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

Witaliński, Wojciech ¹ and Gwiazdowicz, Dariusz J. ²

¹✉ Department of Comparative Anatomy, Institute of Zoology and Biomedical Research, Jagiellonian University, Gronostajowa 9, 30-387 Kraków, Poland.
²Faculty of Forestry and Wood Technology, Poznań University of Life Sciences, Wojska Polskiego 71c, 60-625 Poznań, Poland.

2023 - Volume: 63 Issue: 3 pages: 867-905

https://doi.org/10.24349/0v36-z6a2
ZooBank LSID: AA59D10C-2546-4012-B221-94C29784F723

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Original research

Keywords

mite taxonomy new species new subgenus rare mite fauna Romania, Greece Austria Karawanks Mts

Abstract

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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 & Fenďa, 2018; Juvara-Balş, 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-Balş, 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-Balş (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 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 & Fenďa, 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 & Fenďa, 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, 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 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 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.

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 al1 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 al1 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), 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 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: 38A4A1E0-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 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 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 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 (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, 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 & Fenďa (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 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, 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. 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 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 (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.

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, 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 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 coxae 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 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 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 (JV1), 4–5 (ZV1), 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 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 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 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, which are barbed. Femoral setae al1 and al2 thicker and al2 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 & Fenďa (2018) key to the Parasitidae genera

We propose to have the relevant steps in the key by Hrúzová & Fenďa (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, opisthonotal 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, opisthonotal 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).

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