Key to the world species of Holoparasitus Oudemans, 1936 (Acari: Parasitiformes: Parasitidae) Author Witaliński, Wojciech text Zootaxa 2017 4277 3 301 351 journal article 32824 10.11646/zootaxa.4277.3.1 426319f0-f361-4222-bf48-b093743b0b96 1175-5326 810605 70C1663B-F33F-42C1-99C9-FC3A0A4110DA Key to Holoparasitus species (Females) 1. Sternum split axially into the left and the right part ( Figs 11, 14 )....................... ( inornatus species-group ) … 2 - Sternum entire (e.g. Figs 1, 2 , 33 )......................................................................... 3 2. Endogynium visible as a transversal thin plate, elliptical or slightly triangular, with central thickening bearing on its dorsal side the centrifugally directed tubercles ( Figs 12, 13 )............................ H . inornatus ( Berlese, 1906 ) ( Figs 11–13 ) - Endogynium elliptical, axially oriented ( Fig. 14 )......................... H . megacalcaratus Schmölzer, 1995 ( Fig. 14 ) 3. Sternum with more or less pronounced axial band, granular or reticulated (e.g. Figs 33, 46, 50 , 52, 62 ), and/or the anterior margin of presternal plate serrated or coarsely granular ( Figs 15 , 32 , 52 ) (except H . giganteus ); lateral platelets free........................................................................................ ( mallorcae species-group )...4 - Axial granular or reticulated band on the sternum absent ( Fig. 2 ); anterior margin of the presternal plate smooth, lateral platelets free or fused to the ends of the presternal plate ( Figs 2 , 16, 17 )............................................. 23 4. Central prong of epigynium slender and considerably elongated ( Figs 18–20 ); endogynium with two teeth, covered ventrally by a dentate lamella ( Figs 19–21 ).......................................... H . siculus ( Berlese, 1906 ) ( Figs 18–21 ) - Central prong of epigynium moderately narrow and elongated (e.g. Figs 24, 27 , 34, 41, 48 )........................... 5 5. Gv1 glands absent, epigynium with a rounded apex and moderately sharp lateral prongs ( Figs 24, 27 ), endogynium a small sack ( Figs 25, 28, 31 ).................................................................................. 6 - Gv1 glands present, epigynium with a triangular apex and salient lateral prongs ( Figs 34, 41, 48 )...................... 7 6. Endogynium—a sack with finely dentate anterior margin ( Fig. 25 ).......... H . algiersensis Juvara-Bals, 2008 ( Figs 22–25 ) - Endogynium—a sack with a round, sclerotised part in its middle ( Figs 28, 29, 31 )...................................................................................................... H . eivissa Juvara-Bals, 2008 ( Figs 26–31 ) 7. Endogynium—a narrow tube dorso-ventrally oriented and visible as a very small ring ( Figs 35–39 ), sometimes open on one or both sides, when the tube should be obliquely oriented ( Figs 37, 38 ); ventral margin of the endogynium in most specimens bears triangular, asymmetric lamellar protrusion ( Figs 35, 36 ); ventrally, delicate endogynial lamella dentate anteriorly covers the endogynium ( Figs 35, 36, 39, 40 )................................... H . calpetanus Witaliński, 2016 ( Figs 32–40 ) - Endogynium saccular and large, its diameter similar or larger than the size of epigynium subapical structure (e.g. Figs 43 , 4 9 , 56 )................................................................................................. 8 8. Saccular endogynium with lateral walls forming two large dentated invaginations antero-dorsally directed; stout asymmetric prolongation located in the endogynium midline and protruded anteriorly ( Figs 43, 44 ); epigynium with broad and blunt apex ( Figs 41, 42 ).......................................................... H . lawrencei Hyatt, 1987 ( Figs 41–44 ) – Endogynium otherwise, with curtain-like structures or protrusions on its posterior margin; epigynium with triangular apex.. 9 9. Endogynium with two membranous curtain-like structures or two fleshy lobes ( Figs 49, 51 )......................... 10 - Endogynium with one or two protrusions on the posterior margin (e.g. Figs 56, 60, 66 , 70 ).......................... 11 10. Endogynium with two membranous curtain-like structures ( Fig. 49 ); epigynium with prominent triangular apex; subapical structure anteriorly rounded, with two tiny oval extensions near the apex base ( Fig. 48 )................................................................................................ H . mahnerti Juvara-Bals, 2008 ( Figs 45–49 ) - Endogynium with two arcuate lamellar prolongations ( Fig. 51 ); epigynium with a short, broad tip; subapical structure rectangular, with two membranous triangular extensions near the apex; sternal reticulation shows conspicuous loop-like pattern axially located ( Fig. 50 )..................................................... H . maritimus Hyatt, 1987 ( Figs 50, 51 ) 11. Endogynium with one protrusion on the posterior margin (e.g. Figs 56, 60, 65 ).................................... 12 - Endogynium with two protrusions or one deeply split protrusion on the posterior margin (e.g. Figs 70, 79, 80 )........... 14 12. Endogynium with a single finger-like, occasionally forked out protrusion and 7–8 denticles on its anterior and lateral walls ( Fig. 56 )........................................................... H . rifensis Juvara-Bals, 2008 ( Figs 52–56 ) - Endogynium with a single protrusion only, denticles on its walls absent.......................................... 13 13. Apex of the finger-like endogynial protrusion sharp ( Figs 60, 61 ); apex of the epigynium mucronate, epigynial subapical structure trapezoidal and well sclerotised, with small hyaline lateral extensions ( Figs 58, 59 )................................................................................................... H . lunae Juvara-Bals, 2008 ( Figs 57–61 ) - Apex of a slender endogynial protrusion usually sharp and split, ending at the anterior margin of the endogynium ( Figs 65–67 ); apex of the epigynium large and triangular; epigynial subapical structure formed by a sclerotised rectangular line under the apex, and by the two wing-like hyaline lateral extensions ( Fig. 63 )............. H . malleus Juvara-Bals, 2008 ( Figs 62–67 ) 14. Presternal plate non-serrated ( Fig. 68 ); endogynium with two short posterior protrusions far apart from each other ( Fig. 70 ); big species........................................................ H . giganteus Juvara-Bals, 2008 ( Figs 68–71 ) - Presternal plate more or less distinctly serrated; endogynial protrusions of different shape and size.................... 15 15. Presternal plate with a few minute denticles ( Fig. 73 ); gland pore gv1 located on a soft cuticle or on the posterior margin of the sternum; endogynial protrusions involute, unequal in size and distant of each other ( Fig. 76 ); epigynium with triangular apex and a subapical structure with a conspicuous, rectangular sclerotised line continued by the two trapezoidal membranous wings ( Figs 74, 75 )....................................................... H . lapalma Juvara-Bals, 2008 ( Figs 72–77 ) - Presternal plate distinctly serrated; gland pore gv1 always located on the sternum near the posterior margin; endogynial protrusions well developed; epigynial apex triangular, subapical structure otherwise..................................... 16 16. Endogynium—two protrusions of different length, with their bases very close to each other, or one deeply split protrusion ( Figs 79–81 )...................................................... H . variabilis Juvara-Bals, 2008 ( Figs 78–81 ) - Endogynium with two equally sized protrusions............................................................ 17 17. Gnathotectum large, triangular ( Fig. 82 ); large cup-like endogynium ( Fig. 84 ) with two protrusions, their apices medially oriented and distant of each other; trochanter IV with ventral protuberance.......... H . anaga Juvara-Bals, 2008 ( Figs 82–84 ) - Gnathotectum trispinate; endogynium otherwise; trochanter IV without any protuberance........................... 18 18. Endogynium with the two horn-like or triangular protrusions ( Figs 88, 90, 91 ).................................... 19 - Endogynium with the two finger-like, rather long protrusions ( Figs 95, 98 , 101, 102, 104 )........................... 20 19. Endogynium with two distant horn-like protrusions ( Fig. 88 ); epigynial apex not prominent, subapical structure with oval wings ( Fig. 87 ); palptrochanter seta v1 pectinate, v2 barbed, with a protuberance between them; palpgenu with tubercle close to the anterolateral seta............................................... H . vaucheri Juvara-Bals, 2008 ( Figs 85–88 ) - Endogynium with the two triangular protrusions close to each other ( Figs 90, 91 ); epigynial subapical structure extending into the fan-like wings ( Fig. 92 ).............................................. H . franzi Juvara-Bals, 2008 ( Figs 89–92 ) 20. Endogynium with no teeth on either lateral or anterior walls, endogynial protrusions distant, with a gap of 40–45 µm between their bases ( Figs 95, 96 )........................................... H . canariensis Juvara-Bals, 2008 ( Figs 93–96 ) - Endogynium with teeth on the lateral and anterior walls, long endogynial protrusions straight or sinuous, their tips reaching the anterior margin of the endogynium and their bases close to each other ( Figs 98 , 101, 104 )........................... 21 21. Lamellar endogynial protrusions straight ( Fig. 98 ), distance between their bases ca. 10 µm; palptrochanter with simple v1 , barbed v2 , and a protuberance between them............................. H . mallorcae Juvara-Bals, 1975 ( Figs 97, 98 ) - Lamellar endogynial protrusions sinuous ( Figs 101, 102, 104 ), their bases very close to each other; palptrochanter with pectinate v1 and barbed v2 . ................................................................................ 22 22. Endogynium with several (3–7) denticles and two protrusions extending beyond the anterior endogynial margin ( Figs 101, 102 ).............................................. H . ellipticus Juvara-Bals & Witaliński, 2000 ( Figs 15 , 99–102 ) - Endogynium ( Fig. 104 ) with the two lateral denticles, protrusions ending at the anterior endogynial margin...................................................................... H . gibber Juvara-Bals & Witaliński, 2000 ( Figs 103, 104 ) 23. Endogynium bubble-shaped (e.g. Figs 109, 113, 119 )................................... ( caesus species-group) ...24 - Endogynium non-bubble-shaped......................................................................... 27 24. Tube leading to the endogynial sac relatively wide, ca. 1/2 of the sac diameter ( Figs 107, 109 )......................................................................................... H . ampullaris Witaliński, 1994 ( Figs 105–109 ) - Tube leading to the sac narrow, its diameter not exceeding 1/3 of the sac diameter ( Figs 113, 116, 119 ); subapical epigynial structure compact ( Figs 111, 115, 118 ).................................................................... 25 25. Subapical structure of the epigynium with a central rectangular element ( Figs 111, 112 ); endogynial sac spherical or slightly ellipsoidal ( Figs 112, 113 ).......................................... H . caesus Micherdziński, 1969 ( Figs 110–113 ) - Central rectangular element in the epigynial subapical structure absent.......................................... 26 26. Endogynial sac ellipsoidal ( Fig. 116 )............................... H . tuberculatus Juvara-Bals, 1975 ( Figs 114–116 ) - Endogynial sac posteriorly flattened ( Fig. 119 )......................... H . fortunatus Juvara-Bals, 1975 ( Figs 117–119 ) 27. Endogynium cup-shaped, circular (e.g. Figs 121, 125, 133 , 158 ) or rhombic ( Figs 144, 146 ) in the ventral perspective..... 28 - Endogynium non-cup-shaped, neither circular nor rhombic in its outline (e.g. Figs 201, 208 , 211, 216, 225 )............. 45 28. Endogynium without any thorns, teeth or lamellar protrusions, except a long band-like structure (endogynial lamella?) located posteriorly ( Fig. 121 )................................................... H . inventus Vinnik, 1994 ( Figs 120, 121 ) - Endogynium with thorns, teeth and/or lamellar protrusions.................................................... 29 29. Subapical structure of epigynium with a distinct, "inverted–T" thickening located axially ( Figs 123, 127 )............................................................................................. ( peraltus species-group )...30 - "Inverted-T" thickening absent.......................................................................... 31 30. Endogynium with narrow lamellar protrusions sometimes branched distally; endogynial sac bears many dents or thorns ( Figs 125, 126 ); the anterior thickenings of epigynial subapical structure well pronounced, the posterior thickenings oriented parallelly to each other ( Figs 122, 123 )........................... H . nonaltus Witaliński & Skorupski, 2003 ( Figs 122–126 ) - Endogynium with broad lamellar protrusions indented terminally; endogynial sac with reticulation, but dents or thorns absent ( Fig. 129 ); the anterior thickenings of subapical epigynial structure not discernible, the posterior thickenings run convergently ( Fig. 127 ).......................................................... H . peraltus ( Berlese, 1906 ) ( Figs 127–129 ) 31. Endogynium with thorns gathered in the two lateral (left and right) groups ( Figs 133,137, 141, 146 )................... 32 - Endogynium with the two large thorns located posteriorly, and other thorns or denticles; the latter, if numerous, not arranged in lateral groups but distributed more regularly around the endogynial sac entrance (e.g. Figs 154, 164 , 167, 187 , 195, 197 ).. 35 32. Posterior endogynial margin regular, without two lamellae protruding anteriorly ( Figs 133, 137 ); posterior paragynial lobes in close proximity, or moderately distant of each other......................................................... 33 - Posterior endogynial margin forms the two broad, partially overlapped elliptical or leaf-shaped lamellae covering most of the endogynial entrance ( Figs 141, 144 ); posterior paragynial lobes (and the posterior subapical thickenings of the epigynium) distant ( Figs 138, 139, 143 ).................................................. ( hemisphaericus species-group )...34 33. Endogynial thorns directed anteriorly; posterior paragynial lobes in close proximity; distal margin of Tr IV with lamellar process ending in one or two prongs ( Figs 130, 131 ); endogynium with a pair of double lateral teeth located posteriorly and a pair of smaller teeth located more anteriorly, spherular organ occupies the posterior half of endogynium, whereas the endogynial sac is in the anterior half ( Fig. 133 )........................ H . cornutus Juvara-Bals & Witaliński, 2000 ( Figs 130–133 ) - Endogynial thorns directed posteriorly ( Fig. 137 ); posterior paragynial lobes more distant; lamellar process at the distal margin of Tr IV absent................................................. H . intermedius ( Holzmann, 1969 ) ( Figs 134–137 ) 34. Endogynium roundish in its outline ( Fig. 141 )...................... H . hemisphaericus ( Vitzthum, 1923 ) ( Figs 138–142 ) - Endogynium rhombic ( Figs 144, 146 )............................ H . rhombogynialis Witaliński, 2006 ( Figs 143–146 ) 35. Postcoxal gv2 glands empty into the large foveolae formed by a thickened cuticle ( Figs 147, 151, 152, 156 )............ 35 - Evident foveolae absent ( Fig. 162 ) (but a cuticle surrounding gv2 pores in H . pseudoperforatus may occasionally be thickened)................................................................................................... 38 36. Hyaline endogynial lamella with a thickened, sinuous anterior edge ( Figs 149, 150 ); epigynial subapical structure and central prong relatively large; roundish endogynial sac visible as a distinct structure smaller than the endogynium ( Fig. 149 )......................................................................... H . dallaii Witaliński, 1994 ( Figs 147–150 ) - The edge of endogynial lamella not thickened, frequently poorly visible ( Figs 154, 155, 160 ); epigynial subapical structure and central prong of normal size; endogynial sac as large as the endogynium, not discernible as a smaller distinct structure.... 37 37. Idiosoma strikingly spherical; anterior epigynial margins on the sides of the central prong deeply concaved ( Fig. 153 )........................................................................ H . globosus Witaliński, 1994 ( Figs 151–155 ) - Idiosoma elongated; anterior epigynial margins straight ( Fig. 157 ) … H . fovealis Witaliński & Skorupski, 2007 ( Figs 156–160 ) 38. Endogynium relatively large, circular ( Fig. 164, 165 ); margins of endogynium and endogynial sac distinctly visible as the two concentric circles; both posterior main thorns protrude more or less centripetally at ca. 120o from sac margin; anterior main thorn absent; two to four additional denticles can be distributed bilaterally at the sac entrance, occasionally one or two small denticles, located randomly and much deeper in the sac, are encountered; epigynial central prong weakly sclerotised and less pigmented apically ( Figs 162, 163 )........................... H . amiatus Witaliński & Skorupski, 2007 ( Figs 161–165 ) - Endogynium at least with the two posterior and one anterior main thorns; the anterior main thorn or dent usually smaller, located axially and directed backward; additional denticles usually present and distributed regularly on both sides, between posterior and anterior main thorns (e.g. Figs 167, 173, 186 , 195, 197 )................... ( calcaratus species-group )...39 39. Endogynium with the three main thorns, endogynial sac visible as an elliptical, distinct structure with many long centripetal thorns protruding from the wall ( Figs 167–169 ).......................... H . rotulifer ( Willmann, 1940 ) ( Figs 166–169 ) - The wall of the endogynial sac without any thorns or dents.................................................... 40 40. Endogynium with distinct spherular organ protruding from the posterior surface of the endogynium into the endogynial lumen (e.g. Figs 173–175, 184–187 )........................................................................... 41 - Distinct spherular organ absent (e.g. Figs 189, 193, 195, 197 ).................................................. 42 41. Anterior part of endogynium perimeter not covered from its ventral side with any lamella protruding from the endogynial margin; spherular organ triangular or subtriangular, frequently more or less pointed anteriad and protruding into the endogynial lumen ( Figs 173–175 ); the apex of epigynial plate blunt, tetrangular or rounded ( Figs 170–172 )................................................................................... H . pseudoperforatus ( Berlese, 1906 ) ( Figs 170–175 ) - The anterior margin of the endogynium forms a thin bilobate lamella partly covering the endogynium entrance ( Fig. 183 ), with the lobes separated by a distinct incision (usually with an additional triangular protrusion); spherular organ noticeable, its anterior margin arcuate in its central part and straight laterally ( Figs 184–187 ); the central prong of epigynium present, but located ventrally over the anterior, triangular part of the subapical epigynial thickening, thus poorly discernible ( Figs 179–182 ); the lamellar extensions of the subapical epigynial structure projecting laterally beyond the epigynial apex, occassionally also anteriorly ( Figs 179, 182 )............................................. H . excipuliger ( Berlese, 1906 ) ( Figs 176–187 ) 42. Endogynium with the three main thorns, large and of a similar size; additional 2–3 dents are located on each side between the anterior and posterior main thorns ( Figs 189, 190 ); central epigynial prong long and narrow, lateral protrusions of subapical epigynial structure wing-like and elongated, but practically not extending beyond the epigynial margin ( Fig. 188 ).............................................................. H . kerkirensis Witaliński & Skorupski, 2002 ( Figs 188–191 ) - Endogynium with the posterior main thorns evidently larger than the anterior one; additional lateral dents (1–2 on each side) may also be encountered............................................................................... 43 43. Epigynium ( Fig. 192 ) with the central prong short; lateral hyaline protrusions of the subapical epigynial structure wing-like and elongated, rounded terminally and extending far beyond the epigynium margin; the thickening of the subapical epigynial structure forms the four sides of a regular pentagon, with the anterior sides straight ( Fig. 192 ); endogynium irregularly roundish ( Fig. 193 ).................................................... H . pollicipatus ( Berlese, 1903 ) ( Figs 192, 193 ) - The thickening of the subapical epigynial structure with the anterior parts evidently wavy ( Figs 194, 196 ), lamellar lateral protrusions only moderately extending beyond the epigynium margin, three-sided, with the posterior sides concave ( Figs 194, 196 ); endogynium regularly roundish..................................................................... 44 44. The posterior main thorns of endogynium curved centripetally and settled on a joint, broad base ( Fig. 195 ).................................................................................. H . calcaratus ( Koch, 1839 ) ( Figs 194, 195 ) - The posterior main thorns of the endogynium straight and settled independently; endogynial sac visible as a distinct smaller circle ( Fig. 197 )..................................... H . paradisiacus Witaliński & Skorupski, 2003b ( Figs 196, 197 ) 45. The anterior margin of the sternum between st1 setae better sclerotised, partly overlaps a presternal plate ( Figs 198, 202 ); endogynium visible as an arcuate sclerotised band with an axially located opening ( Figs 200, 201, 204, 205 ).............................................................................................. ( annulus species-group )...46 - Sternum anterior margin developed normally; endogynium different from the above................................ 47 46. Gland pores gv1 present; opening in the cuticular band of endogynium with a subcircular anterior margin, a dentate lamellar structure (endogynial lamella?) located laterally to the opening not discernible............................................................................................ H . annulus Juvara-Bals & Witaliński, 2006 ( Figs 198–201 ) - Gland pores gv1 absent; opening in the cuticular band of endogynium usually with an undulate anterior margin, dentate lamellar structure (endogynial lamella?) can be discernible laterally to the opening........................................................................................... H . sardensis Juvara-Bals & Witaliński, 2006 ( Figs 202–205 ) 47. Endogynium large, irregularly semicircular, with a split lamellar protrusion, anteriorly pointed and dentate ( Fig. 208 ); posterior paragynial lobes distant of each other ( Fig. 206 )......................... H . excisus ( Berlese, 1906 ) ( Figs 206–208 ) - Endogynium very small (e.g. Figs 209, 215, 219, 223 ); posterior paragynial lobes in close proximity of each other or distant.................................................................................................... 48 48. Posterior paragynial lobes distant; central epigynial prong large and sharp, the thickening of subapical epigynial structure not discernible ( Fig. 210 ); endogynium subcircular with two "orifices" ( Fig. 211 )... H . vasilei Juvara-Bals, 1995 ( Figs 209–211 ) - Posterior paragynial lobes in close proximity of each other ( Figs 214, 223 ); central epigynial prong wider than the thickening of the subapical epigynial structure, which is narrow and compact, anteriorly tripartite or rounded (e.g. Figs 213, 218 )..... 49 49. The thickening of the subapical epigynial structure—anteriorly tripartite ( Figs 212–214 ), lateral hyaline protrusions extending a little beyond the epigynial margin....................................... H . fanes Witaliński, 2016 ( Figs 212–216 ) - The thickening of the subapical epigynial structure—anteriorly rounded ( Figs 218, 221 ) … ( crassisetosus species-group ). 50 50. Ti IV with the thickened seta pv1 ( Fig. 217 ); lateral hyaline protrusions of the subapical epigynial structure roundish and extending a little beyond the epigynial margin ( Fig. 218 )... H . crassisetosus Juvara-Bals & Witaliński, 2000 ( Figs 217–220 ) - Ti IV with normal setae pv1 ; prominent lateral hyaline protrusions of the subapical epigynial structure wing-like and directed either laterally, or anterolaterally ( Figs 221 )................ H . digitiformis Juvara-Bals & Witaliński, 2000 ( Figs 221–227 )