Zootaxa · September 2006 DOI: 10.5281/Zenodo.174005

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New Mites Of The Genus Holoparasitus Oudemans, 1936 From Northern Austria And Karavanke (Acari: Parasitidae)

Article in Zootaxa · September 2006 DOI: 10.5281/zenodo.174005

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The user has requested enhancement of the downloaded file. Zootaxa 1320: 15–27 (2006) ISSN 1175-5326 (print edition) www.mapress.com/zootaxa/ ZOOTAXA 1320 Copyright © 2006 Magnolia Press ISSN 1175-5334 (online edition)

New mites of the genus Holoparasitus Oudemans, 1936 from northern Austria and Karavanke (Acari: Parasitidae)

WOJCIECH WITALIÑSKI Institute of Zoology, Jagiellonian University, R. Ingardena 6, PL-30 060 Kraków, Poland. E-mail: [email protected]

Abstract

A new mite species, Holoparasitus rhombogynialis sp. nov. (Acari: Parasitidae) is described from Karavanke (Slovenia-Austria border). H. hemisphaericus (Vitzthum, 1923) is redescribed, including a description of the previously unknown male from the type locality in northern Austria. Both species are considered to be members of the Holoparasitus hemisphaericus species-group, which is newly defined. Ologamasus absoloni Willmann, 1940 is synonymised with H. hemisphaericus, and the female of H. intermedius (Holzmann) sensu Micherdziński is synonymised with H. cornutus Juvara-Bals & Witaliński.

Key words: Acari, Parasitidae, Holoparasitus, new species, new synonyms, redescription, mite taxonomy, Karavanke

Introduction

A well known problem with early taxonomic descriptions is that many "known" species are difficult or even impossible to identify without close examination of the type material. Sometimes, as in the case of Holoparasitus hemisphaericus (Vitzthum, 1923), the description and drawings are misleading. According to the original description based on a single female (Vitzthum, 1923), the key character of the species is a prominent spur on the distal margin of trochanter IV. As mentioned in an earlier paper (Witaliński & Skorupski, 2002), we examined this type specimen and concluded that H. micherdzinskii Witaliński, 1981 is a junior synonym of H. hemisphaericus (Vitzthum). Further, it was found that the peculiar spur on Tr IV was an artefactual interpretation of the normal cuticle of the segment when observed from the side. Therefore, attempts at finding this character and identifying H. hemisphaericus have been unsuccessful. Fortunately, new material collected from the type locality of the species (Waidhofen a.d. Thaya, Austria) made it possible to examine and describe both sexes. It has also become clear that a new,

Accepted by B. Halliday: 21 Aug. 2006; published: 25 Sept. 2006 15 ZOOTAXA undescribed species from the Karavanke mountains, morphologically very similar to H. 1320 hemisphaericus, justifies the formation of a new Holoparasitus hemisphaericus species- group, containing H. hemisphaericus and H. rhombogynialis sp. nov.

Holoparasitus hemisphaericus species-group

Only the specific characters of the new species group are given. For other morphological features of the genus Holoparasitus and its different species groups (annulus, caesus, calcaratus, mallorcae, peraltus) see Juvara-Bals (1975), Hyatt (1987), Juvara-Bals & Witaliński (2000), Witaliński & Skorupski (2002, 2003), Juvara-Bals & Witaliński (2006).

Diagnosis of the species group In both sexes gland pores gv1 are present and pores gv2 are located in unmodified cuticle; gnathotectum is of the trispinate-type. Female. Presternal plate wide, narrowed medially, its anterior margin smooth, lateral platelets free and narrow; sternal shield in one piece; posterior paragynial lobes distant; metagynial sclerites semicircular; posterolateral protrusions “locking” epigynial plate minute or absent; thickening of the underside of anterior paragynial edge facing coxa III absent; central apex of epigynial shield less pigmented and right-angled, lateral prongs large and acute; epigynial subapical structure well pigmented, broad and barrel-shaped in outline; lateral protrusions bipartite, comprised of small, semicircular and well sclerotised posterior structures passing anteriad into less sclerotised hyaline projections; endogynium cup-shaped, with spines gathered in two lateral (left and right) groups, posterior endogynial margin forms two broad, partially overlapping lamellae, elliptical or leaf- shaped, partly covering the endogynial opening; distinct broad endogynial lamella covers posterior part of endogynium ventrally. Male. Sternum with large excipulum open on both ends and formed by three parallel, longitudinally oriented thickenings, accompanied by two additional anterolateral ones (left and right); central part of hypostome regularly triangular and moderately sclerotised; corniculi slender or extended in proximal half; fixed cheliceral digit straight and slender with many tiny denticles followed by lamellar edge, movable digit with large tooth proximally and 4–5 smaller teeth distally; hypostomatic setae on a separate piece of cuticle.

Remarks The proposed species group is most similar to the peraltus species-group (Witaliński & Skorupski, 2003). The best diagnostic characters differentiating both groups are as follows: in females, the epigynial subapical structure possesses an inverted T-shaped thickening (peraltus group) or this structure is absent (hemisphaericus group); in males, the excipulum is hoof-shaped and corniculi are not on a separate piece of cuticle (peraltus

16 © 2006 Magnolia Press WITALIŃSKI group), or the excipulum is “open” at both anterior and posterior ends and flanked by two ZOOTAXA very prominent lines of sternum reticulation running parallel to the anterolateral margins 1320 of the excipulum, and hypostomatic setae are on a separate piece of cuticle (hemisphaericus group). In the following diagnoses specific features complementary to species-group characters are included.

Holoparasitus hemisphaericus (Vitzthum, 1923) (Figs. 1–21)

Ologamasus hemisphaericus Vitzthum, 1923: 101. Holoparasitus absoloni (Willmann, 1940) female, (as Ologamasus absoloni Willmann, 1940: 212), syn. nov. Holoparasitus micherdzinskii Witaliński, 1981: 41, synonymy by Witaliński & Skorupski, 2002: 55.

Material examined Material deposited in the Zoologische Staatssammlung, München: holotype female, slide "V15", left-hand labelled "Ologamasus hemisphaericus Vitz. 1923 & Hypost. Palpi Chel. B. 1 TYPUS" and right-hand labelled "Waidhofen/Thaya, N.-Öst. Muttergang v. Hylesinus fraxini Panz. leg: Wichmann det: Vitzthum", and slide "V22", left-hand labelled "Ologamasus hemisphaericus Vitzth. & TYPUS" and right-hand labelled "Waidhofen, Thaja leg: Wichmann det: Vitzth. 25. 6. 1920", as well as female, slide originally labelled "13", right-hand "Olog. absoloni & Det. C. Willmann" and left-hand "Abs. 775", and additionally labelled as follows "Ologamasus absoloni WILLMANN, 1940 = Holoparasitus absoloni (WILLMANN, 1940)" " rev.: I. Juvara Bals, 1999" "1 f; Holotypus". Material collected by the author and deposited in the Zoological Museum, Jagiellonian University, Cracow: 22 females, 12 males (slides no. 1881–1890), 12.11.2004; Waidhofen a.d. Thaya, Austria, thick layer of moss under group of larches in spruce forest; 48 o 49.18' N, 015o 21.50' E, alt. ca. 650 m a.s.l.

Diagnosis In the female, the endogynium is circular or oval in outline, with two lateral protrusions each bearing several spines directed adaxially and more or less anteriad; two lamellae protruding from the posterior margin of endogynium elliptical, covering most of the endogynial entrance; anterior margin of endogynial lamella convex; epigynial central prong right-angled and evenly but weakly pigmented; subapical epigynial structure with bipartite lateral protrusions, these located posteriorly, semicircular and not extending beyond epigynial margin, anterior protrusions hyaline, extending out of the epigynial margin. In the male, sternal shield with prominent excipulum composed of three parallel thickenings with two depressions in-between and two additional anterolateral thickenings

HOLOPARASITUS © 2006 Magnolia Press 17 ZOOTAXA of sternal reticulation frequently forming an arc delimiting the excipulum anteriorly; 1320 cuticle of excipulum with well visible large pores forming a polygonal network; corniculi slender; spurs on Ge II and Ti II with concave margin, similar in shape, size and location.

Description Female Idiosoma well sclerotised, brownish, 520–570 x 685–755 µm (n=10). Length of podonotal setae: 33–37 µm (j1), 28–32 µm (r3), others 10–37 µm; opisthonotal setae very short, 6–8 µm; length of peritrematal groove 215–220 µm. Venter. Presternal plate wide, narrower medially, lateral platelets wedge-shaped and free (Fig. 1). Sternal reticulation (Figs 1, 2) more pronounced in an area demarcated posteriorly by a line running through the second pair of pores (iv2). Gland pores gv1 present adaxially to setae st3. Setae of sternogenital region finely barbed, 50 µm (st1) to 65 µm (st2, st3) long. Genital region. Paragynial shield (Fig. 2) with posterior paragynial lobes relatively small and far from one another; metagynial sclerite semicircular; posterolateral protrusion “locking” epigynial plate minute, thickening of the underside of anterior paragynial edge facing coxa III absent. Central apex of epigynial shield (Figs. 3, 4) less pigmented and close to right-angled in shape, lateral prongs large and acute. Subapical structure well pigmented, barrel-shaped in outline. Lateral protrusions bipartite; posterior projections are semicircular and well sclerotised, ending more or less at the margin of the epigynium and passing anteriad into less sclerotised hyaline protrusions extending out of the epigynial margin. Endogynium (Figs 2, 5, 7) circular or oval in outline, with two lateral protrusions each bearing several spines directed adaxially and more or less anteriad. Rarely are spines located directly on the endogynium wall. Posterior margin of endogynium protruded into two leaf-shaped lamellae, partly overlapping and covering most of the endogynial opening. Hyaline endogynial lamella (Figs 6, 8) with convex anterior edge extending between metagynial thickenings. Gland pores gv2 in normally sclerotised cuticle. Opisthogaster bears eight pairs of ventral setae 22–39 µm long. Gnathosoma. Gnathotectum trispinate, hypognathal groove with six clearly visible anterior rows of denticles and four posterior rows represented only by 1–2 lateral denticles. Palpcoxal setae finely barbed, hypostomatic setae simple. Distal half of the fixed digit of the chelicera with two teeth and a low tubercle in-between, in front of the pilus dentilis, followed by two larger teeth located on the internal (adaxial) side. There is a lamellar edge on the external (abaxial) side. Movable digit with three teeth. Pedipalp trochanter with seta v1 simple and v2 barbed distally. Legs. Leg structure and setation unremarkable.

Male Idiosoma brownish, well sclerotised, 455–465 x 615–665 µm (n=5). Length of podonotal setae: 28–29 µm (j1), 26–32 µm (r3), others 11–29 µm; opisthonotal setae very

18 © 2006 Magnolia Press WITALIŃSKI short, 7–12 µm; length of peritrematal groove 205–215 µm. ZOOTAXA Ventral side (Figs 9–11). Presternal plates small, triangular, with less sclerotised 1320 projections anteriad. Genital lamina prominent, located in a deep concavity in the anterior sternal margin; its anterior hyaline edge bilobate, lateral pointed projections relatively short and pigmented (Fig. 11). Sternogenital shield with prominent excipulum (Figs 9, 10) composed of three parallel sclerotised ridges; its cuticle shows large pores pentagonally or hexagonally arrayed. The excipulum is demarcated anterolaterally by thickened lines of sternal reticulation. Gland pores gv1 at st3 level, with terminally dilated channels directed laterally. Two pairs of thickenings behind excipulum are more or less discernible. Gland pores gv2 in unmodified cuticle. Sternal setae length from 46 µm (st1) to 35 µm (st3), opisthogastric setae 22–30 µm.

FIGURES 1–8. Holoparasitus hemisphaericus (Vitzthum), female: 1 presternal plate and sternal shield; 2 genital region with paragynia, epigynium and endogynium; 3, 4 epigynium; 5, 7 endogynium; 6, 8 endogynial lamella. 1: slide no. 1881; 2: 1882; 3: 12A; 4, 5, 6: 11A; 7, 8: 1886. Scale bars: 0.1 mm.

FIGURES 9–17. Holoparasitus hemisphaericus (Vitzthum), male: 9 presternal plates and sternum; 10 excipulum; 11 genital lamina; 12–14 gnathotectum, dissected and pressed (12) or in situ (13, 14); 15 gnathosoma, ventrally; 16, 17 left and right chelicera. 9–11, 13: slide no. 1889; 12, 16, 17: 1885; 14, 15: 1890. Scale bars: 0.1 mm.

Gnathosoma (Figs 12–15). Gnathotectum (Figs 12–14) trispinate, but lateral prongs tiny. Hypognathal groove with 10 or 11 rows of clearly visible denticles (Fig. 15); of these, the most proximal row is frequently represented only by lateral denticles. Palpcoxal setae finely barbed, hypostomatic setae simple. Incisions present in cuticle behind hypostomatic setae. Corniculi slender. Chelicera (Figs. 16, 17) with fixed digit narrow, concave in distal third, bearing a row of several tiny denticles; abaxially there is a semicircular incised

20 © 2006 Magnolia Press WITALIŃSKI lamella. Behind pilus dentilis, the ventral border of the digit forms a straight lamellar edge. ZOOTAXA Movable digit with 4–5 teeth in distal third followed proximally by a larger tooth. 1320 Arthrodial membrane with a well-pronounced brush-like process; synarthrodial membrane wavy. Pedipalp trochanter with seta v1 simple, v2 distally barbed.

FIGURES 18–21. Holoparasitus hemisphaericus (Vitzthum), male: 18 femur, genu and tibia of leg II (most setae not shown); 19, 20 rows of denticles on anterolateral side of both coxae II; 21 spurs on Fe II, Ge II and Ti II, ventrally (arrow at anterolateral side). 18–20: slide no. 1884; 21: 1889. Scale bar: 0.1 mm.

Legs. Leg I, III, and IV unremarkable. Leg II (Figs 18–21) with spurs as follows: the main spur on femur short, conical, with tubercle located ventrally at its base, and ending in a very small apical segment (Fig. 18). Axillary process with apical part oval, larger than the apical segment of the main spur. Spurs on Ge II and Ti II with concave margin and similar in shape, size and location. Ventral seta on Ti II larger than that on Ge II. Coxa II bears anterolaterally a row of denticles, 7–9 plus 1; the row ends with a long tooth-like structure directed ventrally (Figs 19, 20).

Remarks H. absoloni (Willmann, 1940) is a junior synonym of H. hemisphaericus (Vitzthum, 1923). A single female mounted on a slide deposited in Zoologische Staatssammlung,

HOLOPARASITUS © 2006 Magnolia Press 21 ZOOTAXA München, was designated as holotype by I. Juvara-Bals in 1999. This specimen is in a 1320 moderate condition, but available features (subepigynial structure, endogynium, sternum) strongly support synonymy with H. hemisphaericus (both types, kindly loaned to the author by Dr. L. Tiefenbacher and Dr. Eva Karl, were examined). The males determined by Willmann as H. absoloni belong in fact to Heteroparasitus tirolensis (Sellnick, 1968) (Karg 1971: 362). The only female determined and described by Micherdziński (1969: 363) as a female of H. intermedius (Holzmann, 1969) was synonymised by Karg (1971: 362) with H. absoloni. However, the female of H. intermedius (Holzmann) sensu Micherdziński actually belongs to H. cornutus Juvara-Bals & Witaliński, 2000 (syn. nov., Micherdziński’s specimen examined).

Holoparasitus rhombogynialis sp. nov. (Figs. 22–41)

Material examined Holotype female (slide no. 1358), 31.07.2000, ca. 2 km north of Loibl Pass, Karavanke mountains, Austria, 46o 26.70' N, 014o 15.80' E, alt. ca. 1000 m a.s.l., litter in beech forest with spruce admixture; 3 females, 2 males paratypes (slides no. 1357, 1359, 1360), ibid.; 1 female, 1 male paratypes (slide no. 1351), 31.07.2000, ca. 4 km south of Loibl Pass, Karavanke, Slovenia, 46o 24.65' N, 014o 16.50' E, alt. ca. 900 m a.s.l., litter in mixed forest (beech, ash, fir, spruce, larch). Types were collected by the author and Dr. M. Skorupski, Agriculture University, Poznań, Poland, and are deposited in the Zoological Museum of the Jagiellonian University, Cracow, Poland.

Diagnosis In both sexes the gnathotectum is trispinate. In the female, the endogynium is rhombic in ventral perspective, with two lateral groups of spines, the majority of which are directed backwards; two lamellae protruding from the posterior endogynial margin leaf-shaped, i.e. narrowing apically; anterior margin of endogynial lamella undulate and concave medially; epigynial central prong weakly pigmented (more so terminally), its margins shallowly concave; anterior part of lateral protrusions of subapical epigynial structure hyaline and membranous. In the male, the excipulum has a ridge-like central thickening, the lateral thickenings usually form cuticular lids covering margins of excipular depressions; anterolateral thickenings of sternal reticulation frequently form lids similar to those of excipulum; setae st1 are located on thickenings; two pairs of distinct button-like thickenings present at the level of sternal pores iv3 and setae st3; corniculi extended proximally; spurs on Ge II and Ti II differ in shape and orientation, that on Ti II arcuate in ventral perspective and located at the margin of less sclerotised cuticle.

22 © 2006 Magnolia Press WITALIŃSKI Description ZOOTAXA Female 1320 Idiosoma brownish, 490–510 x 635–725 µm (n=4). Length of podonotal setae: 29–33 µm (j1), 30–33 µm (r3), others 11–34 µm; opisthonotal setae very short, 8–13 µm; length of peritrematal groove 224–228 µm. Venter. Presternal plate and lateral platelets as in H. hemisphaericus (Figs 22, 23). Sternum relatively longer than in H. hemisphaericus: its width (measured between second leg openings) to length (measured axially) ratio does not exceed 1.9, whereas in H. hemisphaericus it is greater than 2.2. Left and right parts of sternal reticulation line running through the second pair of pores (iv2) usually arcuate (Fig. 22). Gland pores gv1 present. Setae of sternogenital region: 60–61 µm (st1), 64–67 µm (st2), 64–65 µm (st3), 50–52 µm (st4). Genital region. Paragynial shields as in H. hemisphaericus. Central prong of epigynial shield (Fig. 24) weakly pigmented, but more so terminally; its margins shallowly concave. Subapical structure with arcuate lateral thickenings. Lateral protrusions semicircular and well sclerotised, ending at epigynium margin and passing anteriad into membranous protrusions. Endogynium (Figs 25, 27) rhombic in ventral perspective, with two lateral groups of spines directed mostly backwards. From the posterior endogynial margin, two leaf-shaped lamellae protrude anteriad and somewhat ventrally, thus visibly shorter in ventral perspective (Fig. 27). Posterior part of endogynium is covered ventrally by endogynial lamella with anterior margin undulate and concave medially (Fig. 26). Gland pores gv2 in normal cuticle. Opisthogaster with eight pairs of ventral setae 29–46 µm long. Gnathosoma. Gnathotectum trispinate (Fig. 28), corniculi conical. Hypognathal groove with ca. 10 very weakly pronounced rows of denticles. Palpcoxal setae barbed, hypostomatic setae simple. Chelicera (Fig. 29). Fixed digit with two teeth in front and two behind the pilus dentilis, followed by two parallel lamellar edges, similar to those in H. hemisphaericus. Movable digit with three teeth. Pedipalp trochanter with seta v1 simple and v2 barbed distally. Legs. Leg structure and setation unremarkable, except for Co I bearing ventrally two clearly visible tubercles (Fig. 30).

Male Idiosoma brownish, 385–420 x 595–600 µm (n=2). Length of podonotal setae: 26–29 µm (j1), 27–29 µm (r3), others 8–22 µm; opisthonotal setae very short, 8–13 µm; length of peritrematal groove 205–209 µm. Venter (Fig. 32). Presternal plates small, triangular. Genital lamina (Fig. 33) weakly sclerotised, located in a deep concavity of anterior sternal margin, flanked by tubercular protrusions; its anterior hyaline edge circular, lateral acute projections not discernible, and microsclerite under lamina well sclerotised. Excipulum on sternogenital shield prominent (Fig. 32), composed of three largely parallel sclerotised ridges, of these the lateral ones are

HOLOPARASITUS © 2006 Magnolia Press 23 ZOOTAXA slightly arcuate and form lamellar protrusions (lids) covering the excipulum depressions 1320 laterally. The cuticle of excipulum shows distinct pores, but their pattern is similar to that in other areas of the sternum. Anterolaterally, the excipulum is flanked by thickened lines of sternal reticulation, also forming lamellar cuticular lids. On the posterior ends of these thickenings gland pores gv1 are present, whereas more prominent reticulation runs anteriad, ending in thickenings located close to the bases of setae st1. Two pairs of button- like thickenings behind excipulum are present. Gland pores gv2 in normal cuticle. Approximate length of sternal setae: 44–46 µm (st1), 46 µm (st2), 33–39 µm (st3), 35–37 µm (st4), the length of opisthogastric setae is 20–33 µm.

FIGURES 22–29. Holoparasitus rhombogynialis sp. nov., female: 22, 23 presternal plate and sternal shield; 24 epigynium; 25, 27 endogynium; 26 endogynial lamella; 28 gnathotectum; 29 chelicera. 22, 27: slide no. 1357; 23: 1351; 24–26, 28, 29: holotype, 1358. Scale bars: 0.1 mm.

FIGURES 30–36. Holoparasitus rhombogynialis sp. nov.: 30, 31 ventral aspect of coxa I and trochanter I in female (30) and male (31) (arrow at anterolateral side). Male: 32 presternal plates and sternum with excipulum; 33 genital lamina; 34, 35 gnathotectum; 36 corniculus, ventrally. 30: slide no. 1357; 31–34, 36: 1351; 35: 1360. Scale bars: 0.1 mm.

Gnathosoma. Gnathotectum (Figs 34, 35) trispinate as in female. Hypognathal groove with 10–11 rows of clearly visible denticles; palpcoxal setae barbed, hypostomatic setae simple and inserted on piece of cuticle separated by incisions. Corniculi extended in proximal part (Fig. 36). Chelicera (Figs. 37, 38) with fixed digit straight and narrow, in distal third with thin dentate lamella located on adaxial side. Movable digit with four to six teeth followed by larger tooth proximally. Arthrodial membrane with brush-like process, synarthrodial membrane wavy. Pedipalp trochanter with seta v1 simple and shorter than v2, which is barbed. Femur with broad elevation located ventrally at level of anterolateral seta. Legs. Coxa I with two tubercles located ventrally (Fig. 31). Leg II with spurs as follows (Figs 39–41): the main spur on femur short and conical, ending in a low, roundish apical part; ventral seta inserted at base of spur on weakly pronounced elevation. Axillary process slightly curved, larger than apical segment of main spur. Spurs on genu and tibia differ in shape and orientation: the spur on genu straight with slightly concave edge, oriented obliquely in ventral perspective. Spur on tibia arcuate, located at the border of less sclerotised cuticle and oriented axially in ventral perspective (Fig. 40). Ventral seta on Ti II larger than that on Ge II. Coxa II bears anterolaterally 8 + 1 denticles in fan arrangement (Fig. 39). Legs III, and IV unremarkable.

FIGURES 37–41. Holoparasitus rhombogynialis sp. nov., male: 37 chelicera; 38 ibid., crushed; 39 coxa, trochanter, femur, genu and tibia of leg II (only the position of most setae is marked); 40 femur, genu and tibia of leg II in ventral perspective (arrow at anterolateral side); 41 Fe II, ventrally, another aspect. 37–39: slide no. 1360; 40: 1357; 41: 1351. Scale bars: 0.1 mm.

Acknowledgements

The author expresses gratitude to Dr. M. Skorupski, Department of Forest and Environmental Protection, Agriculture University, Poznań, for help during collection of the material in the Karavanke mountains, and M. Pabijan, Institute of Zoology, Jagiellonian University, Cracow, for improving the English. This work was supported by grant no. BW/IZ/28/2005 from the Jagiellonian University.

26 © 2006 Magnolia Press WITALIŃSKI References ZOOTAXA 1320 Holzmann, C. (1969) Die Familie Parasitidae Oudemans, 1901. Acarologie. Schriftenreiche für Ver- gleichende Milbenkunde, 13, 3–55 + 23 Plates. Hyatt, K.H. (1987) Mites of the genus Holoparasitus Oudemans, 1936 (Mesostigmata: Parasitidae) in the British Isles. Bulletin of the British Museum (Natural History), Zoology, 52, 139–164. Juvara-Bals, I. (1975) Sur le genre Holoparasitus Oudemans, 1936 et sur certains caractères mor- phologiques de la famille Parasitidae Oudemans (Parasitiformes). Acarologia, 17, 384–409. Juvara-Bals, I. & Witaliński, W. (2000) Description of five new species of Holoparasitus s. str. with redescription of H. apenninorum (Berlese, 1906) and H. cultriger (Berlese, 1906) from Italy and Spain (Acari, Gamasida, Parasitidae). Revue Suisse de Zoologie, 107, 3–30. Juvara-Bals, I. & Witaliński, W. (2006) Two new species of the genus Holoparasitus Oudemans from the Mediterranean Basin – Algeria and Sardinia (Acari: Gamasida: Parasitidae). Genus, 17, 437–448. Karg, W. (1971) Acari (Acarina), Milben Unterordnung Anactinochaeta (Parasitiformes). Die freilebenden Gamasina (Gamasides), Raubmilben. Die Tierwelt Deutschlands, 59, 1–475. Micherdziński, W. (1969) Die Familie Parasitidae Oudemans, 1901 (Acarina, Mesostigmata). Państwowe Wydawnictwo Naukowe, Kraków, 690 pp. Sellnick, M. (1968) Zwei neue Pergamasus Arten aus Österreich. Berichte des Naturwissen- schaftlich-Medizinischen Vereins in Innsbruck, 56, 463–472. Vitzthum, H. v. (1923) Acarologische Beobachtungen. 7. Reihe. Archiv für Naturgeschichte, Berlin, 89A, 97–181. Willmann, C. (1940) Neue Milben aus Höhlen der Balkanhalbinsel, gesammelt von Prof. Dr. K. Absolon, Brünn (2. Mitteilung). Zoologischer Anzeiger, 130, 209–218. Witaliński, W. (1981) Holoparasitus micherdzinskii sp. nov. – a new species of mite of the family Parasitidae (Acari, Mesostigmata). Bulletin de l'Académie Polonaise des Sciences, Série des Sciences Biologiques, 29, 41–45. Witaliński, W. & Skorupski, M. (2002) Genus Holoparasitus Oudemans, 1936 in Berlese Acaroteca (Acari: Gamasida: Parasitidae). Part I. Redia, 85, 37–60. Witaliński, W. & Skorupski, M. (2003) Genus Holoparasitus Oudemans, 1936 in Berlese Acaroteca (Acari: Gamasida: Parasitidae). Part II. Redia, 86, 17–22.

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