Holoparasitus, Witaliński, 2017

Witaliński, Wojciech, 2017, Key to the world species of Holoparasitus Oudemans, 1936 (Acari: Parasitiformes: Parasitidae), Zootaxa 4277 (3), pp. 301-351 : 310

Key to Holoparasitus species (Males)

1. Sternum with an excipulum (e.g. Figs 229, 230 ); if excipulum is not encountered, a distinct, not-reticulated elliptic area1 is found on the sternum at the coxae II level ( Fig. 231 ).......................................................... 2

- Excipulum or its equivalent, a non-reticulated area on the sternum absent (e.g. Fig. 228 )............................ 15

2. Gnathotectum with the central prong well pronounced, acute, flanked by the two lateral prongs, sometimes minute or altogether absent (e.g. Figs 234, 239 )......................................................................... 3

- Gnathotectum central prong lobate ( Fig. 235 ), rounded or obtuse anteriorly, either spineless, or with minute spines (1–3) apically (e.g. Figs 281 , 310 ); lateral small prongs present; a mobile digit of chelicera much thicker in the basal half (e.g. Figs 282 , 288 , 311 ) (in H. rotulifer ( Willmann, 1940) gnathotectum with a single central prong only, and a movable digit of chelicera with a tubercle on the external (ventral) margin)..................................... ( calcaratus species-group)...9

3. Postcoxal glands gv2 empty into the cuticular foveolae ( Figs 231 , 243 )........................................... 4

- Cuticular foveolae behind coxae IV—not encountered........................................................ 5

4. Sternum with a well-developed excipulum ( Fig. 243 ); corniculi slightly indented; dorsal edge of fixed digit of chelicera normal ( Fig. 244 )........................................................... H. dallaii Witaliński, 1994 ( Figs 243–245 )

1. In H. globosus excipulum is either weakly visible or only not-reticulated area in that place is present - Sternum with a non-reticulated area ( Fig. 231 ); in some specimens the traces of excipulum can be detected; corniculi not indented; dorsal edge of fixed digit of chelicera with an elevation ( Fig. 246 )..................................................................................................... H. globosus Witaliński, 1994 ( Figs 231 , 246, 247 ) 5. Gnathotectum with all prongs similarly well pronounced ( Fig. 248 ); Ti II with additional tubercle dorsally ( Fig. 249 ); fixed digit of chelicera knife-shaped and toothless ( Fig. 250 )...................... H. cultriger ( Berlese, 1906) ( Figs 248–250 ) - Gnathotectum with the central prong well pronounced, whereas the lateral ones are either small or altogether absent ( Fig. 251 ); Ti II with a regular prong only; fixed digit of chelicera either toothless or with several dents.......................... 6 6. Excipulum horseshoe-shaped, its anterior thickened margin complete ( Figs 252 , 259 ); sternal reticulation unremarkable; hypostomatic setae not on a separate piece of cuticle ( Fig. 253 ); fixed digit of chelicera practically toothless............................................................................................ ( peraltus species-group)...7 - Excipulum “open” at both anterior and posterior ends and flanked by two very prominent lines of reticulation running parallel to the excipulum anterolateral margins ( Figs 265 , 270 ); hypostomatic setae on a separate piece of cuticle (like in Fig. 232 ); fixed digit of chelicera with a row of many tiny denticles......................... ( hemisphaericus species-group)...8 7. Spur on Ti II singular, long and low ( Figs 256, 257 ); pedipalp trochanter with seta v1 simple, v2 stout and terminally brushy, settled on eminence not far from v1 ( Fig. 254 )............................. H. peraltus ( Berlese, 1906) ( Figs 251–258 ) - Spur on Ti II bipartite ( Figs 262, 263 ): proximal part with a sigmoidal edge, distal one forming a tooth-like structure; seta v2 pectinate distally, neither stout nor located on eminence close to v1. ................................................................................................... H. nonaltus Witaliński & Skorupski, 2003a ( Figs 259–264 ) 8. Spur on Ti II ( Figs 267–269 ) ending distally at the segment margin and visible as an arcuate tape from both the ventral and the lateral sides; the cuticle located behind the spur (posteriorly) is weakly pigmented ( Figs 268, 269 )................................................................................. H. rhombogynialis Witaliński, 2006 ( Figs 265–269 ) - Spur on Ti II ending far behind the distal margin of the segment ( Figs 272, 273 )................................................................................................ H. hemisphaericus ( Vitzthum, 1923) ( Figs 270–273 ) 9. Excipulum strong, well pronounced, with the lateral margins partially overlapped by a thin cuticular lamellae (e.g. Figs 274, 280 )............................................................................................... 10 - Excipulum weakly to moderately pronounced, its lateral margins not overlapped by the cuticular lamellae (e.g. Figs 298 , 308 )................................................................................................... 11 10. Gnathotectum central lobe pointed; spurs on Ge II large and high, the ones on Ti II smaller and low ( Fig. 278 ); very narrow movable digit of chelicera with semicircular tubercle on the external (ventral) edge ( Fig. 276 ).......................................................................................... H. rotulifer ( Willmann, 1940) ( Figs 274–278 ) - Gnathotectum central lobe obtuse ( Fig. 281 ); fixed digit of chelicera with one-fourth to one-third distal part—narrower ( Fig. 282 ), in dorso-ventral perspective a tubercular to tooth-like protrusion directed laterally at pilus dentilis level is visible ( Fig. 283 ); movable digit without any tubercle on the external (ventral) edge... H. excipuliger ( Berlese, 1906) ( Figs 229 , 279–287 ) 11. The pores of postcoxal glands gv2 surrounded by a thickened cuticle forming foveolae ( Fig. 293 )..................... 12 - Cuticular foveolae around the gv2 gland pores – absent....................................................... 13 12. Axillary process on Fe II asymmetric and bent towards the segment ( Figs 290, 291 ); cheliceral fixed digit with a lateral triangular projection ( Fig. 289 ), seta v2 on pedipalp trochanter pectinate distally on both sides and thicker than v1. ........................................................................ H. pseudoperforatus ( Berlese, 1906) ( Figs 288–292 ) - Axillary process on Fe II symmetric ( Figs 295, 296 ); cheliceral fixed digit without a lateral projection ( Fig. 294 ).......................................................................... H. pollicipatus ( Berlese, 1903) ( Figs 293–297 ) 13. Spurs on Ge II and Ti II very similar, low and ridgelike ( Figs 300, 301 )................................................................................................ H. kerkirensis Witaliński & Skorupski, 2002 ( Figs 298–301 ) - Spurs on Ge II and Ti II different........................................................................ 14 14. Corniculi slender with an adaxial border only slightly sinuous; movable digit of chelicera with one large proximal tooth followed by 4–6 minute distal teeth ( Fig. 302 ); main spur on the femur with an apical segment blunt terminally and quadrilateral, if observed from a ventral perspective ( Figs 303, 306 )................... H. calcaratus ( Koch, 1839) ( Figs 230 , 302–307 ) - Corniculi with a small protuberance on the adaxial border ( Fig. 309 ); movable digit of chelicera with one large proximal tooth followed by 2 smaller distal teeth ( Fig. 311 )............... H. paradisiacus Witaliński & Skorupski, 2003b ( Figs 308–314 ) 15. Ti II with several spurs, the two ventral ones huge ( Fig. 316 )......... H. karawankianus ( Schmölzer, 1991) ( Figs 315, 316 ) - The two huge spurs on Ti II—not encountered.............................................................. 16 16. Hypostomatic setae on a piece of cuticle separated laterally and posteriorly with narrow incisions................... 17 - Incisions in a cuticle laterally to and behind the hypostomatic setae – absent...................................... 46 17. Postcoxal gv2 glands empty into the cuticular foveolae ( Fig. 317 ).. H. fovealis Witaliński & Skorupski, 2006 ( Figs 317–320 ) - Cuticular foveolae behind coxae IV – absent............................................................... 18 18. Hypostome with a central part distinctly sclerotised, tongue-like (e.g. Figs 340 , 391, 398 ) or extended anteriorly2 ( Fig. 349 )............................................................................ ( mallorcae species-group) 3...19 - Central part of the hypostome triangular and weakly sclerotised (e.g. Figs 232, 233 , 451 , 465 )........................ 39 19. Glands gv1 absent; genital lamina rounded, heavily sclerotised on the inner face ( Figs 321, 326, 327, 332 ); microsclerite bearing tritosternum large and trapezoidal..................................................................... 20 - Gland gv1 present; genital lamina of another shape and rather weakly sclerotised, largely transparent (e.g. Figs 373 , 405, 411 ); 2. Similar in appearance to those in the Rhabdocarpais males

Figures

FIGURES 228 – 242. Males. 228 Holoparasitus cornutus, genital lamina and sternogenital shield, II-IV openings for coxae II to IV, iv 1 – iv 3 sternal pores, st 1 – st 4 sternal setae, gv 1 opening of gland gv 1; 229 Holoparasitus excipuliger, genital lamina and sternogenital shield, gv 2 pore of gland gv 2; 230 Holoparasitus calcaratus, genital lamina and sternogenital shield; 231 Holoparasitus globosus, sternogenital shield; 232 Holoparasitus sp., pedipalp trochanter and gnathosoma, ventrally; 233 Holoparasitus crassisetosus, pedipalp trochanter, femur and hypostome; 234 – 240 Holoparasitus spp., gnathotectum, different shapes; 241 genital lamina; 242 femur, genu and tibia of leg II, anterolaterally. Scale bars: 0.1 mm.

FIGURES 270 – 287. Males. Holoparasitus hemisphaericus: 270 sternogenital shield with excipulum; 271 chelicera, antiaxially; 272 Fe II, Ge II and Ti II, anterolaterally; 273 ibid., ventrally (arrow – anterolateral side). Holoparasitus rotulifer: 274, 275 excipulum on sternogenital shield; 276 movable digit of chelicera; 277 fixed digit of chelicera; 278 Fe II, Ge II and Ti II, posterolaterally. Holoparasitus excipuliger: 279, 280 excipulum on sternogenital shield; 281 gnathotectum; 282 chelicera, antiaxially; 283 apical part of chelicera in ventral perspective; 284 Fe II, Ge II and Ti II, anterolaterally; 285, 286 Ge II and Ti II, ventrally (arrow – anterolateral side); 287 Fe II, ventrally (arrow – anterolateral side). Scale bars: 0.1 mm.

FIGURES 302 – 314. Males. Holoparasitus calcaratus: 302 chelicera, adaxially; 303 main spur and axillary process on Fe II, ventral perspective; 304 spurs on Fe II, Ge II and Ti II, posterolaterally; 305 ibid., anterolaterally; 306 Fe II, Ge II and Ti II, ventrally (arrow – anterolateral side); 307 Ge II and Ti II, posterolaterally. Holoparasitus paradisiacus: 308 genital lamina and sternogenital shield with excipulum; 309 gnathosoma, ventrally; 310 gnathotectum; 311 chelicera, adaxially; 312 – 314 Fe II, Ge II and Ti II, respectively, anterolaterally. Scale bars: 0.1 mm.

FIGURES 288 – 301. Males. Holoparasitus pseudoperforatus: 288, 289 chelicera, two aspects; 290 spurs on Fe II, Ge II and Ti II; 291 Fe II, Ge II and Ti II, anterolaterally; 292 Fe II, ventrally (arrow – anterolateral side). Holoparasitus pollicipatus: 293 gv 2 pore in foveola behind opening for coxa IV; 294 chelicera, antiaxially; 295 Fe II, Ge II and Ti II, posterolaterally; 296 ibid, anterolaterally; 297 Fe II and Ge II, ventrally (arrow – anterolateral side). Holoparasitus kerkirensis: 298 excipulum on sternogenital shield; 299 Fe II, posterolaterally; 300, 301 Ge II and Ti II, respectively, posterolateral view. Scale bars: 0.1 mm.

FIGURES 243 – 258. Males. Holoparasitus dallaii: 243 idiosoma ventral side; 244 chelicera, antiaxially; 245 Fe II, Ge II and Ti II, anterolaterally. Holoparasitus globosus: 246 chelicera, adaxially; 247 Fe II, Ge II and Ti II, anterolaterally. Holoparasitus cultriger: 248 gnathotectum; 249 Fe II, Ge II and Ti II, posterolaterally; 250 chelicera. Holoparasitus peraltus: 251 gnathotectum; 252 excipulum; 253 hypostome; 254 pedipalp trochanter, ventrally; 255 chelicera, adaxially; 256 Fe II, Ge II and Ti II, posterolaterally; 257 spurs on Fe II, Ge II and Ti II; 258 main spur and axillary process in ventral perspective (arrow – anterolateral side). Scale bars: 0.1 mm.

FIGURES 259 – 269. Males. Holoparasitus nonaltus: 259 genital lamina and sternogenital shield with excipulum; 260 chelicera, adaxially; 261 movable digit of chelicera; 262 Fe II, Ge II and Ti II, anterolaterally; 263 spur on Ti II, posterolaterally; 264 spur on Ge II, posterolaterally. Holoparasitus rhombogynialis: 265 sternogenital shield with excipulum; 266 chelicera, adaxially; 267 Fe II, Ge II and Ti II, anterolaterally; 268 ibid., ventral aspect; 269 Ti II, ventrally. Scale bars: 0.1 mm.

FIGURES 315 – 320. Males. Holoparasitus karawankianus: 315 gnathosoma, ventrally; 316 leg II, posterolaterally. Holoparasitus fovealis: 317 genital lamina and sternogenital shield; 318 gnathotectum; 319 chelicera, adaxially; 320 Fe II, Ge II and Ti II, anterolaterally. Scale bars: 0.1 mm.

FIGURES 336 – 345. Males. Holoparasitus franzi: 336 pedipalp trochanter; 337 gnathotectum; 338 chelicera, antiaxially; 339 Fe II, Ge II and Ti II, anterolaterally. Holoparasitus lawrencei: 340 gnathosoma and pedipalp trochanters, ventrally; 341 gnathotectum; 342 chelicera, adaxially; 343 movable digit of chelicera, antiaxially; 344 spurs on Fe II, Ge II and Ti II; 345 Fe II, Ge II and Ti II, posterolaterally. Scale bars: 1, 4 – 10 0.1 mm, 2, 3 0.05 mm.

FIGURES 391 – 401. Males. Holoparasitus calpetanus: 391 gnathosoma, pedipalp trochanter and femur in ventral perspective; 392 gnathotectum; 393 chelicera, antiaxially; 394 genital lamina; 395 Fe II, Ge II and Ti II, anterolaterally; 396 Ti II, posterolaterally; 397 Fe II, Ge II and Ti II, ventrally (arrow – anterolateral side). Holoparasitus mallorcae: 398 gnathosoma and pedipalp trochanter, ventrally; 399 gnathotectum; 400 chelicera, antiaxially; 401 Fe II, Ge II and Ti II, anterolaterally. Scale bars: 0.1 mm.

FIGURES 346 – 360. Males. Holoparasitus gibber: 346 chelicera, adaxially; 347 ibid., antiaxially; 348 Fe II, Ge II and Ti II, anterolaterally. Holoparasitus ellipticus: 349 gnathosoma and pedipalp trochanter, ventrally; 350 gnathotectum; 351 chelicera, adaxially; 352 ibid., antiaxially; 353 Fe II, Ge II and Ti II, anterolaterally; 354 Ge II and Ti II, ventrally (arrow – anterolateral side). Holoparasitus anaga: 355 pedipalp trochanter and corniculus; 356 gnathotectum; 357 chelicera, adaxially; 358 ibid., antiaxially; 359 main spur and axillary process on Fe II, anterolaterally; 360 Ge II and Ti II, anterolaterally. Scale bars: 34 6 – 354, 359, 360 0.1 mm, 355 – 358 0.05 mm.

FIGURES 443 – 454. Males. Holoparasitus excisus: 443 gnathotectum; 444 chelicera, adaxially; 445 pedipalp trochanter and femur, ventrally; 446 Fe II, Ge II and Ti II, anterolaterally. Holoparasitus apenninorum: 447 chelicera, adaxially; 448 chelicera, antiaxially; 449 Fe II, Ge II and Ti II, posterolaterally. Holoparasitus amiatus: 450 genital lamina and anterior part of sternogenital shield; 451 pedipalp trochanter and gnathosoma, ventrally; 452 chelicera, antiaxially; 453 spurs on Fe II, Ge II and Ti II in anteroventral perspective (arrow – anterolateral side); 454 Fe II, Ge II and Ti II, anterolaterally. Scale bars: 0.1 mm.

FIGURES 465 – 475. Males. Holoparasitus crassisetosus: 465 gnathosoma with pedipalp trochanters and femurs, ventrally; 466 corniculus; 467 Ti IV, thickened seta; 468 Fe II, Ge II and Ti II, posterolaterally. Holoparasitus annulus: 469 gnathotectum; 470 chelicera, antiaxially; 471 pedipalp trochanter, corniculi and hypognathal groove; 472 corniculus; 473 genital lamina; 474 Fe II, Ge II and Ti II, anterolaterally; 475 ibid., ventrally (arrow – anterolateral side). Scale bars: 0.1 mm.

FIGURES 321 – 335. Males. Holoparasitus singularis: 321 genital lamina; 322 gnathotectum and corniculu; 323 chelicera, antiaxially; 324 ibid., adaxially; 325 Fe II, Ge II and Ti II, anterolaterally. Holoparasitus algiersensis: 326, 327 genital lamina; 328 chelicera, antiaxially; 329 movable digit of chelicera; 330 corniculus; 331 Fe II, Ge II and Ti II, anterolaterally. Holoparasitus eivissa: 332 genital lamina; 333 chelicera, antiaxially; 334 ibid., adaxially; 335 Fe II, Ge II and Ti II, ventrally (arrow – anterolateral side). Scale bars: 0.1 mm.

FIGURES 361 – 375. Males. Holoparasitus giganteus: 361 pedipalp trochanter and corniculus; 362 gnathotectum; 363 chelicera, antiaxially; 364 Fe II, Ge II and Ti II, posterolaterally. Holoparasitus lapalma: 365 pedipalp trochanter and corniculus; 366 gnathotectum; 367 chelicera, antiaxially; 368 Fe II, posterolaterally; 369 Ge II and Ti II, posterolaterally; 3 70 Ti II, ventrally (arrow – anterolateral side). Holoparasitus canariensis: 371 pedipalp trochanter and corniculus; 372 gnathotectum, different aspects; 373 genital lamina; 374 Fe II, Ge II and Ti II, anterolaterally; 375 Ge II and Ti II, ventrally (arrow – anterolateral side). Scale bars: 361, 362, 364 – 366, 368 – 375 0.1 mm, 363, 367 0.05 mm.

FIGURES 402 – 413. Males. Holoparasitus variabilis: 402 gnathotectum; 403 corniculus; 404 pedipalp trochanter and femur, ventrally; 405 genital lamina; 406 chelicera, antiaxially; 407 Fe II, Ge II and Ti II, anterolaterally. Holoparasitus malleus: 408 gnathotectum; 409 corniculus; 410 pedipalp trochanter and femur, ventrally; 411 genital lamina; 412 chelicera, antiaxially; 413 Fe II, Ge II and Ti II, anterolaterally. Scale bars: 402, 404, 405, 407, 410, 411, 413 0.1 mm, 403, 406, 408, 409, 412 0.05 mm.