margin (Figs 9, 10A). Lateral presternal platelets free. Gland
<figure><img src="image_15.png" /><figcaption>**Figure 8** *Ologamasiphis zelkovae n. sp.*, female: idiosoma dorsal side. Abbreviations as in Figure 1.</figcaption></figure>

pores gv1 absent. Paragynial shields (Figs 9, 10B) with arcuate metagynial sclerites, epigynial shield (Figs 9, 10C) irregularly pentagonal in outline. Epigynial anterior prong with a short thickening axially, the anterior margins divided with the lateral lenticular thickenings into the anterior and posterior parts. The posterior epigynial margin longest and straight or slightly arcuate. Epigynial anterior part not reticulated. Endogynium (Figs 9, 10D–F) either circular or triangular in outline, in the latter case the corners are rounded. Pores *iv5* close to the posterior margin of a soft cuticle behind the epigynium, gland openings *gv2* singular, setae *ZV1* short and located at the level of the opisthogastral anterior margin (Fig. 10C). Pores *ivo2* on opisthogaster in the middle between *JV1* and *JV2* setae levels, pores *ivo1*, *ivo3* and *ivp* out of the opisthogaster
<figure><img src="image_16.png" /><figcaption>**Figure 9** *Ologamasiphis zelkovae n. sp.*, female: idiosoma ventral side. Abbreviations as in Figure 2.</figcaption></figure>

on a soft cuticle. Gland pores *gv3* well discernible.

**Gnathosoma** — Gnathotectum (Fig. 10G) with similarly narrow prongs, the central one nearly twice longer. Corniculi conical, hypostome with eight to nine rows of denticles, out of those the posterior ones semicircular. Hypostomatic setae simple and similarly long, palpcoxal setae larger. Palptrochanter *v1* seta simple, *v2* barbed. Chelicera (Fig. 10H) movable digit with three teeth, out of those the proximal one the largest, fixed digit with two teeth ahead and two behind the pilus dentilis.

**Legs** — Coxa I (Figs 11A, 14C) with two distinct anterolateral teeth followed proximally by a third, small and tubercular tooth, as well as two tubercles located anteroventrally, the proximal one weakly pronounced. Coxa III (Fig. 14D) with a tubercular thickening directed postero-ventrally. Leg setation – leg I (Fig. 11A–D): seta *al1* on Tr I somewhat shorter, thicker and terminally barbed; anterolateral setae on Fe I short, posterodorsal seta *ad3* thicker and terminally barbed, setae *ad2* and *av1* barbed, seta *av2* very short; seta *al2* on Ge I barbed, seta
<figure><img src="image_17.png" /><figcaption>**Figure 10** *Ologamasiphis zelkovae n. sp.*, female: A – sternal shield, anterior part; B – paragynium; C – epigynium; D–F – endogynium, different aspects; G – gnathotectum; H – chelicera, antiaxially. Abbreviations as in Figure 3.</figcaption></figure>
<figure><img src="image_18.png" /><figcaption>**Figure 11** *Ologamasiphis zelkovae n. sp.*, female, legs I and II ventrally: A – Co I and Tr I; B – Fe I; C – Ge I; D – Ti I; E – Co II, Tr II, Fe II, Ge II; F – Ti II; G – Ta II. For abbreviations regarding the setae – see Methods.</figcaption></figure>
<figure><img src="image_19.png" /><figcaption>**Figure 12** *Ologamasiphis zelkovae n. sp.*, female, leg IV ventrally: A – Co and Tr; B – Fe, Ge and Ti; C – Ta. For abbreviations regarding the setae – see Methods.</figcaption></figure>

*pv2* on Ti I short and somewhat thickened. Leg II (Fig. 11E–G): seta *al1* on Tr II finely barbed; seta *al2* on Fe II thicker and terminally barbed, setae *ad1–ad3* short; some setae on Ta II bear one barb. Leg III: seta *pd2* on Fe III thicker and shorter. Leg IV (Fig. 12): setae *pd1* and *pd2* on Fe IV short, seta *ad1* of normal length (67–72); seta *al1* on Ti IV relatively small; Ta IV length 265-272, tactile seta (*pd1*) long (160–180), seta *av1* rather short and thickened, whereas *pv1* longer and not so thick. Other aspects of legs I–IV unremarkable.

**Male** (Figures 13, 14A, B, E, F)

**Idiosoma** — Elliptical and weakly sclerotised as in the females, 615–650 x 319–325
<figure><img src="image_20.png" /><figcaption>**Figure 13** *Ologamasiphis zelkovae n. sp.*, male: A – sternal shield; B – anterior sternal margin with genital lamina, another aspect; C – gnathotectum, two aspects; D – chelicera, antiaxially. Abbreviations as in Figure 6.</figcaption></figure>

(length x width, n=2). **Podonotum** – setae length: 35–38 (j1), 41–55 (j2), 81–86 (j3), 82–85 (j4), 56–59 (j5), setae j6 not available, 105–110 (r3). **Opisthonotum** – with long setae, setae length ranging from ca. 55 on the anterolateral up to 77–81 on the posterior part. Dorsal setae simple. Reticulation of the podonotum non-discernible, opisthonotum with a scale-like reticulation. **Peritreme** length 257–259, ending anteriorly as in the females.

**Ventral idiosoma** — Setae length: 41–46 (st1), 37–41 (st2), 35–42 (st3), 31–33 (st4), 24–27 (st5), 31–33 (JV1), 8–9 (ZV1), other opisthogastral setae ca. 26–41. Ventral setae simple. **Sternal region** (Fig. 13A) – presternal plates not visible separately, lateral presternal platelets free. Sternal shield convex anteriorly, with semilunar opening for the genital lamina. In the anterior part of the opening a sclerotised cuticular band running dorsally to the genital lamina joins the left and the right parts of the sternum (Fig. 13B). The anterior part of the genital lamina rectangular, with the anterior corners rounded and the slightly concaved anterior and lateral margins (Fig. 13A, B). Sternal gland pores gv1 absent. In between setae st3 and st4 two pairs of moderately pronounced thickenings of the sternal cuticle are encountered. Pore gv2 with one opening, pores iv5 between st5 and ZV1 setae, but closer to ZV1. Pores ivo1–ivo3, ivp and gv3 weakly discernible.

**Gnathosoma** — Gnathotectum (Fig. 13C) with the central prong more or less larger than the lateral ones which are of different shape. Corniculi conical, hypostome with ca. 8–9 rows of denticles, out of which the posterior rows are more curved. Hypostomatic and palpcoxal setae as in the female. Palptrochanter v1 seta simple, v2 larger and one-sidedly barbed. Chelicera (Fig. 13D) – movable digit with two teeth followed by a high arcuate protrusion proximally,
<figure><img src="image_21.png" /><figcaption>**Figure 14** *Ologamasiphis zelkovae n. sp.*, male (A, B, E, F) and female (C, D): A, B – Fe, Ge and Ti of leg II posterolaterally (A) and anterolaterally (B); C, E – Co I; D, F – Co III. Some setae are marked. Arrow on the anterolateral side.</figcaption></figure>

fixed digit with two teeth ahead and two teeth behind the pilus dentilis. Second and third teeth are located close one to the other and separate with pilus dentilis, the fourth (proximal) tooth more distant and the largest, followed by a margin with an incision.
**Legs** — Coxa I (Fig. 14E) and Co III (Fig. 14F) as in the female. Leg II (Fig. 14A, B) spurred as follows: the femoral main spur narrow, with slightly sinuous external margin, axillary process absent, additional spur low and located at the distal article margin, the genual and tibial spurs conical and straight, the genual one smaller than the tibial one and located closer to the distal article margin. Leg setation – leg I: seta *al1* on Tr I somewhat shorter and thicker, setae *al2* and *av2* on Fe I short, whereas seta *pd3* thickened and terminally barbed. Leg II (Fig. 14A, B): femoral anterolateral setae slightly thickened and terminally barbed, seta *ad3* short and thickened. Leg IV: setae *pd1* and *pd2* on Fe IV shortened, seta *ad1* of normal length (60–64); tactile seta (*pd1*) on Ta IV long (150–155), setae *av1* and *pv1* different, as in the female. Other aspects of legs I–IV unremarkable.

**Material examined**

**Holotype** — Female (slide no. 2990 A), Gerakari, Mt. Kedros, Crete, Greece, 35.1948°N, 24.6067°E, alt. ca. 1255 m a.s.l., 11 Oct. 2018, rotting wood from a crack in *Zelkova abelicea* (Lam.) Boiss. tree. Leg. D.J. Gwiazdowicz.

**Paratypes** — 8 females, 2 males (slides no. 2990 B-D, 2991, 2992 A), ibid.

**Type deposition** — Types are deposited in the Zoological Division of the Nature Education Centre, Jagiellonian University, Kraków, Poland.

**Etymology**

The specific name *zelkovae* refers to *Zelkova abelicea* (Lam.) Boiss, a tree endemic to Crete, where the specimens were encountered.

**Differential taxonomy**

The outline of *Ologamasiphis zelkovae n. sp.* body is evidently elongated rather than globular, hence it substantially differs from the species with the similar length of Ta IV, *O. setosus n. sp.*, *O. parnethortus* and *O. turdetanus*. The other unique feature in *O. zelkovae n. sp.* female are the remnants of the presternal plates incorporated in the anterior margin of the sternal shield. No less characteristic is a central prong of the epigynium with the axial thickening under the anterior tip. As far as the males are compared, the armature of leg II is similar in *O. parnethortus* and *O. zelkovae n. sp.*, but the femoral main spur, encountered in the latter species, is straight. *Ologamasiphis zelkovae n. sp.* and *O. setosus n. sp.* differ in their chelicerae structure: in both species the fixed digit features four teeth, but in the former species, the second and the third tooth is closer to pilus dentilis, whereas in *O. setosus n. sp.* only the second proximal tooth is close to pilus dentilis. Furthermore, Co IV in *O. setosus n. sp.* features a small posterolateral tubercle located distally, which is absent in *O. zelkovae n. sp*.

**Genus Holzmannia Juvara-Balş, 2002**

Type species: *Pergamasus disfistulatus* Athias-Henriot, 1967: 135

**Diagnosis**

Based on subgenus *Holzmannia* Juvara-Balş, 2002 in Hrúzová & Fendá (2018).

**Female and male** — Idiosoma globular; podonotal region with 16–17 pairs of setae, opisthonotum with 12–20 pairs of setae, opisthogaster with seven pairs of setae; adenotaxy with four pairs of podonotal and three opisthonotal gland pores; on ventral idiosoma *gv1* gland pores present or absent; setae *av1* and *pv1* on Ta IV different.

**Female** — Presternal plates completely fused with the sternal shield; lateral presternal platelets present, epigynial shield irregularly heptagonal; peritrematal shield fused with podonotal shield margin.

**Male** — movable digit of chelicera with two teeth; general structure of genital orifice region and leg II armature as in *Ologamasiphis* genus, but a sclerotised cuticular band running dorsally to the genital lamina comprises the left and the right part.
**Subgenus Holzmannia s. s**

Type species: *Pergamasus disfistulatus* Athias-Henriot, 1967: 135

**Diagnosis**

**Female and male** — Podonotal region with 16 pairs of setae, opisthonotum with 20 pairs of setae; on ventral idiosoma *gv1* gland pores present; other characteristics as in the genus.

**Species comprising**

* *Holzmannia* (*H.*) *disfistulatus* (Athias-Henriot, 1967)

*Pergamasus disfistulatus* Athias-Henriot, 1967: 135

*Ologamasus* (*Ologamasiphis*) *minimus*: Holzmann, 1969: 49; Athias-Henriot, 1971: 174

*Ologamasiphis disfistulatus*: Juvara-Balş, 1975: 402

Species content in *Holzmannia* (*Holzmannia*) requires further studies: *Holzmannia* (*H.*) *disfistulatus* (Athias-Henriot, 1967) sensu Juvara-Balş, 1975: 402 is, accordingly to Juvara-Balş (2002: 44) probably a new species, which is still not described. Moreover, Athias-Henriot (1971: 174) considered *Ologamasus* (*Ologamasiphis*) *minimus* synonymous with *Holzmannia* (*H.*) *disfistulatus*. However, in Juvara-Balş (2002: 43) and our opinion in both species some differences acknowledged. Thus status of *Ologamasus* (*Ologamasiphis*) *minimus* should be detailed considered in future.

**Subgenus Juvaria n. subgen.**

Type species: *Holzmannia* (*Juvaria*) *pilosa* n. sp.

**Etymology**

The subgenus name is dedicated to the eminent acarologist Ilinca Juvara-Balş, who designed
the *Holzmannia* genus.

**Diagnosis**

**Female and male** — Podonotal region with 16–17 pairs of long or very long setae, opisthonotum with 12–13 pairs of either short or very long setae; on ventral idiosoma *gv1* gland pores absent; other characteristics as in the genus.

**Species comprising - newly described species**

* *Holzmannia* (*Juvaria*) *pilosa* n. sp., female, male, Romania

* *Holzmannia* (*Juvaria*) *silvestris* n. sp., female, male, Austria

**Description of the new species**

**Holzmannia (Juvaria) pilosa n. sp.**

Zoobank: 11148DE3-C743-4CD6-A770-3952ED736170

(Figures 15–21)

**Diagnosis**

**Female and male** — Idiosoma well sclerotised, globular and oval in dorso-ventral perspective; podonotum and opisthonotum with 17 and 13 pairs of setae, respectively, most of them very long and curved; Tr IV with more or less distinct tubercle located proximally.

**Female** — Gnathotectum with similar prongs, the central one somewhat longer, between the central and lateral ones minute denticles may be encountered; the sternal shield anterior margin thickened, with concavity just behind the tritosternum base; paragynial shields with arcuate metagynial sclerites; epigynial shield anterior prong with convex margins ending with lenticular thickenings, a transversal, not pigmented band of cuticle discernible just behind
<figure><img src="image_22.png" /><figcaption>**Figure 15** *Holzmannia (Juvaria) pilosa* n. sp., female, idiosoma dorsal side. Abbreviations as in Figure 1. Note the peritremes and margin of the holodorsal shield located ventrally.</figcaption></figure>

the thickenings level; endogynium oval; ventral setae rather short; cheliceral fixed digit with two-three teeth in front and two behind the pilus dentilis.

**Male** — Gnathotectum with the central prong somewhat larger than the lateral ones, the latter usually curved; anterior margin of the sternal plate slightly convex; left and right parts of sclerotised cuticular band running dorsally to the genital lamina are axially connected with arcuate ribbon; genital lamina sub-rectangular, with concaved anterior margin and rounded corners; cheliceral fixed digit with one tooth ahead of the pilus dentilis, and a pair of teeth with pilus dentilis inserted in between, broader and lamellar tooth is present proximally; leg II ventrally: femoral main spur narrow and curved, distal additional spur conical and located postero-ventrally, genual spur bacillary and the tibial one conical; leg II laterally: femoral main spur straight, finger-shaped, at the base with convexity of external margin, additional spur rounded and located not far from the distal article margin, genual and tibial spurs elongated, whereas the genual one located closer to the distal margin of the article.

## Description

**Female** (Figures 15–19)

*Idiosoma* (Fig. 15) — Well sclerotised, globular and oval in dorso-ventral perspective,
<figure><img src="image_23.png" /><figcaption>**Figure 16** *Holzmannia (Juvaria) pilosa* n. sp., female: A – idiosoma ventral side; B – location of prosomal pore *ids4* and gland opening *gds4*. Abbreviations as in Figure 2.</figcaption></figure>

485–500 x 325–355 (length x width, n=5), holotype 471 x 343. **Podonotum** – 17 pairs of setae (setae z3, s1, s2, s3, r1, r4 and r6 absent), most of them very long and curved. Setae length in paratypes: 34–39 (j1), 34–39 (j2), 65–72 (j3), 98–105 (j4), 92–97 (j5), 89–93 (j6), 125–131 (r3), in holotype 39 (j1), 39 (j2), 69 (j3), 104 (j4), 93 (j5), 92 (j6), 124 (r3). **Opisthonotum** – with 13 pairs of very long curved setae (100–118), except the lateral ones, which are much shorter (52–59), or only ca. 23 (R1). In axial series setae J1, J3, Z2 and Z5 are absent, whereas in the more marginal series (S, R) seven pairs are not encountered either. However, their precise notation is obvious. Dorsal setae simple, podonotum and opisthonotum not reticulated. **Adenotaxy** – podonotum with four pairs of glands (gdj2, gdz5, gdz6, gds4) and opisthonotum with three pairs of glands (gdZ4, gdZ6, gdS2). **Peritreme** length 195–197 in paratypes and 192 in holotype, ending anteriorly in the region of Co I, just behind *idz1* pore level.

**Ventral idiosoma** (Fig. 16) — Setae length in paratypes: 39–41 (st1), 41–42 (st2), 34–39 (st3), 31–33 (st4), 21–24 (st5), opisthogaster with 7 pairs of setae, ZV1 minute (ca. 5), other opisthogastral setae short, 25 to 31, 18–21 (Aa), 11–12 (Pa), in holotype 42 (st1), 43 (st2), 40 (st3), 33 (st4), 22 (st5), 31 (JV1), 25 (JV2), 26 (JV3), 26 (JV4), 5 (ZV1), 22 (ZV2), 24 (ZV3), 20 (Aa), 10 (Pa). Setae JV5 and ZV4 located out of the opisthogaster shield on a soft cuticle.
<figure><img src="image_24.png" /><figcaption>**Figure 17** *Holzmannia (Juvaria) pilosa* n. sp., female: A – paragynium; B – metagynial thickenings of paragynia; C – epigynium; D – endogynium, two aspects; E – gnathotectum, different aspects; F – chelicera, antiaxially; G, H – Co I, three different orientations, showing proximal tubercle rounded (G) or pointed (H); I – Tr IV with a differently pronounced tubercle. Abbreviations as in Figure 3.</figcaption></figure>

Ventral setae simple, reticulation of the opisthogaster scale-like.

The anterior margin of the sternum wavy and thickened, with concavity just behind the tritosternum base, separate presternal plates absent, lateral presternal platelets free (Fig. 16). Gland pores gv1 absent. Paragynial shields (Figs 16, 17A) with moderately thick and arcuate metagynial sclerites (Fig. 17A, B), epigynial shield heptagonal rather than pentagonal (Figs 16, 17C), its anterior, central prong subtriangular, with more or less convex margins, followed by a quadrilateral posterior epigynial part. In places, where the margins of anterior prong pass into the posterior epigynial part, there is a pair of lenticular thickenings (Fig. 17C). The posterior epigynial margin straight. The anterior prong and the rest of the epigynium are separated with a
<figure><img src="image_25.png" /><figcaption>**Figure 18** *Holzmannia (Juvaria) pilosa* n. sp., female, legs I and II ventrally: A – Co I, Tr I and Fe I; B – Ge I; C – Ti I; D – Co II and Tr II; E – Fe II; F – Ge II; G – Ti II and Ta II. For abbreviations regarding the setae – see Methods.</figcaption></figure>

transversal band of less pigmented cuticle. Endogynium (Figs 16, 17D) oval transversally with a ventrally located opening. Pores *iv5* far from the posterior epigynial margin on a soft cuticle at the *ZV1* setae level, gland openings *gv2* singular. Pores *ivo2* on opisthogaster anteriorly to the *JV1* setae level, pores *ivo1*, *ivo3* and *ivp* out of opisthogaster on a soft cuticle (Fig. 16). Gland pores *gv3* may be discernible.

**Gnathosoma** — Gnathotectum (Fig. 17E) with similar prongs, the central one somewhat longer, frequently between the central and lateral prongs some minute denticles are encountered. Corniculi conical, hypostome with 10 rows of denticles, out of those the posterior-most one is semicircular. Hypostomatic and palpcoxal setae simple. Palptrochanter *v1* seta simple, *v2* barbed. Chelicera (Fig. 17F) with relatively slender digits, the movable digit with three teeth, the proximal one somewhat larger, fixed digit with two-three teeth ahead and two larger teeth behind the pilus dentilis.

**Legs** — Coxa I (Figs 17G, H, 18A) with antero-laterally located three teeth. Between the teeth and the ventral setae there are two tubercles, the proximal one slightly larger and sometimes pointed (Fig. 17H). Trochanter IV with more or less distinct posterolateral tubercle
<figure><img src="image_26.png" /><figcaption>**Figure 19** *Holzmannia (Juvaria) pilosa* n. sp., female, leg IV ventrally: A – Co, Tr and Fe; B – Ge and Ti; C – Ta. For setae abbreviations – see Methods.</figcaption></figure>

(Figs 17I, 19A). Leg setation – leg I (Fig. 18A–C): seta *v1* on Co I finely barbed, seta *al1* on Tr I somewhat shorter and thicker, whereas postero-dorsal short; setae *al1*, *al2* and *ad3* on Fe I short, seta *av2* very short. Leg II (Fig. 18D–H): setae *al1*, *al2* and *ad3* on Fe II slightly thickened; seta *al1* on Ti II barbed; setae *av2*, *al2*, *al3* and *pl2* on Ta II with one barb in the central part. Leg IV (Fig. 19): setae *pd1* and *pd2* on Fe IV small, seta *ad1* elongate (120–125); setae *pl1* and *pd3* on Ge IV small; Ta IV ca. 244 long, tactile seta (*pd1*) very long (190–195), seta *av1* short and thickened, whereas *pv1* twice longer and slim. Other aspects of legs I–IV unremarkable.
<figure><img src="image_27.png" /><figcaption>**Figure 20** *Holzmannia (Juvaria) pilosa* n. sp., male: A – sternal shield; B – thickening of the anterior sternal margin; C – genital lamina; D – gnathotectum, two aspects; E – aberrant gnathotectum; F – chelicera, antiaxially. Abbreviations as in Figure 6.</figcaption></figure>

**Male** (Figures 20, 21)

**Idiosoma** — Well sclerotised, globular and oval in outline, 425–445 x 280–290 (length x width, n=4). **Podonotum** — setae length: 28–33 (j1), 26–30 (j2), 59–65 (j3), 90–93 (j4), 72–78 (j5), 69–75 (j6), 112–118 (r3). **Opisthonotum** — setae length from ca. 51 up to 112, the shortest in the anterolateral parts. Dorsal setae simple. Reticulation of dorsal cuticle non-discernible. **Peritreme** length 186–193, ending anteriorly as in the females.

**Ventral idiosoma** — Setae length: 26–29 (st1), 27–30 (st2), 26–27 (st3), 22–24 (st4), 18–20 (st5), 21–22 (JV1), 2–3 (ZV1), other opisthogastral setae ca. 17–24. Ventral setae simple. **Sternal region** (Fig. 20A) – presternal plates not present, lateral presternal platelets free. Sternal shield anteriorly with semilunar opening for the genital lamina. In the anterior part of the opening, a sclerotised cuticular band running dorsally to the genital lamina comprises the left and the right parts connected axially with arcuate ribbon (Fig. 20A, B). The genital lamina rectangular, with the anterior corners rounded and a concaved anterior margin (Fig. 20A, C). Sternal gland pores gv1 absent. In between setae st3 and st4 two pairs of weakly pronounced thickenings of the sternal cuticle are present. Gland pore gv2 with one opening, pores iv5 between st5 and ZV1 setae, but distinctly closer to ZV1. Pores ivo1–ivo3, ivp and gland pores gv3 well discernible.

**Gnathosoma** — Gnathotectum (Fig. 20D) with the central prong triangular and somewhat larger than the lateral ones, which are triangular or more frequently curved adaxially. In one specimen the one-prong type gnathotectum was observed as an abnormality (Fig. 20E). Corniculi conical, hypostome with ca. nine rows of denticles. Hypostomatic and palpcoxal setae simple. Palptrochanter v1 seta simple, v2 barbed. Chelicera (Fig. 20F) – movable digit with two teeth followed proximally by a low arcuate edge, the proximal tooth larger and slightly rounded. Fixed digit with one tooth ahead of the pilus dentilis, and a pair of teeth with the pilus dentilis inserted between the teeth in the pair. Proximally broader and lamellar tooth is present.

**Legs** — Coxa I (Fig. 21C) and Tr IV (Fig. 21D) as in the female. Leg II (Fig. 21A,
<figure><img src="image_28.png" /><figcaption>**Figure 21** *Holzmannia (Juvaria) pilosa* n. sp., male: A, B – Fe, Ge and Ti of leg II ventrally (A) and anterolaterally (B) with the spurs in different aspects; C – Co I; D – Co IV. Some setae are marked. Arrows on the anterolateral side.</figcaption></figure>

B) spurred as follows: when viewed from the ventral side (Fig. 21A), femoral main spur is narrow and curved, additional spur conical and located postero-ventrally near the distal femoral margin, genual spur bacillary and the tibial one conical. In the lateral perspective (Fig. 21B), femoral main spur straight, finger-shaped, with convexity of the external margin at the base. The additional spur is rounded and located not far from the distal margin of the article, genual and tibial spurs are elongated and the former one located closer to the distal margin of the article. Leg setation – legs I, III as in the female. Leg II (Fig. 21A, B): setae *al1*, *al2*, *ad2* and *ad3* on Fe II short and thickened, *al1* and *al2* terminally barbed. Leg IV: setae *pd1* and *pd2* on Fe IV short, seta *ad1* elongate (114–117), tactile seta (*pd1*) on Ta IV very long (179–182), some setae with one or several barbs. Other aspects of legs I–IV unremarkable.

**Material examined**

**Holotype** — Female (slide no. 2253 A), Apuseni Mts, Transylvania, Romania, 46.3295°N, 23.0618°E, alt. ca 570 m a.s.l., 12 April 2010, litter in a common hornbeam forest. Leg. Dr. M. Pabijan, Jagiellonian University, Kraków, Poland.

**Paratypes** — 6 females, 6 males (slides no. 2249–2252 A-C, 2253 B,C, 2254), ibid.

**Type deposition** — Types are deposited in the Zoological Division of the Nature Education
Centre, Jagiellonian University, Kraków, Poland.

## Etymology

The specific name *pilosa* refers to the distinctly long dorsal setae.

## Differential taxonomy

Differential taxonomy for *H. (J.) pilosa n. sp.* is addressed further below, jointly with *H. (J.) silvestris n. sp.*.

## Holzmannia (Juvaria) silvestris n. sp.

Zoobank: 7F5E9B78-FF09-4019-A28D-A4047D12F94F

(Figures 22–28)

## Diagnosis

**Female and male** — Idiosoma moderately to well sclerotised, pyriform and oval in both female and male; podonotum with 17 pairs of moderately long setae, opisthonotum with 12–13 pairs of much shorter setae; Co III with posterolateral tubercle, Tr IV with posterolateral distinct tubercle located proximally.

**Female** — The sternal shield anterior margin thickened centrally, with concavity just behind the tritosternum base; lateral presternal platelets present; paragynial shields with anteriorly divergent metagynial sclerites terminally s-shaped; epigynial shield anterior prong with convex margins and arcuate thickenings, connected by a wavy, sclerotised line; endogynium circular in the ventral perspective, and cup-shaped when viewed from the posterior idiosoma end; sternal setae moderately long, opisthogastral ones short; cheliceral fixed digit with two teeth ahead and two behind the pilus dentilis, the proximal one the largest.

**Male** — Anterior margin of the sternal plate thickened, flanked with concavities facing to Co I; semicircular genital opening containing the genital lamina located in the anterior part of the sternum, sclerotised cuticular band running dorsally to the genital lamina more or less distinctly split in axial line; anterior part of the genital lamina is sub-rectangular, with more or less concaved anterior margin and rounded anterior corners; cheliceral fixed digit with two teeth ahead of the pilus dentilis, and one tooth behind it, followed by a larger, lamellar tooth proximally; leg II ventrally: femoral main spur narrow and slightly curved, distal additional spur conical and located postero-ventrally, genual and tibial spurs conical and pointed; leg II laterally: femoral main spur straight, at the base with convexity of the external margin, additional spur rounded and located close to the distal article margin, genual and tibial spurs hooked and pointed, the genual one located closer to the distal margin of the article than the tibial spur.

## Description

### Female (Figures 22–26)

**Idiosoma** (Fig. 22) — Pyriform, moderately to well sclerotised, 455–495 x 330–355 (length x width, n=5), holotype 476 x 325. **Podonotum** — 17 pairs of setae (setae *z3*, *s1*, *s2*, *s3*, *r1*, *r4* and *r6* absent), setae length variable, setae *j3–j6*, *z4*, *z5* and *r3* evidently longer than the others, axial setae in paratypes as follows: 30–33 (*j1*), 23–26 (*j2*), 51–55 (*j3*), 59–68 (*j4*), 44–50 (*j5*), 38–45 (*j6*), setae *r3* 97–102, in holotype 31 (*j1*), 26 (*j2*), 52 (*j3*), 60 (*j4*), 38 (*j6*), *j5* and *r3* not available. **Opisthonotum** — with ca. 13 pairs of setae, in axial series setae *J1*, *J3*, *Z2* and *Z5* are absent, whereas in the more marginal series (*S*, *R*) seven pairs are not encountered either. However, their precise notation is obvious. In general, opisthonotal setae are short and their length ranges from ca. 10–12 (lateral setae) up to 19–27 (axial setae). Dorsal setae simple, reticulation on the podonotum and the opisthonotum not discernible, despite a narrow marginal parts of the podonotum. **Adenotaxy** — podonotum with four pairs of glands (*gdj2*, *gdz5*, *gdz6*, *gds4*); opisthonotum with three pairs of glands (*gdZ4*, *gdZ6*, *gdS2*). **Peritreme** is running along
<figure><img src="image_29.png" /><figcaption>**Figure 22** *Holzmannia (Juvaria) silvestris* n. sp., female, idiosoma dorsal side. Note the peritremes, holodorsal shield margin, pores *ids4* and *gds4* located ventrally. Abbreviations as in Figure 1.</figcaption></figure>

podonotal margin and its length is 202–208 in paratypes and 209 in holotype, ending anteriorly in the region of Co I, just behind the *idz1* pore level.

**Ventral idiosoma** (Fig. 23) — Sternogenital setae length in paratypes: 36–40 (*st1*), 34–39 (*st2*), 34–37 (*st3*), 29–31 (*st4*), 23–26 (*st5*), opisthogaster with seven pairs of setae, *ZV1* minute (3–5), length of other opisthogastral setae 22–25 (*JV1*), 21–24 (*JV2*), 18–22 (*JV3*), 18–20 (*JV4*), 4–6 (*ZV1*), 15–18 (*ZV2*), 11–13 (*ZV3*), 18–22 (*Aa*), 10–12 (*Pa*), in holotype 34 (*st1*), 33 (*st2*), 33 (*st3*), 28 (*st4*), 24 (*st5*), 24 (*JV1*), 23 (*JV2*), 21 (*JV3*), 18 (*JV4*), 3 (*ZV1*), 14 (*ZV2*), 18 (*Aa*), 12 (*Pa*), *ZV3* not available. Ventral setae simple, reticulation of the sternum (Fig. 23) scale-like, reticulation on the opisthogaster discernible in the anterior part and in the perianal area only. The anterior margin of the sternal shield with a central concavity just behind tritosternum and two shallow concavities behind coxeae I. Presternal plates incorporated into the sternal shield, small lateral platelets present. The pores *gv1* absent. Paragynial shields (Figs 23, 24A) with divergent metagynial sclerites distinctly sigmoidal in the anterior part. Epigynial shield (Figs 23, 24B) anterior prong with convex margins ending with two arcuate sclerotisations connected with a wavy line. Endogynium (Figs 23, 24C, D) circular in the ventral perspective, with some non-distinct internal structures, but cup-shaped when viewed from the posterior idiosoma end (Fig. 24D). Gland pores *gv2* with one opening, *iv5* poorly visible, on a soft cuticle behind epigynium, *ivo2* well visible on the opisthogaster, *ivo1*, *ivo3* and *ivp* located out of the shield
<figure><img src="image_30.png" /><figcaption>**Figure 23** *Holzmannia (Juvaria) silvestris n. sp.*, female: idiosoma ventral side. Abbreviations as in Figure 2, *s6* podonotal seta.</figcaption></figure>

on a soft cuticle, *gv3* non-discernible. Pore *iv5*, microseta *ZV1* and gland pore *gv2* located close to the others in a horizontal row (Figs 23, 24B).

**Gnathosoma** — Gnathotectum (Fig. 24E) trispinate, all prongs similar, narrow and acute, the central one somewhat longer. Corniculi conical, hypostome with 11 rows of denticles, hypostomatic and palpcoxal setae simple, hypostomatics similarly long, palpcoxal setae somewhat longer. Palptrochanter *v1* seta simple, *v2* barbed. Chelicera (Fig. 24F) — movable digit with three teeth, the proximal one slightly larger, fixed digit with two teeth ahead and two teeth behind the pilus dentilis, followed by undulate lamellar margin. The proximal tooth somewhat larger than others.

**Legs** — Coxa I (Figs 24G, 25A) with anterolaterally located three teeth, between teeth and ventral setae there are two tubercles. Coxa III with posterolateral tubercle (Fig. 24H), Tr IV with posterolateral distinct tubercle located proximally (Figs 24I, 26A). Leg setation — leg I (Fig. 25A–C): seta *al1* on Tr I shorter, setae *av2* and *ad3* on Fe I very short, setae *al1* and *al2* twice smaller than the posterolateral setae, seta *av1* barbed. Leg II (Fig. 25D–F): setae *al1*, *al2*, *ad2* and *ad3* on Fe II shorter and thicker, anterolateral setae on Ge II and Ti II barbed. Seta *al2* on Ta II features one barb located centrally. Leg IV (Fig. 26): setae *pd1* and *pd2* on Fe IV shorter, posterolateral setae on Ge IV and Ti IV small; Ta IV length ca. 231, seta *pl3* may be
<figure><img src="image_31.png" /><figcaption>**Figure 24** *Holzmannia (Juvaria) silvestris n. sp.*, female: A – paragynium; B – epigynium; C, D – endogynium ventrally (C) and in the caudal perspective (D); E – gnathotectum; F – chelicera, antiaxially; G – Co I, ventrally; H – Co III, ventrally; I – Tr IV, ventrally. Abbreviations as in Figure 3.</figcaption></figure>

barbed on both sides, *av2* and *pv2* with one barb in the central part, seta *pv1* somewhat longer than *av1*, seta *ad1* length 85–88, tactile seta (*pd1*) very long (190–197). Other aspects of legs I–IV unremarkable.

**Male** (Figures 27, 28)

**Idiosoma** — Oval and sclerotised as in the female, 435–445 x 280–290 (length x width, n=5). **Podonotum** — setae length: 25–30 (*j1*), 24–26 (*j2*), 27–31 (*j3*), 50–58 (*j4*), 33–42 (*j5*), 33–37 (*j6*), 90–97 (*r3*). **Opisthonotum** — setae length from ca. 14 up to 21. Dorsal setae simple. Reticulation of the dorsum non-discernible. **Peritreme** length 199–203, ending anteriorly as in the females.

**Ventral idiosoma** — Setae length: 26–29 (*st1*), 27–30 (*st2*), 29–33 (*st3*), 26–29 (*st4*), 20–24 (*st5*), 18–21 (*JVI*), 4–5 (*ZVI*), 16–20 (*Aa*), 13–16 (*Pa*), other opisthogastral setae ca. 13–18. Ventral setae simple. **Sternal region** (Fig. 27A) – presternal platelets free, anterior margin of the sternal shield thickened between the concavities for Co I, more or less distinctly split in the axial line into lateral parts (Fig. 27A–C). Genital opening located in the anterior
<figure><img src="image_32.png" /><figcaption>**Figure 25** *Holzmannia (Juvaria) silvestris n. sp.*, female, legs I and II ventrally: A – Co I, Tr I and Fe I; B – Ge I; C – Ti I; D – Co II, Tr II and Fe II; E – Ge II and Ti II; F – Ta II. For abbreviations regarding the setae – see Methods.</figcaption></figure>

part of the sternum contains a genital lamina inserted through the genital opening under the sternal shield (Fig. 27A, B). The anterior part of the genital lamina is sub-rectangular, with more or less concaved anterior margin and rounded anterior corners (Fig. 27A–C). Sternal gland pores *gv1* absent. In between setae *st3* and *st4* two pairs of weakly pronounced elongated thickenings of the sternal cuticle are encountered. Gland pore *gv2* with one opening, pores *iv5* between *st5* and *ZV1* setae, but distinctly closer to *ZV1*. Pores *ivo1*, *ivo2*, *ivo3*, and also *gv3* well discernible.

**Gnathosoma** — Gnathotectum (Fig. 27D) trispinate with the central prong somewhat wider and longer. Corniculi conical, hypostome with ca. 10 rows of denticles. Hypostomatic and palpcoxal setae simple. Palptrochanter *v1* seta simple, *v2* barbed. Chelicera (Fig. 27E) — movable digit with two teeth followed proximally by an arcuate edge, the proximal tooth slightly larger. Fixed digit with two teeth ahead and one tooth behind the pilus dentilis, followed
<figure><img src="image_33.png" /><figcaption>**Figure 26** *Holzmannia (Juvaria) silvestris* n. sp., female, leg IV ventrally: A – Co and Tr; B – Fe, Ge and Ti; C – Ta. For abbreviations regarding the setae – see Methods.</figcaption></figure>

by larger lamellar tooth.

**Legs** — Coxa I, Co III and Tr IV (Fig. 27F–H) as in the female. Leg II (Fig. 28A, B) spurred as follows: when viewed from the ventral side (Fig. 28A), femoral main spur is narrow and slightly curved, axillary process absent, distal additional spur conical, genual and tibial
<figure><img src="image_34.png" /><figcaption>**Figure 27** *Holzmannia (Juvaria) silvestris* n. sp., male: A – sternal shield with anterior thickening, genital opening and genital lamina; B, C – anterior margin of the sternum, two aspects; D – gnathotectum; E – chelicera, antiaxially; F – Co I, ventrally; G – Co III, ventrally; H – Tr IV, ventrally. Abbreviations as in Figure 6.</figcaption></figure>

spurs conical and pointed. Femoral additional spur, as well as the genual and tibial ones are located ventrally along the leg's axis. From the lateral perspective (Fig. 28B), femoral main spur straight, at the base, with convexity of the external margin, additional spur rounded and located close to the distal article margin, genual and tibial spurs hooked and pointed, the genual one located closer to the distal margin of the article than the tibial spur. Leg setation – legs I, III as in the female. Setae on leg II (Fig. 28A, B) simple but genual and tibial anterolateral setae,
<figure><img src="image_35.png" /><figcaption>**Figure 28** *Holzmannia (Juvaria) silvestris n. sp.*, male: A, B – Fe, Ge and Ti of leg II ventrally (A) and anterolaterally (B). Some setae are marked. Arrow on the anterolateral side.</figcaption></figure>

which are barbed. Femoral setae *al1* and *al2* thicker and barbed terminally, *ad1* with one barb, *ad2* and *ad3* thick and short. Leg IV setation as in the female, tarsal antero-dorsal seta (*ad1*) of normal length (75–78), tactile seta (*pd1*) long (168–172), seta *pl4* finely barbed. Other aspects of legs I–IV unremarkable.

**Material examined**

**Holotype** — Female (slide no. 2112 A), Karawanks Mts, Austria, 46.4616°N, 14.6576°E, alt. ca. 860 m a.s.l., 14 Sept. 2006, wet deciduous litter and moss on the rocks.

**Paratypes** — 11 females, 14 males (slides no. 2112 B-E, 2106 A, 2114), ibid.; 2 females, 1 male (slides no. 2094 A, 2099 A), near Freibach Stausee, Karawanks Mts, Austria, 46.5056°N, 14.4470°E, alt. ca. 750 m a.s.l., 14 Sept. 2006, litter in the mixed forest; 2 males (slide no. 2103 A), Karawanks Mts, Austria, 46.4616°N, 14.6576°E, alt. ca. 860 m a.s.l., 14 Sept. 2006, litter in the deciduous forest; 1 female (slide no. 2131 A), Felsentore, Karawanks Mts, Austria, 46.4531°N, 14.6610°E, alt. ca. 1350 m a.s.l., 23 July 2007, litter in the beech forest. Holotype and paratypes leg. W. Witaliński.

**Type deposition** — Types are deposited in the Zoological Division of the Nature Education Centre, Jagiellonian University, Kraków, Poland.
## Etymology

The specific name *silvestris* (lat. *silvestris* means forester) refers to large forests in the Karawanks Mountains where the types were encountered.

## Differential taxonomy

The newly described species *H. (J.) pilosa n. sp.* and *H. (J.) silvestris n. sp.* possess a peritrematal shield joined to the lateral margin of the dorsal shield in the female, so it effectively matches up, along with its other features, the characteristics for *Holzmannia* genus. However, in both species the sternal gland pores *gv1* are absent, as in the case of *Juvaria n. subgen.*, what actually distinguishes the species from *H. (H.) disfistulatus* (Athias-Henriot, 1967). Furthermore, the anterior prong of the epigynium, as well as the metagynial sclerites of the paragynial shields are quite different between *Holzmannia* s.s. and *Juvaria n. subgen.* The *Holzmannia* genus may likely also embrace the species described by Schmölzer as *Ologamasus longisetosus* Schmölzer, 1995, encountered in South Austria. Both sexes were considered in describing the species, although both the actual description, and especially the drawings appear rather confusing. The ventral side of the female shows the large, free presternal plates never encountered in *Ologamasiphis* and *Holzmannia*, whereas in the male, the second leg is also unusual: Fe II has calcar and axillary process instead of an additional distal spur, and on Ge II, there are no spurs whatsoever. Such a combination of characteristics was never observed in the case of both genera under study.

*Holzmannia (J.) pilosa n. sp.* female differs from *H. (J.) silvestris n. sp.* in many respects. Most evident are the following ones: in *H. (J.) pilosa n. sp.* the dorsal setae are very long, whereas in *H. (J.) silvestris n. sp.* the podonotal setae are moderately long, but the opisthonotal ones very short. In the female, metagynial sclerites of the paragynial shields are different, arcuate in the former species, and terminally sigmoidal in the latter. In *H. (J.) pilosa n. sp.* males, both parts of thickened anterior margin of the sternum are axially connected by an arcuate thickening, whereas in *H. (J.) silvestris n. sp.* there is an axial cleft in the thickening of the sternum. The armature of the second leg is similar, yet the tibial spur in *H. (J.) pilosa n. sp.* is relatively short and rounded apically, in *H. (J.) silvestris n. sp.*, although it is much longer, arcuate in the lateral view, and pointed.

## A modification of Hrúzová & Fend'a (2018) key to the *Parasitidae* genera

We propose to have the relevant steps in the key by Hrúzová & Fend'a (2018) to be modified, with a view to having them brought in line with the new taxonomic concept, as addressed further below:

27. Female peritrematal shields fused with holodorsal shield margin for their whole length, opisthogaster free; male without transverse suture on the dorsal shield....................28
— Female peritrematal shield only anteriorly joined with the holodorsal shield margin, opisthogaster either fused with the peritrematal shield or free; male with or without transverse suture on the dorsal shield ....................................................................................32

28. Idiosoma not globular; gnathotectum with three or five prongs; female with two large triangular presternal plates, podonotal region with 18–22 pairs of setae, opisthonotal region hypertrichous, opisthogaster with 11–32 setae pairs, epigynium triangular or subpentagonal; femur II of the male with large apophysis (main spur) and axillary process....................
.... *Pergamasus* Berlese, 1903. Type species: *Acarus crassipes* Linneaus, 1758 sensu Berlese, 1906 – 29

— Idiosoma globular; gnathotectum three- or one-prong type; female with the presternal plates completely fused with the sternal shield, or only small triangular plates remaining, podonotal region with 16–20 pairs of setae, opisthonotal region with 12–23 pairs of setae, opisthogaster with seven or eight pairs of setae, epigynium pentagonal or heptagonal; femur II of the male
with large apophysis (main spur), axillary process setiform or absent..................28a

28a. Podonotal region with 20 pairs of setae, opisthonotum with 21-23 pairs of setae; opisthogaster with seven or eight pairs of setae; epigynium heptagonal; in the male, femur II with setiform axillary process, an additional spur in the distal region of the article absent......
.. *Heteroparasitus* Juvara-Balş, 1975. Type species: *Pergamasus tirolensis* Sellnick, 1968 – 31
— Podonotal region with 16-17 pairs of setae, opisthonotum with 12-20 pairs of setae, opisthogaster with seven pairs of setae; epigynium pentagonal or heptagonal; in the male, on Fe II axillary process absent, an additional spur in the distal region of the article present......
. *Holzmannia* Juvara-Balş, 2002. Type species: *Pergamasus disfistulatus* Athias-Henriot, 1967 – 28b

28b. On ventral idiosoma *gv1* gland pores present...................subgenus *Holzmannia* s. s.
— On ventral idiosoma *gv1* gland pores absent....................................................
...................subgenus *Juvaria* n. subgen. Type species: *Holzmannia* (*Juvaria*) *pilosa* n. sp.

32. Female peritrematal shield posteriorly fused with the opisthogaster; extension of the peritrematal shield behind the stigmata discernible in the male; the male without transverse suture on the dorsal idiosoma .................................................................................. 33
— Female peritrematal shield free posteriorly; extension of peritrematal shield behind the stigmata not recognizable in the male; the male with or without transverse suture on the dorsal idiosoma.................................................................................................................. 40

40. Podonotal region with 22-23 or 31-45 pairs of setae, opistonotal region hypertrichous, with approx. 60 pairs of setae; opisthogaster with 11-30 pairs of ventral setae; paraxial setae on palpgenu fringed; female genital pores *iv5* near the posterior margin of the epigynium; presternal plates of the female triangular, contiguous; movable digit of male chelicera with two teeth............ *Amblygamasus* Berlese, 1906. Type species: *Gamasus dentipes* Koch, 1839
— Podonotal region with 16-21 pairs of setae, opistonotal region oligotrichous or holotrichous; opisthogaster with 8-11 pairs of ventral setae; paraxial setae on the palpgenu entire, spatulate; female genital pores *iv5* on a soft cuticle between the epigynium and the opisthogaster; presternal plates of the female triangular or ribbon-like, or in the shape of small triangular sclerites with intermediate sclerotizations, or fused with the sternal shield; movable digit of male chelicera with one or two teeth .................................................................................. 41

41. Idiosoma usually globular; podonotal region with 18-20 pairs of setae, opisthonotum with 19-22 pairs of setae; opisthogaster with seven pairs of setae; on the ventral idiosoma *gv1* gland pores absent; in the female, presternal plates fused with the sternal shield; in the male, movable digit of chelicera with two teeth, the genital opening is located just behind the anterior sternal margin and contains the genital lamina, instead of separate subgenital sclerite, a cuticular ribbon joining the left and the right sternal parts is present; Fe II bears a main spur and an additional spur in the distal region of the article, but the axillary process is absent....................
. *Ologamasiphis* Athias-Henriot, 1971. Type species: *Pergamasus epigynialis* Willmann, 1940
— Idiosoma oval; podonotal region with 20-21 pairs of setae, opisthonotal region with 23-24 pairs of setae, opisthogaster with 8-11 pairs of setae; on the ventral idiosoma *gv1* gland pores present; in the female, presternal plates free, triangular, ribbon-like, or in the shape of small sclerites with intermediate sclerotizations; in the male, movable digit of chelicera with one or two teeth, the genital opening with subgenital sclerites is located in front of sternal anterior margin; Fe II bears a main spur and axillary process.................... *Paragamasus* s. l. – 43

42. deleted
## Acknowledgements

The Authors appreciate the assistance extended to them by Dr M. Pabijan, Jagiellonian University, Kraków, Poland, who kindly made available to them the material originally collected both in Greece (Peloponnesus), and Romania, as well as express their gratitude to the two anonymous reviewers for their incisive and perceptive comments, which substantially contributed to enhancing overall quality of the manuscript. The study was partly supported by the grant allocated by the Jagiellonian University, Kraków, Poland (Grant Ref. No N18/DBS/000005).

## ORCID

Wojciech Witaliński https://orcid.org/0000-0001-6396-0728

Dariusz J. Gwiazdowicz https://orcid.org/0000-0002-0064-2316

## References

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