New Leptogamasus mite species (Parasitiformes: Parasitidae) from Europe. IV. Austria

Three new species of Parasitidae are described from Austria, namely Leptogamasus (Leptogamasus) chelatus n. sp., Leptogamasus (L.) coronarius n. sp. and Leptogamasus (L.) trispinus n. sp.


Introduction
Mites in the genus Leptogamasus Trägårdh, 1936, and subgenus Leptogamasus s. str., are commonly encountered in forest litter across Europe. Their general features have been outlined in previous papers (Witaliński, 2019; Witaliński, 2020. Nine new species originating from northern and central Italy (Witaliński, 2021a,b) were recently described. In the present study, three Leptogamasus (Leptogamasus) species from Lower Austria and Styria are described.

Materials and Methods
The research methods applied in the present study were described in detail in the previous paper in this series (Witaliński, 2020). Since the structure of the podonotum in these species is very uniform, it is not described in depth for every species. The material under study was originally collected by the author.

Diagnosis
Female and male -Gnathotectum trispinate with pointed prongs, the central one somewhat longer, the lateral ones slightly divergent; gland pore gv1 present; podonotum with 21 pairs of setae, opisthonotum with 24 pairs of setae plus either one supplementary pair or to seven supplementary setae; Tr IV without tubercle.
Female -All three prongs of gnathotectum similarly narrow; presternal plates subtriangular, with an irregular posterior margin; anterior margin of the sternal shield slightly concave; gland pores gv1 located far from one to another, occasionally in an asymmetric arrangement; epigynial shield without teeth on the internal (dorsal) surface, anterior margins nearly straight, a lenticular unpigmented area located subapically; spherules of the endogynium subspherical, well discernible anterior walls of the endogynial sac featuring several small teeth directed axially, stipule usually made up of two parts, the anterior one minute, the posterior one larger and tooth-like, usually curved.
Male -Gnathotectum prongs wider and shorter than in the female; genital lamina with rounded anterior corners and lateral margins running backward convergently; presternal plates subtriangular with small anterior protrusion and posterior margin more or less regularly arcuate; corniculi slim but the adaxial margin partly more sclerotised and sinuous when flattened; cheliceral fixed digit wavy in ventral perspective, in the lateral one narrow and sigmoidal with obliquely cut apex, either edentate or with small tooth between apex and pilus dentilis; laterally, leg II femoral main spur straight and axillary process half-moon shaped, genual and tibial spurs conical, in ventral perspective, the tip of genual spur located closer to the article distal margin than the tibial one.
Gnathosoma -Gnathotectum ( Fig. 3E) trispinate, with pointed, similarly narrow and moderately long prongs. Corniculi conical, hypostome with 11 rows of denticles, hypostomal and palpcoxal setae simple. Palptrochanter v1 seta simple, v2 barbed. Chelicera (Fig. 3F)movable digit with four teeth, the proximal one larger. Fixed digit with two distant teeth in front of pilus dentilis, and two behind it, followed by two lamellar and one solid edge projections.
Legs -Setae al1, al2 on Fe II short and thick, whereas the anteroventral seta enlarged and barbed. Ti II with thickened and finely barbed ventral setae. Leg IV: posterodorsal and posterolateral setae on the femur thick and short, antero-and posteroventral setae on the tibia thickened, the latter finely barbed. Some ventral and posterolateral setae on the tarsus, as well as as the posterolateral seta on the basitarsus thickened and terminally barbed. Tr IV without the dorsal tubercle. Other aspects of legs I-IV unremarkable.
Pores gv2 with a double, hardly discernible opening, pores iv5 equally distant from setae st5 and ZV1, or slightly shifted toward st5 setae. Sternum and opisthogaster reticulation scale-like.
Legs -Ventral seta v1 of Tr I simple, seta v2 barbed. Leg II is spurred as follows: when observed from the ventral side ( Fig. 5A-C), femoral main spur is finger-shaped and slightly curved posterolaterally, axillary process hooked and pointed posterolaterally, genual and tibial spurs are conical and rounded apically. The genual spur is oriented along the leg axis, whereas the tibial spur is slightly obliquely oriented. Moreover, the latter is more distant from the distal article margin than the genual one. When viewed from the lateral side, leg II ( Fig. 5D) shows straight femoral main spur, axillary process half-moon shaped, and spurs on the genu and tibia similarly conical, with somewhat rounded tips. Genual spur apex terminating at the article margin level, whereas the tibial one ends at some distance from the margin level. Most of the setae on leg II are simple, but setae al1 and al2 on tibia and al1 on genu can be finely barbed. Setae al1, al2 and ad2 on the femur shorter and thicker, whereas ad3needle-like. Anterolateral setae on basitarsus and tarsus thickened and barbed. Leg IV setation as in the female. Tubercle on Tr IV absent. Other aspects of legs I-IV unremarkable.
Type deposition -The types are deposited in the Zoological Division of the Nature Education Centre, Jagiellonian University, Kraków, Poland.

Etymology
The specific name chelatus refers to the peculiar chelicerae with fixed digits, wavy in a ventral view.

Diagnosis
Female and male -Gnathotectum trispinate with pointed prongs, the central one somewhat longer; gland pore gv1 present; podonotum with 21 pairs of setae, opisthonotum with 24 pairs of setae plus two pairs of supplementary setae, in holotype female one seta S5 is missing; Tr IV without tubercle.
Female -Gnathotectum prongs narrow; presternal plates subtriangular, with convex posterior margin; anterior margin of the sternal shield slightly concave; gland pores gv1 located axially not far from one another; epigynial shield bears two small teeth on the internal (dorsal) surface, anterior margins almost straight; spherules elongated, V-like oriented, stipule crown-shaped, with many teeth.
Male -Gnathotectum prongs short, triangular; genital lamina with the anterior margin concave, and the lateral ones convex, the anterior corners rounded; presternal plates subrect- angular, anterior protrusions large and divergent; corniculi slim, the adaxial margin slightly sinuous; cheliceral fixed digits straight in ventral perspective, with subapical small protrusion antiaxially, in the lateral view narrow and straight, with a small elevation between apex and pilus dentilis, and a row of ca. eight minute denticles behind pilus dentilis; leg II ventrally: femoral main spur short and rounded apically, genual spur located closer to the distal article margin than the tibial spur, in lateral view, the main femoral spur rather short and slightly tapered, the axillary process half-moon to finger-shaped, the genual and tibial spurs with slightly concave margin and rounded distal apex.
Legs -Seta v1 on Co I simple, v2 finely barbed, setae ad on Tr I and ad2 on Fe I short and somewhat thickened. Fe II anteroventral seta larger and finely barbed, setae al1 and al2 short and thick. Seta al on Ti II finely barbed. Leg IV: posterodorsal and posterolateral setae on the femur shorter and thicker, posteroventral seta on the genu finely barbed, posteroventral seta on the tibia enlarged and barbed. Some setae on the tarsi barbed. Tr IV without tooth or tubercle. Other aspects of legs I-IV unremarkable.
Male ( (Fig. 9A) -the anterior margin of genital lamina concave, the lateral margins convex and the anterior corners rounded. Presternal plates subrectangular, anterior protrusions large and divergent. Gland pores gv1 slightly anteriorly to the st3 setae level, followed by two pairs of thickenings of the sternal cuticle. The anterior thickenings are quite fine, the posterior ones rounded and distinct. Pores gv2 with two openings, pore iv5 halfway between st5 and ZV1 setae, ivo2, ivo3, and gv3 distinct.
Gnathosoma -Gnathotectum ( Fig. 9B) with similar triangular prongs. Corniculi (Fig.  9C,D) narrow, the adaxial side slightly sigmoidal, when pressed with adaxial low elevation (Fig. 9D). Hypostome (Fig. 9C) with 11 rows of denticles and thickenings between corniculi at the base of the internal malae. Hypostomal and palpcoxal setae simple. Palptrochanter anterolateral side with a fine thickening ending between setae v1 and v2, v1 seta simple, v2 thicker and barbed. Chelicera (Fig. 9E,F) -ventrally, fixed digits straight, with subapically located small elevation directed antiaxially, the movable digits curved adaxially. Laterally, the movable digit with one tooth followed by an arcuate edge, the fixed digit narrow and straight, with small lamellar elevation between apex and pilus dentilis and a row of ca. eight minute denticles behind pilus dentilis.
Legs -Leg II (Fig. 10) spurred as follows: when viewed from the ventral side (Fig.  10A), the femoral main spur short and curved posterolaterally, the axillary process pointed also posterolaterally. From the lateral perspective (Fig. 10B), the main femoral spur relatively short and slightly tapered, the axillary process half-moon to finger-shaped. The genual and tibial spurs with slightly concave margin and rounded distal apex, the genual spur located closer to distal article margin than tibial one. Setae on leg II simple except anterolateral setae on genu and tibia which barbed, femoral setae al1 thicker, al2 and ad2 thicker and shorter, whereas ad3 needle-shaped. Leg IV: trochanter without tooth or tubercle, posterodorsal and posterolateral setae on the femur shorter and thicker, posteroventral seta on the genu and tibia thickened and barbed. Other aspects of legs I-IV unremarkable.  Type deposition -Types are deposited in the Zoological Division of the Nature Education Centre, Jagiellonian University, Kraków, Poland.

Etymology
The specific name coronarius (lat. corona means wreath or crown) refers to the crown-like outline of the endogynial stipule.

Diagnosis
Female and male -Idiosoma weakly to moderately sclerotised; gland pore gv1 present; podonotum with 21 pairs of setae, opisthonotum with variable number of setae: from 23 pairs to 24 pairs and four supplementary setae located marginally; Tr IV with distally located low arcuate elevation.
Female -Gnathotectum vary, one-prong type (holotype) or three-prong type (paratype); presternal plates with long adaxial protrusions; the sternal shield anterior margin with shallow concavity; the anterior margins of the epigynial shield nearly straight or slightly wavy, posterolateral ones short and convex, the transverse band of less pigmented cuticle is located subapically, the internal (dorsal) surface of the epigynium without teeth; endogynial spherules bean-shaped and convergent anteriorly, in front of the spherules, a U-shaped structure with a pair of adaxially directed thorns present, thorn-shaped stipule (?) is located between the spherules.
Male -Gnathotectum trispinate with similar triangular prongs; genital lamina with the anterior margin concave and anterior corners rounded; presternal plates subrectangular, with the short anterior protrusions and rounded posterior margins; corniculi slim but with adaxial elevation; cheliceral fixed digit straight and narrow, its apex cut obliquely, apical part separate dorsally by an incision from the rest of the digit, between the apex and the pilus dentilis a lamellar protrusion followed by denticle is visible, behind the pilus dentilis the row of ca. 8 minute denticles is present; leg II femoral main spur and the axillary process finger-shaped, genual and tibial spurs conical, the former one somewhat smaller and located closer to the distal article margin.
Gnathosoma -Gnathotectum (Fig. 13D,E) variable in shape, in the paratype female is trispinate (Fig. 13E), all prongs similar, narrow and acute, in holotype it culminates aberrantly into a single prong (Fig. 13D). Corniculi conical, hypostome with 10 rows of denticles, hypostomatic and palpcoxal setae simple. Palptrochanter v1 seta simple, v2 barbed. Chelicera (Fig. 13F) -movable digit with four teeth, the proximal one much larger, fixed digit with two distant teeth in front of the pilus dentilis, two teeth behind the pilus dentilis, followed by an undulate lamella and a more solid edge proximally.
Legs -Setae ad on Tr I somewhat shorter and thicker. Leg II: anterolateral setae al1, al2 on Fe II shorter and thicker, anteroventral setae on Fe II and Ti II larger and barbed. Leg IV: posterolateral setae on the femur thick and short, posteroventral seta on the tibia and posterolateral setae on basitarsus and tarsus thickened and terminally barbed. Tr IV with a low, arcuate elevation located distally on the article dorsal side (Fig. 15C). Other aspects of legs   (Fig. 14A) -the anterior margin of genital lamina concave, the lateral ones slightly concave and the anterior corners rounded (Fig. 14A,B). Presternal plates with short anterior protrusions and posterior margins more or less rounded. Sternum with gland pores gv1 close to st3 setae level and relatively close to these setae bases, followed by two pairs of moderately visible thickenings of the sternal cuticle. Pore gv2 with two closely located openings, pores iv5 ca. halfway between st5 and ZV1 setae, ivo2, ivo3, and gv3 well discernible.
Gnathosoma -Gnathotectum (Fig. 14C) trispinate with all prongs similar and triangular. Corniculi (Fig. 14D,E) with arcuate elevation on the adaxial margin, hypostome with nine rows of denticles, hypostomatic and palpcoxal setae simple. Palptrochanter v1 seta simple, v2 finely barbed. Chelicera -when observed from the ventral side, mobile digits slightly curved adaxially, the fixed ones straight. Laterally (Fig. 14F,G), mobile digit with one tooth followed by an arcuate edge proximally, fixed digit straight and narrow with obliquely cut apex and dorsal incision at pilus dentilis level. The arcuate lamellar protrusion between the apex and the pilus dentilis followed by a denticle is present, with a row of ca. eight denticles located behind the pilus dentilis (Fig. 14G).
Legs -Leg II (Fig. 15A,B) spurred as follows: when viewed from the ventral side (Fig.  15A), the femoral main spur short, finger-shaped and nearly straight, the axillary process pointed posterolaterally, genual spur lenticular to roundish, the tibial one lenticular. The genual spur located closer to the article margin than the tibial one. From a lateral perspective (Fig.  15B), the femoral main spur and the axillary process finger-shaped and straight. Genual and tibial spurs conical, the second one slightly larger. Setae on leg II simple, except genual and tibial anterolateral setae (al1, al2) which can be finely barbed, femoral seta al1 thicker, al2 and ad2 thicker and shorter, whereas ad3 needle-like. Leg IV: trochanter with a weakly pronounced dorsal arcuate elevation in distal part of the article (Fig. 15D). Posterolateral setae on the femur thickened and short, posteroventral seta on tibia thickened and barbed terminally, some setae on basitarsus and tarsus thickened. Other aspects of legs I-IV unremarkable.
Type deposition -Types are deposited in the Zoological Division of the Nature Education Centre, Jagiellonian University, Kraków, Poland.

Etymology
The specific name trispinus refers to prominent three spines in the female genital region.

Taxonomic remarks
The examined material for Leptogamasus trispinus n. sp. is rather modest, since only two females (the holotype and one paratype) are available. The holotype female shows some unusual characters as compared with the paratype, i.e. lacking of podonotal setae R6 and the gnathotectum one-prong type. Both characters in the paratype female are typical for the genus. However, the holotype female was selected as the holotype since the other features are available and typical, especially all legs, which are incomplete in the female paratype.

Differential taxonomy
The Leptogamasus (Leptogamasus) new species currently described are more or less similar to several previously acknowledged species. As far as the females are concerned, L. (L.) chelatus n. sp. is somewhat similar to Leptogamasus (Leptogamasus) termitatus (Athias-Henriot, 1967) since both species share the following characteristics: internal (dorsal) surface of epigynium smooth, i.e. without teeth, the spherules subspherical and the endogynial stipule made up of two parts, i.e. anterior and posterior. However, the most distinct difference consists in the actual size and shape of the stipule. In L. (L.) termitatus posterior part of the stipule is very large and furcate, whereas in L. (L.) chelatus n. sp. is small and tooth-shaped. The males are also different, i.e. leg II spurs in both species are similar with regard to the femur and the tibia, to some extent, but genual spur in L. (L.) chelatus n. sp. is conical, whereas in L. (L.) termitatus is low and elongated axially. Chelicerae offer more distinct characteristics since the cheliceral fixed digit in L. (L.) chelatus n. sp. bears one tooth in front of the pilus dentilis, followed by edentate edge behind the pilus dentilis, whereas in L. (L.) termitatus fixed digit bears ca. two teeth in front and several fine teeth behind the pilus dentilis. Unfortunately, the fixed digit in a newly described species shows a conspicuous undulation when observed in the ventral perspective, even though this particular property is not encountered in L. (L.) termitatus.
Leptogamasus (L.) coronarius n. sp. and the most similar Leptogamasus (Leptogamasus) bidentoides (Athias-Henriot, 1967) females share some of the following characteristics: internal (dorsal) surface of epigynium with two teeth, the wall of endogynial sac without teeth and stipule of similar length and width, richly dentated on the surface and the anterior margin, Tr IV without tubercle or a tooth. These two species can be easily distinguished, owing to the shape of spherules, which are in L. (L.) coronarius n. sp. regularly rounded anteriorly and in L. (L.) bidentoides possess several tooth-like protrusions. The male in L. (L.) coronarius n. sp. cannot be compared with L. (L.) bidentoides male which is unknown, even though it is acknowledged to share some similarities with L. (L.) belligerens Witaliński, 1973 male with the armature of leg II somewhat similar. In the latter species, the cheliceral fixed digit features several fine teeth in front and behind of the pilus dentilis, whereas in L. (L.) coronarius n. sp. minute teeth are present behind the pilus dentilis only, in contrast to the anterior digit part bearing a single low tooth only. ) falsiviatus a row of denticles behind the pilus dentilis is absent and Ti II spur is L-shaped, whereas in L. (L.) sinuforceps fixed digit bears denticles but spurs on Ge II and Ti II are baton-shaped rather than conical as in L. (L.) trispinus n. sp. In L. (L.) cortinis a shallow incision of fixed digit is located some distance behind the pilus dentilis level, and digit dorsal margin bears a tubercle more apically. Moreover, the corniculi bear a distinct tubercle on the ventral side. In L. (L.) renogynialis cheliceral fixed digit differs from L. (L.) trispinus n. sp. since in the former species the dorsal incision is smaller and located between the pilus dentilis and the digit apex. Some similarities of females in L. (L.) trispinus n. sp. and L. (L.) paracarpaticus might suggest a comparison of the males. However, in the latter species, the cheliceral fixed digit does not show both the dorsal incision and the denticles behind the pilus dentilis.