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Maxiniini tribe n. Type genus: Maxinia Lindquist & Makarova, 2012. Diagnosis. Adult dorsal shield entire, robust, broad, without lateral incisions. Idiosomal setae simple. Female with sternal shield consolidated with endopodal sclerites between coxae I���II, and with free endopodal fragments between coxae II���III and free strongly developed strips between coxae III���IV; male sternitigenital shield fused with all endopodal sclerites beside coxae II���IV. Female with an apomorphically expansive ventrianal shield encompassing metapodal platelets anterolaterally and including setae JV 1 anteriorly. Male with sternitigenital shield abutting an expansive ventrianal shield that incorporates metapodal platelets. Peritreme slightly shortened, its margins smooth. Subcapitulum with narrow deutosternal groove; hypostomatic setae hp -1 and hp -3 not elongated and whip-like. Palp trochanter setae similar in size. Genu III with only seven setae (lacking pv -1). Male spermatodactyl short, tube-like. Tarsus I with claws. None of setae on tarsi II to IV conspicuously differentiated by shape or size. Genera included: Maxinia., Published as part of Makarova, Olga L., Marchenko, Irina I. & Lindquist, Evert E., 2021, Distribution, habitats, and redescription of the rare mite species Iphidonopsis sculptus Gwiazdowicz, 2004 (Mesostigmata: Ascidae), pp. 448-464 in Zootaxa 4952 (3) on page 451, DOI: 10.11646/zootaxa.4952.3.2, http://zenodo.org/record/4690531, {"references":["Lindquist, E. E. & Makarova, O. L. (2012) Review of the mite subfamily Arctoseiinae Evans with a key to its genera and description of a new genus and species from Siberia (Parasitiformes, Mesostigmata, Ascidae). ZooKeys, 233, 1 - 20. https: // doi. org / 10.3897 / zookeys. 233.3862"]}

Zerconopsini tribe n. Type genus: Zerconopsis Hull, 1918. Diagnosis. Adult dorsal shield entire, often fully covering body dorsum, with or without lateral incisions, its setae mostly simple but usually some (1–8 pairs) paddle-shaped, rarely setae j 1, z 1 and often seta J 5 serrated or pilose. Sternal or sternitigenital shield consolidated with endopodal sclerites between coxae I–II and II–III but usually not connected with free endopodal strips between coxae III–IV. Female with ventrianal shield bearing 1–6 pairs of opisthogastric setae but never expansive enough to include metapodal platelets and setae JV 1; rarely, broad anal shield present bearing only three circumanal setae (Iphidonopsis). Peritreme long, its margins often crenulate. Subcapitulum with deutosternum wide or of moderate width; hypostomatic setae hp -1 and usually hp -3 elongated, often whip-like, 2–5 times longer than distance between them. Palp tarsus without distinct macroseta but apically with two distinct sensillae—large erect one with truncated tip, and curved “S”-like sensilla (Fig. 10). Palp trochanter internal seta whip-like, often much (3–6 times) longer than external seta (as an exception, 1.5 times in one undescribed Zerconopsis species). Male spermatodactyl of different length, tube-like, sometimes with membranous frenulum. Tarsus I with or without claws. Genu III usually plesiomorphically with eight setae (pv -1 present); tarsi II–IV each with dorso-proximal setae ad -2, pd -2 elongated, curved; males of few Zerconopsis species have ventral seta mv on tarsus II modified, thickened and attenuated. Genera included: Zerconopsis, Xenoseius, Iphidonopsis.

Zerconopsini tribe n. Type genus: Zerconopsis Hull, 1918. Diagnosis. Adult dorsal shield entire, often fully covering body dorsum, with or without lateral incisions, its setae mostly simple but usually some (1���8 pairs) paddle-shaped, rarely setae j 1, z 1 and often seta J 5 serrated or pilose. Sternal or sternitigenital shield consolidated with endopodal sclerites between coxae I���II and II���III but usually not connected with free endopodal strips between coxae III���IV. Female with ventrianal shield bearing 1���6 pairs of opisthogastric setae but never expansive enough to include metapodal platelets and setae JV 1; rarely, broad anal shield present bearing only three circumanal setae (Iphidonopsis). Peritreme long, its margins often crenulate. Subcapitulum with deutosternum wide or of moderate width; hypostomatic setae hp -1 and usually hp -3 elongated, often whip-like, 2���5 times longer than distance between them. Palp tarsus without distinct macroseta but apically with two distinct sensillae���large erect one with truncated tip, and curved ���S���-like sensilla (Fig. 10). Palp trochanter internal seta whip-like, often much (3���6 times) longer than external seta (as an exception, 1.5 times in one undescribed Zerconopsis species). Male spermatodactyl of different length, tube-like, sometimes with membranous frenulum. Tarsus I with or without claws. Genu III usually plesiomorphically with eight setae (pv -1 present); tarsi II���IV each with dorso-proximal setae ad -2, pd -2 elongated, curved; males of few Zerconopsis species have ventral seta mv on tarsus II modified, thickened and attenuated. Genera included: Zerconopsis, Xenoseius, Iphidonopsis., Published as part of Makarova, Olga L., Marchenko, Irina I. & Lindquist, Evert E., 2021, Distribution, habitats, and redescription of the rare mite species Iphidonopsis sculptus Gwiazdowicz, 2004 (Mesostigmata: Ascidae), pp. 448-464 in Zootaxa 4952 (3) on page 450, DOI: 10.11646/zootaxa.4952.3.2, http://zenodo.org/record/4690531

Iphidonopsis sculptus Gwiazdowicz, 2004 (Figs 1���36) Iphidonopsis sculptus Gwiazdowicz, 2004: 155. Specimens examined. CANADA, ALBERTA PROVINCE: 2 females, 1 male, 2.5 mi. (4 km) SE Kananaskis Forest Experiment Station, ex bracket fungi on spruce log, 12.VIII.1970, leg. E. Lindquist; 2 females, Harvey Pass, Banff National Park, ex Dryocoetes affaber (Mann.) [Coleoptera: Curculionidae: Scolytinae], 12.VII.1968, leg. unspecified. RUSSIA, TVER��� REGION: 1 female, Central Forest State Reserve, spruce forest, ex Fomitopsis pinicola (Fungi: Fomitopsidaceae), 7.VIII.2019, leg. A.V. Alexandrova; KARELIA, PETROZAVODSK REGION: 16 females, 2 males, Kivach Nature Reserve, spruce forest, ex Fomitopsis pinicola, 26.IX.2020, leg. A. Zuyev; MOSCOW REGION: 1 female, Istra City vicinity, Novorakovo vil., ex Fomitopsis pinicola on dead Betula pendula (coexistant with Zerconopsis remiger (Kramer, 1876) and Z. michaeli Evans et Hyatt, 1960), 17.VI.2010, leg. O. Makarova; 1 female, Moscow city, Pokrovsko-Streshnevo Park, ex Fomitopsis pinicola on Alnus glutinosa log (coexistent with Zerconopsis remiger and Z. michaeli), 20.VII.2020, leg. O. Makarova; KOMI REPUBLIC: 4 females, 8 males, 3 deutonymphs, Pechoro-Ilychskii Nature Reserve, Yaksha vil. vicinity, mossy spruce forest, ex Trametes hirsuta (Fungi: Polyporaceae), 27.IX.2003, leg. O. Makarova; ALTAI: 1 female, Central Altai, Seminskii Pass, 1700 m a.s.l., 51��02��� N 85��36��� E, Pinus sibiricus ��� Picea obovata forest, in litter, 20.VII. 2001, leg. I.I. Marchenko; 1 female, West Altai, Ust-Koksa District, Krasnaya Mt., 1900 m a.s.l., 50��05��� N 85��11��� E, alpine meadow with Pinus sibiricus, in litter, 23.VII.2017, leg. I.I. Marchenko; WEST SAYAN: 2 females, 1 male, Khakassia, 40 km from Abaza, along river Korasibo, 700 m a.s.l., 52��35��� N 90��08��� E, Abies sibirica ��� Picea obovata ��� Pinus sibiricus taiga, in litter, 25.VI.2013, leg. I.I. Marchenko; KHABAROVSKII KRAI: 1 female, Bikin District, environs of Boytsovo vil., 120 m a.s.l., 46��58��� N 134��19��� E, mixed forest with Pinus koraiensis, in litter, 5.IX.1991, leg. I.I. Volonikhina (Marchenko); PRIMORSKII KRAI: 2 females, 2 males, Ussurijskii Nature Reserve, 43��33��� N 132��16��� E, mixed forest with Pinus koraiensis, in litter, 15.VII.1988, leg. L.G. Grishina. Redescription. Adults. Middle-sized mites with idiosoma rather pentagonal (especially pronounced in unfed mites), its anterior part triangular, lateral and posterior margins straight (Figs 1, 3). Mature adults dark-yellow or brownish in colour. Dorsal and anal/ventrianal shields strongly sclerotised and ornamented (Figs 1���4). Dorsal shield covering entire dorsal and often lateral idiosoma, without lateral incisions, with lateral margins notched in males and unfed females (Fig. 17) but almost smooth midlaterally in swollen females (Fig. 7). Anterior and lateral areas of dorsal shield with sharp comb-like ornamentation, medial area with six pairs of deep depression with angular margins (three smaller pairs of depression on podonotal region, three larger pairs on opisthonotal region) (Fig. 1). Most dorsal setae smooth, of moderate size or long, with hair-like tips (Fig. 7). Five pairs of dorsal gland pores well developed (gdj 2, gdj 4, gdz 6, gdZ 3, gdZ 4); dorsal lyrifissures hardly discernible, six pairs on opisthonotal region and one pair, Rp, on soft cuticle (Fig. 7). Female. Idiosomal dorsum (Figs 1, 7). Dorsal shield length 484���612 by 292���360 width, moderately broad, length/width 1.59���1.75, its maximal width at level of setae J 1 (Fig. 7). Podonotal region normally bearing 22 pairs of setae including all r -series setae (j 1���6, z 1���6, s 1���6, r 2���5). Opisthonotal region with 14 pairs of setae (S 2 always absent, R 1, R 2 on soft cuticle). Among dorsal setae, j 1 (9���13) and z 1 (8���11) shortest, awl-like, sometimes attenuated, inserted ventrally on ventrally bent anterior shield margin (Fig. 11); setae s 1, s 2 (both 16���20), r 2���4 (20���28) very thin; seta J 5 (24���34) ciliated basally; caudal setae S 5 and Z 5 (44���56), directed posteromedially; other setal lengths from 26���38 (z 2, j 3) to 48���70 (S 4, Z 4); marginal setae R 1 and R 2 24���34, inserted laterally or ventrally on soft cuticle. Idiosomal venter (Figs 2, 11, 12, 14, 15). Base of tritosternum slightly longer (20���24) than wide (13���16); laciniae 88���100 long, fused for nearly one third of entire length, with sparse long barbs along free part and short lateral spicules on fused area (Fig. 14). Presternal sclerotisation undeveloped.All ventral setae smooth, needle-like. Sternal shield 90���108 length by minimal width between coxae II 66���76, consolidated with endopodal platelets between coxae I���II (shield width at this level 128���144) and between coxae II���III (shield width at this level 112���134) (Fig. 11); anterior and posterior margins of shield weakly concave or straight; reticulation usually hardly visible, slightly lineate along lateral margins; shield with typical setae st 1���3 (17���20) and lyrifissures iv 1���3, vestigial gland pores gv 1 rarely visible on its posterior margin (Fig. 12). Setae st 4 (16���19) on soft cuticle. Endopodal strips between coxae III and IV free. Epigynal shield length 114���136 by 66���76 wide, faintly reticulate, with lateral margins slightly widened posteriorly; seta st 5 (20���22) and paragenital poroids iv 5 off shield; anterior hyaline flap short, not extending to posterior margin of sternal shield. Epigynal shield flanked by strong subcutaneous sclerites connected with pericoxal strips behind coxa IV. Four postgenital platelets vestigial, hidden in common transverse dermal fold; one more pair of bacilliform platelets formed between setae JV 1 and ZV 1. Two pairs of bacilliform metapodal platelets present, smaller anterior platelets 11���18 x 2���5, larger posterior platelets 38���42 x 8���9. Exopodal platelets between coxae I���II and II���III small, triangular or rhomboid, hardly visible in ventral aspect; their apices encompassing opening of cutaneous glands gvb 1 and gvb 2, respectively. Anal shield expansive, generally subtrapezoidal or broadly suboval, well sclerotised, with convex anterolateral margins variably smooth or crenulate (Figs 11, 15) and wide cribrum between poorly visible openings of gland pores gv 3, wider (164���168) than long (108���124), length/width 0.64���0.76, clearly pitted and slightly reticulate; paranal setae (19���22) inserted at level of posterior margin of anus, shorter than tubercle-based postanal seta (24���33), all circumanal setae needle-like; distance between anal opening and anterior shield margin 2���3 times smaller than distance between anal opening and posterior shield margin. Opisthogastric region with ten pairs of setae (JV 1���5, ZV 1���5), openings of gland pores gv 2 and four pairs of poroids; length of opisthogastric setae varying from 19���20 (ZV 1) to 30���42 (JV 5, Z V 5). Peritrematal shield rather wide, smooth, fused with ventrally bent dorsal shield margin anteriorly and exopodal platelet enveloping coxa IV posteriorly (Fig. 11); lyrifissures ip 1���3 and glands gp 1���2 present. Peritreme with crenulate margins, relatively wide (268���308 x 10���14), reaching seta j 1 anteriorly. Spermathecal apparatus not visible. Gnathosoma (Figs 8���10, 13, 16, 26). Gnathotectum narrow (15���18), triramous; middle process apically split (3���5 partite), slightly longer (18) than side laterally dentate processes (Fig. 9). Subcapitulum (Fig. 16) as long (82��� 85) as wide (79���84). Deutosternum of even, moderate width (14���15) (Fig. 16) or broadened posteriorly up to 19���20 (as on Fig. 27), with seven rows of numerous denticles (9���21 denticles in each row), clearly bordered. Hypostomatic pairs hp 1 (41���44) and hp 3 (34���36) sometimes undulate, much longer than hp 2 (16���19) and pc (19���20), all setae attenuate. Corniculi rather small, 32���35 x 9���11. Internal malae somewhat longer than corniculi, sharply narrowing, with lateral margins very shortly fimbriated proximally. Buccal cavity with fimbriated paralabra well developed (Fig. 26). Chelicera weak, its length without basal segment 143���159; basal segment length 54���66, with paraxial wall more elongated (as on Fig. 21b); cheliceral digits of moderate size, ca 1.5 longer than corniculus. Fixed digit of chela with one large blunt tooth in paraxial position and thin needle-like pilus dentilis in antiaxial position; distal part of fixed digit grooved, ending in apical hook and two small opposed denticles on sides of groove margins, with circular depression between them (Fig. 8); dorsal seta (16���17) lanceolate; lateral and dorsal lyrifissures narrow, hardly visible. Movable digit (37���40) bidentate, its arthrodial membrane smooth. Palp length 128���152; internal seta of trochanter (40���48) resembling hp 1 in form and length but swollen basally, much longer than external seta (14���16); palp with three typically specialised thick setae on femur and genu; femoral al and genual al -1 12���17 long, conical apically; genual al -2 much longer (20���23), flattened apically, with blunt oblique tip (Fig. 13); palp tarsus with two distinctive setae���one thick truncate (20���22) and one sinuous (S-like) (Fig. 10). Legs (Figs 28, 29, 33, 34). Legs of moderate length (I 304���396, II 272���384, III 260���344, IV 332���468); leg I clearly shorter than dorsal shield. Length of tarsus I 64���94, tarsus IV 100���150. Leg chaetome as described for genus Zerconopsis (Lindquist and Evans, 1965); setation of legs I, II, III, IV, genua: 12���10���8���7; tibiae: 12���9���7���7; chaetotactic formulae for genua I���II���III���IV: (2���3/2, 2/1���2) (2���3/1, 2/1���1) (1���2/1, 2/1���1) (1���2/1, 2/0���1); tibiae: (2���3/2, 2/1���2) (2���2/1, 2/1���1) (1���1/1, 2/1���1) (1���1/1, 2/1���1); seta av -2 on genu I and tibia I present; femur I with seta pl -2 moved ventrally (as on Fig. 24). Setae of legs smooth, spiniform or hair-like distally, generally of moderate length or short, only tarsus I distally with four long thin setae and tarsi II���IV each with dorso-proximal setae ad -2, pd -2 curved, almost as long as interval from their insertion to apex of tarsus (Figs 33, 34); most dorsal and several lateral and ventral setae on femora, genua and tibiae of all legs inserted on tubercles (as on Figs 24, 25, 28, 29); distal verticil setae on tarsi II���IV rather small, almost evenly distant from apex; av -1, pv -1 (11���15) slightly shorter than al -1, pl -1 (13���18) (as on Fig. 36). Ambulacrum I on short pedicel, claws I (8���10) twice smaller than claws II���IV (14���17). Ambulacra of legs II���IV (length 24���32) with short thin paradactyli (9���14) hardly extending beyond apices of claws (as on Fig. 36). Tarsi II���IV with apical setae ad -1 and pd -1 small (10���15), about half as long as pretarsus to base of claws. Tarsus I with 7 rod-like solenidia (5 of them inserted apically). Male. Idiosomal dorsum (Fig. 3). Dorsal shield length 398���456 by 234���280 width, length/width 1.61���1.82. Chaetotaxy and ornamentation of dorsal shield similar to that of female but setae j 2���6, z 2, z 5, J 1���5 (18���28) much shorter than setae in series Z and S (40���50). Seta j 1 awl-like, short (8���10); z 1 very thin (6���11); length of s 1, s 2 14���15, r 2���4 16���20; posterior marginal series only with one seta, R 1 (18���20). Idiosomal venter (Figs 4, 17, 18, 22, 23). Tritosternum slightly smaller than in female: base 14���18 long, 11���13 wide; laciniae 68���76 long, distally with long barbs. Sternitigenital shield united with endopodal platelets developed between coxae I���II and II���III (Figs 17, 23); its anterior margin medially concave, tightly abutting genital valve, posterior margin convex; shield length 160���174, width before coxae II 114���120, width at midlevel of coxae II 60���66, width at level of middle angles 108���118, width at midlevel of coxae IV 44���54; glands gvb 1 and gvb 2 associated with triangular exopodal platelets between coxae I���II and II���III, respectively (Fig. 23). Endocoxal angular platelets between coxae III���IV free. Sternitigenital shield with vestigial reticulate ornamentation, three pairs of lyrifissures iv 1���3, and setae st 1���3, st 5 (17���19) usually slightly longer than seta st 4 (13���17). Genital valve transverse oval (11��� 13 x 24���32), inner funnel weakly narrowing (Fig. 18). Leg acetabula II and III with lateral margins strengthened by row of small exopodal sclerites (Fig. 17). Poroids iv 5 inserted on soft cuticle between sternitigenital and ventrianal shields. Ventrianal shield 144���160 long and 170���211 wide at level of setae ZV 1, length/width 0.75���0.91, clearly reticulated anteriorly and coarsely punctate behind setae JV 2, subtrapezoidal in form (Fig. 17), with anterior margin generally convex, undulate and lateral margins often bent dorsally, consolidated with metapodal platelets (Fig. 22). Ventrianal shield with 7���8 pairs of opisthogastric setae (JV 1���5, ZV 1���3; ZV 1 and nearest lyrifissure rarely on soft cuticle), three circumanal setae, two pairs of poroids and well developed openings of glands gv 3. Cribrum narrow (7���10), usually bent dorsally. Setae ZV 4 on soft cuticle.Among opisthogastric setae, ZV 1 shortest (11���17), JV 5, ZV 4 longest (up to 24���31); circumanal setae 16���20. Peritrematal shield anteriorly reduced, peritreme 248���268 x 8���11. Gnathosoma (Figs 5, 19���21, 26a, 27, 30). Form and size of gnathotectum (width 13���17, middle process length 16���20) similar to female; subcapitulum (70���72 x 68���76) and corniculus (23���25 x 9���10) smaller, hypostomatic seta hp 2 (14���16) and subcapitular seta pc (15���19) shorter than in female (Fig. 30), length of hypostomatic seta hp 1 40���48, hp 3 35���39; internal malae in contrast to female distinctly ciliated; salivary stylus apically narrowing, 25 long (Fig. 26a). Deutosternum often widening posteriorly from 10���12 to 18���20 (Figs 27, 30). Chelicera length without basal segment 116���128; basal segment length 52���62; fixed digit of chela with one large tooth and small denticle in paraxial position and thin needle-like pilus dentilis in antiaxial position; distal part of fixed digit grooved (Fig. 21a); movable digit 32���38 long, with one tooth and long tube-like spermatodactyl protruding far beyond tip of digit (length of free part 50���58) and directed downward (Figs 5, 21, 30). Palp (112���118 long) shorter than in female; trochanter internal seta 36���44 in length, trochanter external seta 11���13, genual al -1 and femoral al 12���13, genual al -2 18���21, tarsal apical macroseta 19���21. Legs (Figs 24, 25, 36). Length of legs I���IV: 276���292 (much shorter than idiosoma), 240���268, 236���256, and 298���324, respectively; length of tarsus I 64���75, tarsus IV 96���158. Chaetotaxy as in female including ventral position of seta pl -2 on femur I (Fig. 24). Legs without dimorphically modified setae. All ambulacral structures as in female but smaller (ambulacra II���IV 21���24, their paradactyli 8���11, claws 11���14; claw I 8���10). On tarsi II���IV, length of apical setae ad -1 and pd -1 10���12; among subapical setae, av -1, pv -1 (9���11) slightly shorter than al -1, pl -1 (13���14) (Fig. 36). New description. Deutonymph. Idiosomal dorsum (Figs 6, 31). Dorsal shield 392���408 x 228���332, with lateral incisions behind setae s 6, ornamentation similar to that of female (Figs 6, 31), with prominent openings of glands gdz 6, gdZ 3, gdZ 4; podonotal region with 18 pairs of setae, opisthonotal region with 14 pairs of setae; setae j 1 (9���11) and z 1 (6���9) shortest, setae j 5, j 6, z 5, z 6, J 1���5 14���22 long, other setae longer, up to 42���48 (Z 3���5, S 3���5). Setae s 1, s 2, r 2���5 (14���19) and R 1, R 2 (17���20) inserted on soft cuticle. Idiosomal venter (Figs 37, 38). Tritosternum base 18���20 x 11���12; laciniae 74 long, with sparse long barbs in free part and shorter barbs (not spicules as in adults) in fused area (Fig. 37). Sternal shield narrow, with acuminate posterior margin, weakly sclerotised (Fig. 38), 188���192 long, its width at midlevel of coxae II 44, width at level of middle angles 88���100; shield bearing setae st 1���3 (18���19), st 4 (13���14) and lyrifissures iv 1���3. Setae st 5 (17), paragenital poroids iv 5 and sometimes asymmetrically poroids iv 3 on soft cuticle. Exopodal platelets not developed, openings of glands gvb 1 on soft cuticle, gvb 2 not visible. Anal shield transversely oval (60���64 x 114���118), well formed, coarsely punctate, with narrow (8) cribrum and circumanal setae 17���22 long. Opisthogastric region with 10 pairs of setae (JV 1���5, ZV 1���5) and three pairs of lyrifissures; seta ZV 1 (10���15) shortest, JV 5 (22���25) longest; metapodal platelets narrow (21���29 x 6���7); a pair of transversal bacilliform platelets developed between setae JV 1 and ZV 1 (Fig. 38). Peritrematal shield reduced, peritreme long and narrow (232���264 x 6), almost extending to seta z 1, with smooth margins (Fig. 31). Gnathosoma (Figs 32, 35). Gnathosomal structures in general as in adults. Tectum 14���16 wide with middle process 14���17 long (Fig. 32). Subcapitulum 70���76 x 64���66; deutosternum widening posteriorly from 12 to 16; hp 1 36���40, hp 2 14���17, hp 3 30���36, pc 17���20; corniculus 20���22 x 10���11; chelicera without basal segment 120���136 long, similar to female (Fig. 35); basal segment len, Published as part of Makarova, Olga L., Marchenko, Irina I. & Lindquist, Evert E., 2021, Distribution, habitats, and redescription of the rare mite species Iphidonopsis sculptus Gwiazdowicz, 2004 (Mesostigmata: Ascidae), pp. 448-464 in Zootaxa 4952 (3) on pages 452-462, DOI: 10.11646/zootaxa.4952.3.2, http://zenodo.org/record/4690531, {"references":["Gwiazdowicz, D. J. (2004) Iphidonopsis sculptus gen. n. sp. n., a new mite genus of the family Ascidae (Acari, Gamasida) from Poland. Biologia, Bratislava, 59, 153 - 158.","Lindquist, E. E. & Evans, G. O. (1965) Taxonomic concepts in the Ascidae, with a modified setal nomenclature for the idiosoma of the Gamasina (Acarina: Mesostigmata). Memoirs of the Entomological Society of Canada, 47, 1 - 64. https: // doi. org / 10.4039 / entm 9747 fv"]}

Maxiniini tribe n. Type genus: Maxinia Lindquist & Makarova, 2012. Diagnosis. Adult dorsal shield entire, robust, broad, without lateral incisions. Idiosomal setae simple. Female with sternal shield consolidated with endopodal sclerites between coxae I–II, and with free endopodal fragments between coxae II–III and free strongly developed strips between coxae III–IV; male sternitigenital shield fused with all endopodal sclerites beside coxae II–IV. Female with an apomorphically expansive ventrianal shield encompassing metapodal platelets anterolaterally and including setae JV 1 anteriorly. Male with sternitigenital shield abutting an expansive ventrianal shield that incorporates metapodal platelets. Peritreme slightly shortened, its margins smooth. Subcapitulum with narrow deutosternal groove; hypostomatic setae hp -1 and hp -3 not elongated and whip-like. Palp trochanter setae similar in size. Genu III with only seven setae (lacking pv -1). Male spermatodactyl short, tube-like. Tarsus I with claws. None of setae on tarsi II to IV conspicuously differentiated by shape or size. Genera included: Maxinia.

Key to species of the genus Iphidonopsis Gwiazdowicz, 2004 1. Gnathotectum with middle process simple, acuminate; female anal shield as long as wide, with anal opening evenly distant from anterior and posterior shield margins; metapodal platelets oval; four postgenital sclerites well formed; setae ZV 1 absent on opisthogaster. Female 494 ��m. China.......................................... magnanalis (Ma & Yin, 1999) - Gnathotectum with middle process split, 3���5 partite, apically; female anal shield wide, width/length ratio 1.41���1.56; distance between anal opening and anterior shield margin 2���3 times smaller than distance between anal opening and posterior shield margin; metapodal platelets bacilliform or narrowly oval; four postgenital sclerites hardly discernible; ZV 1 present, laterad JV 1 on opisthogaster. Female 484���612 ��m. Holarctic..................................... sculptus Gwiazdowicz, 2004, Published as part of Makarova, Olga L., Marchenko, Irina I. & Lindquist, Evert E., 2021, Distribution, habitats, and redescription of the rare mite species Iphidonopsis sculptus Gwiazdowicz, 2004 (Mesostigmata: Ascidae), pp. 448-464 in Zootaxa 4952 (3) on page 463, DOI: 10.11646/zootaxa.4952.3.2, http://zenodo.org/record/4690531, {"references":["Gwiazdowicz, D. J. (2004) Iphidonopsis sculptus gen. n. sp. n., a new mite genus of the family Ascidae (Acari, Gamasida) from Poland. Biologia, Bratislava, 59, 153 - 158.","Ma, L. & Yin, X. (1999) Four new species and two new record genera of the family Aceosejidae from China (Acari: Gamasina). Acta Arachnologica Sinica, 8 (1), 1 - 7. [in Chinese]"]}

Iphidonopsis -Zerconopsis relations Members of Iphidonopsis much resemble those of Zerconopsis species but in contrast to the latter lack in particular paddle-formed setae on the dorsal shield and females have an anal (not ventrianal) shield, bearing only the three circumanal setae. Based on out-group comparisons with other genera of Arctoseiinae, both of these characteristics may be considered as initial, plesiomorphic conditions. Nevertheless, the genus Iphidonopsis seems to be an advanced derivate of Zerconopsis (probably of the mycophilous michaeli -group) in secondarily losing the paddleshaped form of certain dorsal setae as well as reducing the size of the ventrianal shield, leaving opisthogastric setae on soft cuticle. Both of these tendencies are already found among species of Zerconopsis. Thus, Z. apodius Karg, 1969 has only one pair of paddle-like dorsal setae (Z 5) ���remaining��� from eight pairs that are possible among other genus members (eight pairs of such modified dorsal setae were revealed in a new undescribed Zerconopsis species from North America and Europe). Moreover, another new species of Zerconopsis from California has only one pair of opisthogastric setae on the ventrianal shield while other members bear two to six pairs of setae in addition to the circumanal setae. Similar to species of the michaeli -group of Zerconopsis, adults of Iphidonopsis have a rigid and expansive dorsal shield with extremely complex ornamental structure, relatively reduced and anteriorly rounded female ventrianal/ anal shield, and legs I���II reinforced by tuberculate setal sockets. As advanced features of Iphidonopsis within Zerconopsis, we consider the simplification of paddle-shaped dorsal setae, diminution of the ventrianal shield to a broad anal shield in female, and presence of an apomorphically elongated spermatodactyl in the males. Both Iphidonopsis and michaeli -group members of Zerconopsis inhabit bracket fungi, sometimes together., Published as part of Makarova, Olga L., Marchenko, Irina I. & Lindquist, Evert E., 2021, Distribution, habitats, and redescription of the rare mite species Iphidonopsis sculptus Gwiazdowicz, 2004 (Mesostigmata: Ascidae), pp. 448-464 in Zootaxa 4952 (3) on page 462, DOI: 10.11646/zootaxa.4952.3.2, http://zenodo.org/record/4690531, {"references":["Karg, W. (1969) Untersuchungen zur Kenntnis der Ascaoidea Karg, 1965 (Acarina: Parasitiformes) mit der Beschreibung von acht neuen Arten. Zoologischer Anzeiger, 182, 393 - 406."]}

Subfamily Arctoseiinae Evans, 1963 We propose a subdivision of the subfamily Arctoseiinae Evans, 1963 into three tribes, Arctoseiini Evans, 1963 (numbering about 60 species), Zerconopsini tribe n. (about 25 species), and monobasic Maxiniini tribe n. The strongly evident relationship of Iphidonopsis, Zerconopsis, and Xenoseius is based on apomorphic attributes (dorsoproximal setae ad -2 and pd -2 of tarsi II to IV elongated, curved, somewhat whip-like; hypostomal setae hp -1 and to some extent hp -3, as well as the internal palp trochanter seta elongated, whip-like) and on a plesiomorphic attribute (genu III with eight setae, including pv -1). Besides, females of these genera possess an often expansive ventrianal shield usually bearing one to six pairs of opisthogastric setae in addition to the circumanal setae (female Iphidonopsis exceptional, having a large anal shield bearing only the circumanal setae) along with one or two pairs of free metapodal platelets. All these characteristics argue for the deriving of a tribe Zerconopsini tribe n. Segregation of the tribe Arctoseiini Evans, 1963 (including Arctoseius and Iphidozercon) is supported by such apomorphic attributes as reduced ventral sclerotisation (epigynal shield slender, anal shield bearing only three circumanal setae, sternal shield sometimes separated from endocoxal sclerites) and genu III with seven setae, lacking pv -1. Recognition of the monobasic Maxiniini tribe n. is strongly based apomorphically on a narrowed deutosternum, a highly expanded ventrianal shield incorporating metapodal platelets and bearing setae JV 1, along with the absence of setae ZV 1 in both sexes. Although the single known species of Maxiniini seems to be free-living in soil litter throughout all its broad range (Lindquist & Makarova, 2012), numerous members of the two other tribes display some clear tendencies in their ecological specialisation associated phoretically with winged dipterans, namely an inclination to fungal substrata in Zerconopsini and to different wet habitats enriched by nematoceran juveniles (eggs, early stage larvae) in Arctoseiini., Published as part of Makarova, Olga L., Marchenko, Irina I. & Lindquist, Evert E., 2021, Distribution, habitats, and redescription of the rare mite species Iphidonopsis sculptus Gwiazdowicz, 2004 (Mesostigmata: Ascidae), pp. 448-464 in Zootaxa 4952 (3) on pages 449-450, DOI: 10.11646/zootaxa.4952.3.2, http://zenodo.org/record/4690531, {"references":["Evans, G. O. (1963) Observations on the chaetotaxy of the legs in the free-living Gamasina. Bulletin of the British Museum (Natural History), Zoology, 10, 275 - 303. https: // doi. org / 10.5962 / bhl. part. 20528","Lindquist, E. E. & Makarova, O. L. (2012) Review of the mite subfamily Arctoseiinae Evans with a key to its genera and description of a new genus and species from Siberia (Parasitiformes, Mesostigmata, Ascidae). ZooKeys, 233, 1 - 20. https: // doi. org / 10.3897 / zookeys. 233.3862"]}

Arctoseiini Evans, 1963 Type genus: Arctoseius Thor, 1930. Diagnosis. Adult dorsal shield entire, often parallel-sided, with or without lateral incisions. Idiosomal setae simple. Sternal or sternitigenital shield consolidated or not with endopodal sclerites between coxae I���II and II���III, and usually with free endopodal strips between coxae III���IV. Females with free metapodal platelets and anal shield bearing only three circumanal setae (as an exception, anal shield of Arctoseius magnanalis Evans, 1958 with one pair of opisthogastric setae). Male with ventrianal shield incorporating metapodal platelets or not, rarely with ventral and anal shields. Peritreme of varying length, sometimes long and sharply curved anteriorly, its margins smooth. Subcapitulum with deutosternum wide or of moderate width, with setae weakly differing in length. Palp trochanter setae similar in size. Palp tarsus sometimes with distinct apical macroseta. Genu III with only seven setae (lacking pv -1). Male spermatodactyl short or of middle length; tube-like, rarely undulate or spoon-like. Tarsus I with claws. None of setae on tarsi II to IV conspicuously differentiated by thinness, elongation, or curved shape (as an exception, males of several Arctoseius species have some ventral setae modified, thickened on tarsus and femur II). Genera included: Arctoseius, Iphidozercon., Published as part of Makarova, Olga L., Marchenko, Irina I. & Lindquist, Evert E., 2021, Distribution, habitats, and redescription of the rare mite species Iphidonopsis sculptus Gwiazdowicz, 2004 (Mesostigmata: Ascidae), pp. 448-464 in Zootaxa 4952 (3) on page 450, DOI: 10.11646/zootaxa.4952.3.2, http://zenodo.org/record/4690531, {"references":["Evans, G. O. (1963) Observations on the chaetotaxy of the legs in the free-living Gamasina. Bulletin of the British Museum (Natural History), Zoology, 10, 275 - 303. https: // doi. org / 10.5962 / bhl. part. 20528"]}

Review of Iphiopsididae as a definable family-group The genus Berlesia was first placed in the controversial family Iphiopsidinae (now Iphiopsididae) by Vitzthum (1943) where it has remained. This placement is unsatisfactory for several reasons. Firstly the Iphiopsididae itself needs examination. Kramer (1886) mentioned the new family as Iphiopsidae (sic), just once, after stating that the available gamasid families were not sufficient to encompass known species, and thereby granting family status to Iphiopsis mirabilis (Berlese, 1882). Kramer (1886) did not specifically include any other genera in his Iphiopsidae, and his proposal of a family for Iphiopsis was overlooked until noted by Vitzthum (1943). Berlese (1892, p.16) provided a diagram of relationships between genera (Genera Mesostigmatum Secundum Ordinem Naturalem Disposita) which clearly showed that Berlesia (along with the ant-associated Neoberlesia Berlese, 1892) was considered to be distinctly separated from Iphiopsis, Iphis and genera considered similar at that time. The associated key (p. 37) did not diagnose Berlesia, as the key was explictly restricted to European taxa. However the diagnostic information in that key, separating Iphiopsis based on loss of peritreme, re-appears later where it is then used in a much broader geographic context (Vitzthum 1943), and thereafter for many more genera (Evans 1955). Oudemans (1904) was struck by the similarity of female Berlesia chelicerae to those of Varroa Oudemans, 1904, and their modifications for parasitism (although which cheliceral digit regresses was noted as dissimilar); however, he rejected a close relationship between these genera because of differences in ventral shields. Vitzthum (1943) modified Kramer’s (1886) monobasic Iphiopsididae, adding Jacobsonia Berlese, 1910b and Berlesia to form a subfamily Iphiopsidinae within Laelaptidae (sic). The only significant character given in sup- port of this subfamilial grouping was the almost absent peritreme (along with a weak attribute of the idiosoma being colourless or brownish). However, this criterion was sometimes clearly subordinated to others, since the genus Dinogamasus Kramer, 1898, which lack peritremes, was placed in the newly-created Hypoaspidinae, while Neolaelaps Hirst, 1926, with similarly vestigial peritremes, was placed in the Laelaptinae. Keegan (1950) added Dynatochela, a beetle-associate, to the Iphiopsidinae. Its reduced peritreme was the sole basis for subfamily placement, despite other notable discordant features of Dynatochela (including well-developed claws on all tarsi). Clearly, the concept for any supraspecific grouping must include the type species. Evans (1955) invoked this point at the genus level, noting that the most important characters of Iphiopsis were the reduction of the peritreme and the absence of claws on leg I (the species original description by Berlese, 1882, indeed states that the first pair of legs are destitute of claws, and his accompanying figure 1, plate 64, clearly shows that leg I lacks claws albeit provided with a well-developed ambulacrum). Potential implications at the subfamily level were avoided by Evans (1955) noting that claws of one or more pairs of legs are usually reduced in Iphiopsidinae (our italics). This qualification allowed Evans to retain Berlesia and Dynatochela, and include Dinogamasus in Iphiopsidinae. However, by considering primarily myriapod hosts, Evans’ discussion did not account for the growing heterogeneity in this taxon. Also, iphiopsidines were stated to be “paraphages of arthropods”, thus overlooking statements by previous authors (Oudemans 1904, Vitzthum 1943) that Berlesia is parasitic. Evans (1955) makes a strong assertion that Iphiopsidinae “with little doubt arose from” the Hypoaspis -group—if so, Hypoaspidinae is rendered paraphyletic, and this creates another question as to the validity of Iphiopsidinae. The Iphiopsidinae was again defined according to a reduced peritreme for the placement of Dyscinetonyssus hystricosus Moss & Funk, 1965; however, this was qualified with acknowledgement that some authorities considered the characters used to define Iphiopsidinae as convergent (Moss & Funk 1965). In the first published cladistic morphological analysis of the Laelapidae, Casanueva (1993) recovered a phylogenetic estimate where all sampled myriapod associates clade together. Thus, the earlier concept of the Iphiopsidinae was recovered in part, but became substantially modified by including Julolaelaps Berlese, 1916 and Scissuralaelaps Womersley, 1945. The former iphiopsidine Dyscinetonyssus was recovered in a separate clade that included all beetle and cockroach associates, while all hymenopteran associates including Neoberlesia and Dinogamasus were recovered in a greatly expanded Melittiphidinae. Berlesia was not analysed. These are interesting results, which might be intuitively appealing if one believes dermanyssines show a largely inflexible pattern of cophylogeny with their hosts at most levels ranging from species to family. However, the combination of character and taxon sampling does not permit full confidence in the results. For instance, while the loss of hypostomatic seta hp3 has relevance to some myriapod associates, it was used to define a clade with Julolaelaps Berlese, 1916 subtending the myriapod-associated Iphiopsidinae. The sampling entailed a strong assumption that the exclusion of Julolaelaps which retain hp3 (e.g., J. moseri Hunter & Rosario, 1986) would not substantially alter the results, including grouping with the similar Hypoaspis as previously hypothesized (Evans 1955). This result and similar ones appear to be reinforced by sampling taxa in a way that creates superficially distinct characters. For instance, the defining synapomorphy reconstructed for most of the Hypoaspidini is loss of podonotal z3 which is a poor, mostly unrepresentative attribute to characterize many taxa intended for this clade including Hypoaspis sensu stricto. Podonotal z3 is an ontogenetically weak deutonymphal seta whose suppression is subject to much homoplasy (Lindquist & Evans 1965), so it would be surprising if its loss could define any sizeable clade. Also the five putative synapomorphies that define a clade equivalent to Iphiopsidinae (i.e. myriapod associates other than Julolaelaps and Scissuralaelaps) plus an additional four other putative synapomorphies that largely define a yet more inclusive clade, are all regressive leg characters that are also found in hypotrichous melittiphines, notably Myrmozercon brevipes Berlese, 1902 (with this genus actually not included in the analysis). Note that the leg chaetotaxy of Berlesia species is relatively holotrichous compared to most of the myriapod-associated Iphiopsididae sampled by Casanueva (1993) and bears only two of the nine reductive characters reconstructed by Casaneuva (1993; Fig 8, Stem 3) as putative synapomorphies for the relevant group containing the type species (Group VII Iphiopsini). However, an alleged sister group to the main iphiopsidid clade proposed by Casanueva (1993) is relevant to consider with respect to Berlesia as it contains potential relatives but also reveals possible pitfalls in uncovering these with morphological analysis. In this clade the mygalomorph-associated genus Ljunghia Oudemans, 1932 and the crab-associated genus Cyclothorax von Frauenfeld, 1868 are recovered together. Casaneuva (1993) noted that the synapomorphies linking Ljunghia to iphiopsidids were weak with low confidence in this expansion proposed for the Iphiopsididae. Ljunghia has similarities with Berlesia which may or may not indicate relationship. Deserving attention amongst these are: the usually pos- teriorly-positioned paranal setae, the shared absence of gv3 on the anal shield, and the stout movable digit. For the latter, however, it is important to note that the movable digit of Berlesia appears rotated laterally, with its teeth flared away from the fixed digit, whose base is swollen (somewhat as in Ixodida). This contrasts with Ljunghia where the teeth on the movable digit still oppose what remains of the fixed digit, and the fixed digit base is not swollen. Thus Ljunghia movable digits lack some special features found in Berlesia that would make it more difficult to argue for similarity due to relationship under a structural criterion (Riedl 1978, p 34). The above highlights the rampant homoplasy due to convergence and parallelism that plagues all potential candidates for Iphiopsididae. When both Moraza & Kazemi (2012) and Seeman & Alberti (2015) tried to apply characters proposed by Casaneuva (1993) to define/diagnose Iphiopsididae, they found that these characters had widespread occurrences elsewhere in the Laelapidae, such that both sets of authors rejected the family-ranking of Iphiopsididae. Nemati et al. (2015) also found the inclusion of Julolaelaps to be problematic. Characters such as hypotrichy, reduced peritremes and reduced claws found in the “ Iphiopsididae ”, are known to be correlated with a wide variety of intimate symbioses in various Dermanyssoidea and are regressive in nature (as realized 65 years ago by Evans 1955). Iphiopsididae, at whatever rank, will remain an unstable concept, unless future molecular analyses, backed by unexpected morphological attributes indicate otherwise. Currently, it can only be considered satisfactory for taxa that are highly similar morphologically to the type genus Iphiopsis. Better knowledge of the type species would be useful here (Halliday & Juvara-Bals 2016), insofar as clarifying the core taxa related to Iphiopsis. Other piecemeal renovations to Iphiopsididae will not help, unless they include new diagnostic characters that reliably link included iphiopsidid taxa. However, new suprageneric groupings of taxa that are clearly not related to Iphiopsis, when these are supported by convincing synapomorphies, will advance overall systematic perspectives partly because they will reduce the number of taxa to be considered for placement in Iphiopsididae., Published as part of Lindquist, Evert E., Oconnor, Barry M., Shaw, Matthew D. & Sidorchuk, Ekaterina A., 2020, Review of the genera Berlesia Canestrini, 1884, and Katydiseius Fain & Lukoschus 1983, the subfamily Katydiseiinae Fain & Lukoschus, 1983, and their family group relationships (Acari: Mesostigmata: Gamasina), with description of three new species parasitic on gryllacridid crickets (Orthoptera), pp. 5-70 in Zootaxa 4857 (1) on pages 49-51, DOI: 10.11646/zootaxa.4857.1.4, http://zenodo.org/record/4396474, {"references":["Vitzthum, H. (1943) Acarina. Bronn's Klassen und Ordnungen des Tierreichs, Bd. 5, Abt. 4, Buch 5, Lief 5, 641 - 800.","Kramer, P. (1886) Ueber Milben. Archiv fur Naturgeschichte, 52, 241 - 268, pl. XII.","Berlese, A. (1882) Acari, Myriopoda et Scorpiones hucusque in Italia reperta. Ordo Mesostigmata (Gamasidae). Fasc. 1. Typis Seminarii, Padua, 190 pp., XV tabs.","Berlese, A. (1892) Acari, Myriapoda, et Scorpiones hucusque in Italia reperta. Ordo Mesostigmata (Gamasidae). Typis Seminarii, Padua, 143 pp., 11 pls.","Evans, G. O. (1955) A review of the laelaptid paraphages of the Myriapoda with descriptions of three new species (Acarina: Laelaptidae). Parasitology, 45, 352 - 368. https: // doi. org / 10.1017 / S 0031182000027694","Oudemans, A. C. (1904) Note VIII. On a new genus and species of parasitic Acari. Notes from the Leyden Museum, 24 (4), 216 - 222.","Berlese, A. (1910 b) Brevi diagnosi di generi e specie nuovi di Acari. Redia, 6, 346 - 388.","Kramer, P. (1898) Gamasiden aus Deutsch Ostafrika. Zoologischer Anzeiger, 21, 416 - 418.","Hirst, S. (1926) Descriptions of new mites including four new species of \" red spider \". Proceedings of the Zoological Society of London, 96, 825 - 841.","Keegan, H. L. (1950) Dynatochela primus, n. sp. n. gen. (Acarina: Laelaptidae: Iphiosinae). Journal of Parasitology, 36, 511 - 514.","Moss, W. W. & Funk, F. C. (1965) Studies on the developmental chaetotaxy of Dyscinetonyssus hystricosus, n. g., n. sp. (Acari: Mesostigmata: Laelaptoidea). Acarologia, 7, 235 - 267.","Casanueva, M. E. (1993) Phylogenetic studies of the free-living and arthropod-associated Laelapidae (Acari: Mesostigmata). Gayana Zoologia, 57, 21 - 46.","Berlese, A. (1916) Centuria prima di Acari nuovi. Redia, 12, 19 - 67.","Womersley, H. (1945) An interesting and primitive new genus of Laelaptidae (Acarina) from Australia and New Guinea. Records of the South Australian Museum, 8, 226 - 228.","Lindquist, E. E. & Evans, G. O. (1965) Taxonomic concepts in the Ascidae, with a modified setal nomenclature for the idiosoma of the Gamasina (Acarina: Mesostigmata). Memoirs of the Entomological Society of Canada, 47, 1 - 65.","Berlese, A. (1902) Specie di Acari nuovi. Zoologischer Anzeiger, 25, 697 - 700.","Oudemans, A. C. (1932) Opus 550. Jubileum nummer voon Dr. J. Th. Oudemans 70 Jaar. Tijdschrift voor Entomologie, 75 (Supplement), 202 - 210.","von Frauenfeld, G. R. (1868) Zoologische Miscellen XV. Verhandlungen der zoologisch-botanischen Gesellschaft in Wien, 18, 885 - 902.","Riedl, R. (1978) Order in living organisms. A systems analysis of evolution (Translation of Riedl 1975 by R. P. D. Jeffries). Wiley Interscience Publication, New York, 313 pp.","Moraza, M. L. & Kazemi, S. (2012) Description of a new millipede-associated species (Acari: Mesostigmata: Laelapidae) from Iran and a key to species of Julolaelaps Berlese. International Journal of Acarology, 38, 6 - 17. https: // doi. org / 10.1080 / 01647954.2011.583273","Seeman, O. D. & Alberti, G. (2015) A new species of Scissuralaelaps (Acari: Mesostigmata: Laelapidae) from millipedes in the Philippines. Systematic and Applied Acarology, 20, 707 - 720. https: // doi. org / 10.11158 / saa. 20.6.12","Nemati, A., Riahi, E. & Gwiazdowicz, D. J. (2015) Description of a new species of Julolaelaps (Acari, Mesostigmata, Laelapidae) from Iran. ZooKeys, 526, 105 - 116. https: // doi. org / 10.3897 / zookeys. 526.5946","Halliday, R. B. & Juvara-Bals, I. (2016) Systematics and biology of the mite genus Ljunghia Oudemans in Southeast Asia (Acari: Laelapidae). Systematic & Applied Acarology, 21, 830 - 864. https: // doi. org / 10.11158 / saa. 21.6.10"]}

Berlesia hospitabilis new species (Figures 1–6, 18, 19, 21) Berlesia, undescribed species, host of Podapolipus gryllacridi Lindquist & Sidorchuk, 2019. Diagnosis Adult: dorsal shield faintly reticulated posteriorly, smooth elsewhere, with irregular indication of lateral incisions at level of setae s6; setae j1, mid-dorsals (j4–j6, z5, J1–J3) and J4 very short (e.g., j4 about one-fourth as long as interval between j4 and j5); dorsal shield setae j3 and z4 moderately long, but shorter than diagonal interval between them, and shorter than transverse interval between j3; setae z1 and J2 absent. Dorsolateral margins of idiosoma with setae of r-R and S series sparse, not hypertrichous. Soft ventral cuticle with 7 and 6 pairs of opisthogastric setae on female and male, respectively. Leg I genu and tibia each with 12 setae, pd-3 absent, av-2 present. Tarsus IV with seta pv-1 asymmetrically shorter but similar in attenuated form to av-1. Peritremes simple, not elaborated with internal cellular structure. Description Adult: Idiosomatic dorsum. Adult female (Fig. 1A). Idiosoma 550–730 long, 288–377 wide between insertions of setae r3; dorsal shield 507–550 long, greatest width 250–287 at level between insertions of s4 (n=4), faintly ornamented with subdermal reticula outlined by sparse puncta (mostly not evident in micrographs of surface structure); shield with irregular indication of lateral incisions at level of setae s6. Dorsal shield with usually 10 or 11 pairs (7 podonotal, 3 or 4 opisthonotal) of smooth setae of which s4 or Z5 asymmetrically on/off shield margin and J4 present or absent; setae j3 (58–75), z4 (67–85), s4 (70–85), Z5 (70–75) of similarly long lengths; other shield setae (j4–j6, z5, J1, J3, J4) tiny (5–12), occasionally asymmetrically rudimentary. Soft dorsal peripheral cuticle with tiny setae j1 (10–15) anteriorly, z2 (40–50), s6 (77–83) and r2, r3, r5 (50–70) anterolaterally, and Z2 (70–83), Z4 (80–85), S2 (55–67), S3 (62–68), S4 (72–83), S5 (75–90) and R1 (48–62) posterolaterally. Dorsal shield with complement of 15 pairs of discernible pore-like structures (5 podonotal, 10 opisthonotal), of which five pairs (two podonotal, three opisthonotal) superficially appear secretory (gland pores) and 10 pairs (three podonotal, seven opisthonotal) non-secretory (poroids). Adult male (Fig. 3A). Idiosoma 566–570 long, 322–330 wide between insertions of setae r3; dorsal shield 512–524 long, greatest width 350–358 at level between insertions of r5 (n=2), faintly ornamented and with irregular lateral incisions as in female, margins widened to hold setae j1 anteriorly and r2–r5, Z2, Z4, S2–S4 on lateral subcutaneous extensions. Dorsal shield with 21 or 22 pairs (13 podonotal, 8 or 9 opisthonotal) of smooth setae of which S3 asymmetrically present or absent; setae j1 (10–12), j4–j6, z5 (7–9), J1, J3, J4 (5–7) tiny as in female, occasionally asymmetrically rudimentary; other setal lengths j3 (30–37), z2 (14–18), z4 (47–50), s4 (65–71), s6 (55–60), r2 (25–28), r3 (28–30), r5 (43–45), Z2 (70–72), Z4 (62–67), Z5 (62–63), S2 (44–50), S3 (50–53), S4 (80–82). Soft dorsal peripheral cuticle with setae R1 (37–39) laterally and S5 (68–78) posteriorly. Dorsal shield with complement of ca 22 pairs of discernible pore-like structures (9 podonotal, 13 opisthonotal), many of which are difficult to discern whether secretory (gland pores) or non-secretory (poroids). Idiosomatic venter. Adult female (Fig. 1B). Tritosternum base 27–32 long and 22–27 wide at its base, with paired, shortly ciliated laciniae fused for about 8–15 of entire length (150–162) (Fig. 1F). Presternal region without platelets. Sternal shield mid-length (75–82) much less than narrowest width (125–145) between legs II, unornamented; sternal setae st1, st2 attenuated, of similar length (53–62). Sternal setae st3, st4 similarly attenuated (45–57), with poroid iv3 on soft cuticle. Endopodal strips absent alongside coxae III–IV. Epigynal shield smooth, its rounded hyaline anterior margin reaching or slightly overlapping posterior edge of sternal shield, its lateral margins slightly widened at insertions of attenuated setae st5 (55–62), its posterior margin slightly convex; shield 198–220 long, 55–63 at narrowest width between legs IV, 68–75 at widest level at setae st5. Postgenital groove with elements of one pair of platelets widely separated at level behind paragenital poroids. Opisthosomatic venter with two pairs of small baculiform platelets, longitudinal metapodal platelets 27–34 long, 5–7 wide and nearly transversal postgenital platelets. Anal shield subrhomboidal, unornamented; shield greatest width (118–135) slightly less than length (128–160, including cribrum 12–15), cribrum width 43–50; postanal seta (52–65) about 1.5 longer than paranal setae (35–40) inserted in line with posterior margin of anus. Soft cuticle with seven pairs of opisthogastric setae flanking anal shield, JV1 (40–43), JV2 (38–45), ZV1 (35–40), ZV2 (32–40) anteriorly and JV3 (32–38), ZV3 (27–32), JV5 (52–60) posterolaterally; of three pairs of pore-like structures on soft cuticle, paranal gland pores well separated from lateral margins of anal shield. Peritrematal shield and peritreme extending barely to mid-level of coxae III; peritreme structure simple, without embellished internal chambers (Fig. 1J), its length 52–55, including stigmatal diameter of ca 15; peritrematal shield continuous posteriorly with short exopodal strip extending partly around posterior margin of coxa IV. Exopodal elements absent alongside coxae II–IV. Spermathecal apparatus with simple, elongate (65–75) tubulus leading from solenostome in soft cuticle between bases of legs III–IV to an unpaired sacculus (Fig. 1K). Adult male (Fig. 3C). Tritosternum formed as in female, with laciniae fused for only about one-tenth of entire length (125–140). Presternal region with a pair of platelets preceding genital opening. Sternitigenital shield length 255, narrowest widths 115 at level between coxae II, 100 between coxae III; posterior margin width 63; shield unornamented, narrowly connected to anterior edge of endopodal strips beside coxae IV; alveoli of setae st5 barely contiguous with rounded posterolateral corners of shield; setae st1–st5 attenuated as in female, st4 (26–30) clearly shorter than others (42–50). Paragenital poroids on soft cuticle flanking posterolateral corners of sternitigenital shield. Postgenital groove not developed. Inguinal region with five or six pairs of sigilla indicating unusual muscular development connected with deeply retractable bases of chelicerae.Anal shield similar in rhomboid form, lack of ornamentation as in female, its anterior and lateral apices more irregular; shield longer (136–138, including cribrum ca 12 long) than wide (118–120, cribrum width 45–50); postanal seta (62–65) nearly twice as long as paranal setae (28–37). Soft cuticle with six pairs of opisthogastric setae flanking anal shield, ZV1 absent (present on female); other than JV5, these setae much shorter than in female; JV1 (23–30), JV2 (18–25), JV3 (29–32), ZV2 and ZV3 tiny (7–8), JV5 (49–57); six pairs of pore-like structures evident on soft cuticle, paranal gland pores well separated from lateral margins of anal shield. Peritremes, and peritrematal shield with brief poststigmatal exopodal extension, as in female. Gnathosoma. Adult female. Gnathotectum anterior margin with a single smooth, medial anterior projection extending from a convex margin which smooth or slightly serrate at base of projection (Figs 1G, H). Cheliceral shaft, excluding basal section, 175–178 long; fixed digit with truncated apex bearing two or three tiny serrae, alveolar remnant of pilus dentilis, and minute subapical tooth; movable digit (33–35) with three coarse teeth flared laterally (Figs 1C, D). Subcapitulum with 11 to 13 rows of deutosternal denticles, each similarly narrow, with a single denticle (Fig. 1E); subcapitular setae hp1 (76–80) and hp3 (82–90) similar in attenuated lengths, hp2 (22–27) and capitular setae pc (20) similarly short, but pc strongly bulbiform basally. Corniculi somewhat hyaline, blade-like, narrowly rounded apically, moderately long (50–52), but exceeded by elongated tubular salivary stylets (70–80) and even longer, slender, finely fimbriated internal malae (80–90). Palp length 155–162; palptrochanter setae dissimilar in length, v1 53–63, v2 17–20; palpus with femoral seta al-1 and genual setae al-1, al-2 blunt, not spatulate; palpfemoral seta d (31–35) slightly spine-like, nearly as long as femur; palptarsal apotele two-tined (Figs 2 L–N). Adult male. Gnathotectum with simple anterior margin as in female, but sides more angular, leading to single median projection (Fig. 3E). Cheliceral shaft, excluding short basal section ca 130 long, including fixed digit projection; fixed digit straight, more slender and pointed than in female; movable digit reduced, except for greatly elongated, sinuous spermatodactyl 375–430 long, its base extending anteroventrally beyond apex of fixed digit, gradually narrowing apically, without elaborations (Figs 3B, 18A, 19B). Unlike female, corniculi shorter (25–26), more truncated apically, greatly exceeded by elongated salivary styli (96–100), and capitular setae pc setiform (Fig. 3D). Form and length of other gnathosomatic structures much as in female: internal malae (115–134); setae hp1 (ca 50) and hp3 (70–76) much longer than hp2 and pc (16–18); deutosternum with 11–12 rows of single or rarely doubled denticles. Palp length 150–152. Form and length of palpal setae as in female. Legs. Adult female (Figs 2 A–K). Legs I (462–500) nearly as long as legs IV (495–537), and about 0.9 as long as dorsal shield; other leg lengths (excluding pretarsi): II (395–427), III (405–440). Leg I pretarsus (40–45) to apex of pulvillus, tarsus (125–137) about 1.7–2.1 longer than each of the similarly long femur (73–80), genu (60–65), and tibia (65–70). Legs II–IV with tarsus/tibia length ratios about 1.8–2.1. Complement of setae on leg segments holotrichous for femora I-II-III-IV (12-11-6-6) as presented for the genus; chaetotactic formulae for genua I-II-III-IV: 12 (2-3/2, 2/1-2), 11 (2-3/1, 2/1-2), 9 (2-2/1, 2/1-1), 10 (2-2/1, 3/1-1); tibiae: 12 (2-3/2, 2/1-2), 10 (2-2/1, 2/1-2), 8 (2-1/1, 2/1-1), 10 (2-1/1, 3/1-2); genu I and tibia I without pd-3; tibia III without pl-2. Tarsus I with seta md (65–70) longer than other setae on basal half of segment (Fig. 2G); dorsal subapical cluster of sensory structures with a row of four, tightly adjacent, tiny, pointed setae surrounded by three solenidia, one of which biramous (Figs 2J, K). Tarsi II–IV with a finely serrated ventroapical ridge basal to a short, rounded process (Figs 2E, F), and with setae (d-1) (10–15) nearly half as long as pretarsus to base of claws (22–30). Coxal seta av on leg I and pv on legs II, III strongly bulbiform like subcapitular pc (Figs 2 A–C); coxa IV seta v setiform (Fig. 2D). Trochanters II–IV with basal seta pv-2 attenuated (50–55), and III–IV ad slightly spinelike. Femur IV with seta v large, spine-like (40). Tarsus IV with apical seta pv-1 asymmetrically shorter, but not thicker than av-1 (Fig. 2E). Legs I–IV each with particular dorsal setae enlarged, spinelike: I Fe ad-1 (55–62), pd-1 (37–42); I Ge ad-1 (50–52), pd-1 (55–57); II Fe ad-1 (60–65), pd- 1 (55–60); II Ge ad-1 (34–35), pd-1 (55–60); II Ti pd-1 (29); III Fe ad-1 (45–52); III Ge ad-1 (55–60), pd-1 (60–63); III Ti ad (55), pd-1 (61); IV Fe ad-1 (50–66); IV Ge ad-1 (62–70), pd-1 (65–70); IV Ti ad-1 (52–55), pd-1 (57–60). Relative lengths of other leg setae as shown in Figures 2 A–G. Adult male (Figs 4 A–E). Legs similar in relative dimensions to those of female. Lengths I 440–452, II 367–400, III 392–415, IV 494–495. Legs without noticeable male dimorphism in thickness of segments, with same chaetotaxy as female; legs I–III with form and relative sizes of setae somewhat similar to those of female, including attenuated form of basal seta pv-2 of trochanter, but particular dorsal spine-like setae generally shorter and ventral setae more spine-like on femora, genua, tibiae. Other than trochanter, leg IV segments with nearly all dorsal and ventral setae spine-like; tibia and basal half of tarsus with pd- setae formed as a column of five blunt spine-like setae; femur IV with spine-like seta v distinctly smaller (20) than on female. Lengths of elongated dorsal spine-like setae: I Fe ad-1 (56), pd-1 (20); I Ge ad-1 (27–32), pd-1 (34–37); II Fe ad-1 (65), pd-1 (45); II Ge ad-1 (23), pd-1 (44); III Fe ad-1 (23); III Ge ad-1 (52), pd-1 (41); III Ti ad (37), pd-1 (33); IV Fe ad-1 (67); IV Ge ad-1 (75), pd-1 (50); IV Ti ad-1 (60). Tarsi II and III with setae (v-1), and mv bulbiform; tarsus IV with setae (d-2) and basitarsal (d-3) thickened, spine-like. Deutonymph. Idiosomatic dorsum (Fig. 5A). Idiosoma ca 520 long, ca 290 wide between insertions of setae r3 (n=2); dorsal shields weakly sclerotized, unornamented, completely divided by transverse strip of soft cuticle between setae j6 and J1. Podonotal shield 245–285 long, ca 260–290 wide at its greatest width at level of setae r5, with 10 or 11 pairs of (j1, j3–j6, z2, z4, z5, s4, s6, sometimes r5); opisthonotal shield ca 175–240 long and 210–225 at widest level anteriad setae Z2, with five or six pairs of setae (J1 J3, J4, Z2, Z5, sometimes Z3). Soft cuticle with two or three pairs of podonotal setae (r2, r3, sometimes r5) and five or six pairs of opisthonotal setae (S2–S5, R1, sometimes Z3). Form and relative lengths of dorsal setae as in adult female, all smooth. Setal lengths on podonotum: j1 (11), j3 (35–40), j4 (10–12), j5–j6 (6–7), z2 (17–20), z4 (ca 60), z5 (5–7), s4 (ca 60), s6 (57), r2 (ca 30), r3 (37), r5 (32–35); on opisthonotum: J1 (5–7), J3–J4 (3–5), Z2, Z4 (50–60), Z5 (50–60), S2–S3 (40–42), S4 (45–48), S5 (68–70), R1 (25). Peritreme short (50 including stigma), extending anteriorly to mid-level of coxae III, similar in form to adult. Idiosomatic venter. Tritosternum similar in form to that of adult, with laciniae fused for about 22–29 of entire length (98–112 excluding base). Presternal area devoid of ornamentation. Intercoxal area length 210–225, weakly sclerotized (lateral margins not clearly discerned), bearing five pairs of setae and three pairs of poroids (Fig. 5C); setae st1 (50), st2–st3 (42–43) longer than st4 (27) and st5 (32–35); paragenital poroid on soft cuticle flanking setae st5. Opisthogaster with oboval, smooth anal shield (Fig. 5C); length ca 90, greatest width ca 70; shield with postanal seta (49) stouter and about 1.4 longer than paranal setae (35). Soft opisthogastric integument with seven pairs of setae; JV1, JV2 (30–32), ZV1 (8–10), ZV2 (15), JV3, ZV3 (10–12), JV5 thicker (33–36). Paranal gland pores on anal shield laterally. Gnathosoma. Gnathotectum elongate-angular as in male (Fig. 5B); chelicerae, palpal structures, and subcapitulum much as in female, except setal pair pc simple. Cheliceral shaft, excluding basal section, 145 long; movable digit 28–33 long. Subcapitulum with 11 or 12 rows of deutosternal denticles formed as in female; subcapitular setae hp1 (60–69) and slightly longer hp3 (65–70) similarly attenuated, hp2 (18–22) and capitular setae pc (20–21) similarly short. Corniculi (55) with slightly narrower apices than in female, exceeded by elongated tubular salivary styli (80) and even longer, slender, finely fimbriated internal malae (ca 100). Palpus length 135; palptrochanter setae v1 10, v2 28; palpgenual seta d (10) slightly spine-like. Legs. Leg lengths, excluding pretarsi, I (430–490), II (346–372), III (385–414), IV (450–481). Leg setation with normal deutonymphal complement of setae as given for adult female. Tarsi II to IV with ventroapical process and pretarsal structures formed as in female; dorsoapical setal processes (d-1) relatively short (12–13, about 0.6 as long as pretarsus to base of claws 20–25) as in female; form and relative lengths of setae much as in female, with particular setae enlarged, spine-like: I Fe ad-1 (45–48), pd-1 (31–33); I Ge ad-1 (40–43), pd-1 (47–50); II Fe ad-1 (57), pd-1 (38–42); II Ge ad-1 (36), pd-1 (46–50); III Fe ad-1 (53); III Ge ad-1 (46–50), pd-1 (40); IV Fe ad-1 (56); IV Ge ad-1 (67), pd-1 (61); IV Ti ad-1 (ca 50), pd-1 (ca 50). Tarsus IV with asymmetry in lengths of setae (v-1) as in female. Claws of all legs strongly formed (length ca 15), projecting freely from pulvillus (Fig. 19E), in contrast to adult and protonymph. Protonymph. Idiosomatic dorsum. Idiosoma 405 long, 250 wide at widest level of setae r5 (n=1), with weakly sclerotized dorsal shields (Fig. 6A); podonotal shield 208 long and 225 at level between setae s4, Published as part of Lindquist, Evert E., Oconnor, Barry M., Shaw, Matthew D. & Sidorchuk, Ekaterina A., 2020, Review of the genera Berlesia Canestrini, 1884, and Katydiseius Fain & Lukoschus 1983, the subfamily Katydiseiinae Fain & Lukoschus, 1983, and their family group relationships (Acari: Mesostigmata: Gamasina), with description of three new species parasitic on gryllacridid crickets (Orthoptera), pp. 5-70 in Zootaxa 4857 (1) on pages 9-18, DOI: 10.11646/zootaxa.4857.1.4, http://zenodo.org/record/4396474, {"references":["Lindquist, E. E. & Sidorchuk, E. A. (2019) A new species of Podapolipus (Acari: Heterostigmata: Podapolipidae) from an Australian gryllacridid cricket (Orthoptera), with keys to orthopteran-associated species of the genus. Zootaxa, 4647 (1), 115 - 133. https: // doi. org / 10.11646 / zootaxa. 4647.1.11","Rentz, D. C. F., Su, Y. N. & Bethoux, O. (2018) A new raspy cricket from the northern Australian rainforests (Orthoptera: Gryllacrididae). Zootaxa, 4514 (1), 41 - 52. https: // doi. org / 10.11646 / zootaxa. 4514.1.3","Lindquist, E. E., Bethoux, O., Robin, N. & Sidorchuk, E. A. (2016) The first case of mite-on-mite hyperparasitism? In: 8 th Symposium of the European Association of Acarologists, Valencia, 11 - 15 July 2016. Universitat Politecnica de Valencia, Valencia, pp. 88."]}

Berlesia multisetosa new species (Figures 7–12) Diagnosis Adult: dorsal shield entirely smooth, female’s with irregular indication of lateral incisions at level of setae s6; vertex setae j1, mid-dorsals (j4–j6, z5, J1–J3) and J4 of moderate length (e.g., j4 nearly as long as interval between j4 and j5); dorsal shield setae j3 and z4 slightly longer, but shorter than diagonal interval between them, and shorter than transverse interval between j3; setae z1 and J2 present. Dorsolateral areas of idiosoma with setae of r–R and S series hypertrichous. Soft ventral cuticle hypertrichous, with about 15 pairs of opisthogastric setae. Leg I genu and tibia each with 11 setae, pd-3 and av-2 absent. Tarsus IV with seta pv-1 bluntly spine-like and asymmetrically shorter than attenuated av-1. Peritremes embellished with internal cellular structures. Description Idiosomatic dorsum. Adult female (Fig. 7A). Idiosoma 625–700 long, 412–432 wide between insertions of setae r3; dorsal shield 520–530 long, greatest width 290–320 at level between insertions of s4 (n=2), unornamented except for subdermal sigillar patterns; shield with irregularly formed lateral incisions at level of setae s6. Dorsal shield with usually 10 to 14 pairs (5 to 8 podonotal, 4 to 6 opisthonotal) of smooth setae of which any of z2, s5, J4 or Z4 asymmetrically on/off shield margin; of shield setae, j3 (60–65), z4 (62–75), s4 (62–75), s5 (60–70) of similar lengths; j4 (45–65), j5 (40–60), j6 (31–53), z5 (33–52), J1–J4 somewhat shorter but not tiny (22–40), with z4 and s4 (62–75), s5 (60–67), Z1 (57–70) and Z4 (60–72) similarly longer. Soft dorsal peripheral cuticle with setae j1 (47–52), z1 (40–43), z2 (54) anteriorly, s6 (60–65) and a hypertrichy of 12–15 r -setae (50–62) anterolaterally, and Z2 (57–65), S1–S4 (55–70), S5 (75) and hypertrichy of 10–15 R -setae (52–65) posterolaterally. Dorsal shield with complement of nine pairs of discernible pore-like structures (four podonotal, five opisthonotal), of which three pairs (one podonotal, two opisthonotal) superficially appear secretory (gland pores) and six pairs (three podonotal, three opisthonotal) non-secretory (poroids). Adult male (Fig. 9A). Idiosoma 580–612 long, 370–437 wide between insertions of setae r3; dorsal shield 520– 532 long, greatest width 337–365 at level between insertions of s6 (n=3), unornamented, with irregular but entire lateral margins, which widened to hold setae j1, z1, s3 anteriorly and Z2, S2 laterally. Dorsal shield with usually 23 to 25 pairs (14 or 15 podonotal, 9 or 10 opisthonotal) of smooth setae of which s6, Z2, Z4 often asymmetrically off shield margin; relative setal lengths much as in female, except J1–J4 shorter (20–30) compared to j4–j6 (37–57); other setal lengths j1 (35–42), j3 (58–68), z1 (25–32), z2 (55–62), z4 (60–65), s4–s6 (57–67), about eight r -setae (40–52), Z1–Z2 (60–65), Z3 (60–72), Z4–Z5 (65–77), S1–S3 (62–68), S4 (65–72), S5 (68–85), about ten R -setae (52–68). Dorsal shield with complement of eleven pairs of discernible pore-like structures (4 podonotal, 7 opisthonotal), most of which are difficult to discern whether secretory (gland pores) or non-secretory (poroids). Idiosomatic venter. Adult female (Fig. 7B). Tritosternum base 23–25 long and 23–26 wide at its base, with paired ciliate laciniae fused for about 5–7 of entire length (ca 100). Presternal area with a delineated, lightly sclerotized, streaky region without clear platelets. Sternal shield mid-length (98–100) much less than narrowest width (148–155) between legs II, faintly scabrous; sternal setae st1, st2 attenuated, of similar length (55–67). Sternal setae st3, st4 similarly attenuated (50–60), with poroid iv3 on soft cuticle. Endopodal strips rudimentary alongside coxae III–IV. Epigynal shield smooth, its rounded hyaline anterior margin reaching or slightly overlapping posterior edge of sternal shield, its lateral margins slightly widened at insertions of attenuated setae st5 (57–65), its posterior margin slightly convex; shield 205–223 long, 60–63 at narrowest width between legs IV, 77–85 at widest level at setae st5. Postgenital groove undeveloped, without elements of platelets. Opisthosomatic venter with two pairs of weak metapodal platelets, lateral pair 30–32 x 8–12, inner pair 17–20 x 7–10. Anal shield subrhomboidal, lightly lineated; shield greatest width (128) slightly less than length (132–147, including cribrum 20), cribrum width 75; postanal seta (60–70) about 1.5 longer than paranal setae (37–50) inserted in line with posterior margin of anus. Soft cuticle with ca 15 pairs of opisthogastric setae of similar lengths (43–57) flanking anal shield, JV5 not distinguishable posterolaterally; of four or five pairs of pore-like structures on soft cuticle, paranal gland pores well separated from lateral margins of anal shield. Peritrematal shield and peritreme extending barely to mid-level of coxae III; peritreme with embellished internal chambers (Fig. 7F), its length 45, including stigmatal diameter of ca 22; peritrematal shield continuous posteriorly with short exopodal strip extending partly around posterior margin of coxa IV. Exopodal elements absent alongside coxae II–IV. Spermathecal apparatus with simple, elongate (90–95) tubulus leading from solenostome between bases of legs III–IV to an unpaired sacculus (Fig. 7G). Adult male (Fig. 9B). Tritosternum formed as in female, with laciniae fused for only about one-tenth of entire length (ca 100). Presternal region with two pairs of small platelets anterior to genital opening. Sternitigenital shield length 245–255, narrowest widths 115–125 at level between coxae II, 100–107 between coxae III; posterior margin width 62–80; shield unornamented, narrowly separated from anterior edge of endopodal strips beside coxae IV; alveoli of setae st5 on corners of irregularly shaped posterior margin of shield; setae st1–st5 similarly attenuated as in female, st1 and st5 (53–67) slightly longer than st2–st4 (50–65). Paragenital poroids on soft cuticle flanking posterolateral corners of sternitigenital shield. Postgenital groove not developed. Inguinal region with five or six pairs of sigilla indicating unusual muscular development connected with deeply retractable bases of chelicerae. Anal shield similar in rhomboid form, slight ornamentation as in female, its anterior and lateral apices more irregular; shield longer (130–135, including cribrum ca 15–20 long) than wide (103–112, cribrum width 60–62); postanal seta (57–62) ca 1.5 to nearly twice as long as paranal setae (30–40). Soft cuticle with about 15 pairs of opisthogastric setae flanking anal shield; these setae gradually slightly lengthening posteriorly, those in JV1-JV2-ZV1-ZV2 area 43–48, those mid-posteriorly 45–62, and posterior-most JV5 (62–75); of five pairs of pore-like structures evident on soft cuticle, paranal gland pores well separated from lateral margins of anal shield. Short but embellished peritremes (40–45), and peritrematal shield with brief poststigmatal exopodal extension as in female (Figs 9J, 20B). Gnathosoma. Adult female. Gnathotectum anterior margin obtusely smoothly triangular, with a blunt, medial anterior projection (Figs 7H, J). Cheliceral shaft, excluding basal section, 155–158 long; fixed digit with blunt apex, vestige of pilus dentilis, and a vestigial tooth midway on paraxial face (Fig. 7E); movable digit (35) with three coarse teeth flared laterally (Figs 7D, E). Subcapitulum with 11 to 13 rows of deutosternal denticles, each similarly narrow, with a single denticle, or any row occasionally with two contiguous denticles (Fig. 7C); subcapitular setae hp1 (60–63) and slightly longer hp3 (75–80) similarly attenuated, hp2 (20–25) and capitular setae pc (22–27) similarly short, but pc strongly bulbiform basally. Corniculi somewhat hyaline, blade-like, bluntly pointed apically, moderately short (35–38), closely adjacent to similarly short tubular salivary styli (ca 37–40) and longer, slender, finely fimbriated internal malae (87–90). Palp length 150–162; palptrochanter setae dissimilar in length, v1 22–25, v2 38–52; palpus with femoral seta al-1 and genual setae al-1, al-2 blunt, not spatulate; palpfemoral seta d (30–32) slightly spine-like, shorter than femur length; palptarsal apotele two-tined. Adult male. Gnathotectum with simple, triangular anterior margin much as in female, but sides acutely angular, tapering to single median projection (Fig. 9E). Cheliceral shaft, excluding short basal section 120–125 long; fixed digit straight, short (30–42) more slender and pointed than in female; movable digit bluntly pointed beyond apex of fixed digit, reduced, amalgamated into spermatodactyl base which extends anteroventrally; spermatodactyl greatly elongated, sinuous, gradually tapering, 425–465 long, without elaborations (Figs 9C, D). Unlike female, corniculi shorter (22–25), more truncated apically, greatly exceeded by elongated salivary styli (63–70), and capitular setae pc nearly setiform, slightly bulbous basally. Form and length of other gnathosomatic structures much as in female: internal malae (120–130); setae hp1 (62–63) and hp3 (65–77) much longer than hp2 (18–20) and pc (23–25); deutosternum with 11–12 rows of mostly contiguously doubled denticles (Fig. 9F). Palp length 160–165; palpfemoral seta d (25–30) similar in form as in female. Legs. Adult female (Figs 8 A–H). Legs I (425–450) nearly as long as legs IV (460–470), and about 0.8 as long as dorsal shield; other leg lengths (excluding pretarsi): II (375–380), III (385–400). Leg I pretarsus (37–38) to apex of pulvillus, tarsus (115–125) about 1.7–2.1 longer than each of the similarly long femur (67–70), genu (53–63), and tibia (60–65). Legs II–IV with tarsus/tibia length ratios about 1.8–2.3. Complement of setae on leg segments holotrichous for femora II-III-IV (11-6-6) as presented for the genus, but femur I with 11 setae, lacking pv-2; chaetotactic formulae for genua I-II-III-IV: 11 (2-3/1, 2/1-2), 11 (2-3/1, 2/1-2), 9 (2-2/1, 2/1-1), 10 (2-2/1, 3/1-1); tibiae: 11 (2-3/1, 2/1-2), 10 (2-2/1, 2/1-2), 8 (2-1/1, 2/1-1), 10 (2-1/1, 3/1-2); genu I and tibia I without pd-3 and av-2; tibia III without pl-2. Tarsus I with seta md (55–60) longer than other setae on basal half of segment (Fig. 8B); dorsal subapical cluster of sensory structures with a cluster of three, tightly adjacent, tiny, pointed setae surrounded by three solenidia, one of which biramous (Fig. 8C). Tarsi II–IV with a finely serrated ventroapical ridge basal to a short, rounded process (Figs. 8E, H), and with dorsoapical setae (d-1) (12–15) nearly half as long as pretarsus to base of claws (ca 25). Coxal seta av on leg I and pv on legs II, III strongly bulbiform basally, like subcapitular pc; coxa IV seta v setiform. Trochanters I–IV with basal seta pv-2 attenuated (50–55), and III–IV ad slightly spinelike. Tarsus IV with apical seta pv-1 asymmetrically shorter, bluntly spine-like, than attenuated av-1 (Fig. 8H). Legs I–IV each with particular setae enlarged, spine-like: I Fe ad-1 (65), pd-1 (45); I Ge ad-1 (50–60), pd-1 (55–58); II Fe ad-1 (57–63), pd-1 (47–48); II Ge ad-1 (50–52), pd- 1 (47–48); II Ti pd-1 not differentiated; III Fe ad-1 (52–54); III Ge ad-1 (57–62), pd-1 (50–52); III Ti ad (42), pd-1 (30); IV Fe ad-1 (57–62), pv not differentiated; IV Ge ad-1 (66–68), pd-1 (62–65); IV Ti ad-1 (53–60), pd-1 (55–60). Relative lengths of other leg setae as shown in Figs 8A, D, F–H. Adult male (Figs 10 A–D). Legs similar in relative dimensions to those of female; lengths I 415–430, II 360– 378, III 375–400, IV 440–470. Legs I and IV without dimorphism in thickness of segments, legs II and III thickersegmented than in female. Leg I with form and relative sizes of setae similar to those of female, except femur I with 12 setae, including pv-2, and with av slightly bulbiform. Legs I–IV retain attenuated form of basal seta pv-2 of trochanter. Legs II–IV with particular setae dimorphically modified: leg II Fe av spine-like; Ge av, Ti av, and tarsus mv, av-2, av-1, pv-1 bulbiform; leg III coxal pv simple; Tr ad markedly arched; Fe al less spine-like, and tarsus mv, av-2, pv-2, av-1, pv-1 bulbiform; leg IV Tr ad markedly arched, av, pv-1 thickened, more spine-like; Fe pd, pl more spine-like; Ge pd-2, pd-3 and Ti pl-1 larger, more spine-like; tarsus av-2, pv-2, av-1, pv-1 bulbiform. Lengths of spine-like setae similarly enlarged as in female: I Fe ad-1 (48–62), pd-1 (37–45); I Ge ad-1 (45–52), pd-1 (52–60); II Fe ad-1 (55–65), pd-1 (42–49); II Ge ad-1 (38–45), pd-1 (50–57); II Ti pd-1 not differentiated; III Fe ad-1 (40–50); III Ge ad-1 (43–53), pd-1 (47–50); III Ti ad (35), pd-1 (27); IV Fe ad-1 (57–70), pv not differentiated; IV Ge ad-1 (67–70), pd-1 (67–75); IV Ti ad-1 (55–65), pd-1 (60–67). Deutonymph. Idiosomatic dorsum (Fig. 11A). Idiosoma 475–500 long, 362–365 wide between insertions of setae r3 (n=3); dorsal shields weakly sclerotized, unornamented, faintly divided by transverse strip of soft cuticle between setae j6 and J1. Podonotal shield 210–225 long, ca 215–225 wide at its greatest width at level of setae s5, with ca 12 pairs of setae (j1, j3–j6, z1, z2, z4. z5, s4–s6); opisthonotal shield ca 175–195 long and 175–205 at widest level near setae Z2, with nine pairs of setae (J1–J4, Z1–Z5). Soft cuticle with ca 25 pairs of dorsolateral setae, including uncertain homologies of the s -and S -series among hypertrichous elements of the marginal r–R -series (Fig. 11A). Form and relative lengths of dorsal setae more homologous than in adult, many of them slightly barbed as shown in Fig. 11A; other than z1 clearly shortest (18–28), setal lengths range from j1 (35–40) to Z5 longest (65). Peritreme similarly short (length 37–38, including stigma diameter 15–17) and embellished as in adult (Fig. 11D). Idiosomatic venter (Fig. 11B). Tritosternum similar in form to that of adult, with laciniae fused for only about 5–10 of entire length (87–92 excluding base). Presternal area devoid of ornamentation. Intercoxal shield 182–200 long, weakly sclerotized, bearing five pairs of setae and three pairs of poroids, seta st5 sometimes barely off shield edge; setae st1–st4 (43–50) slightly shorter than st5 (52–57); paragenital poroids on soft cuticle flanking posterior edge of shield. Opisthogaster with oboval, smooth anal shield; length 95–100, greatest width 75–80; shield with postanal seta (45–50) about 1.3 longer and stouter than paranal setae (35–37). Soft cuticle with hypertrichous complement and relative lengths of setae flanking anal shield much as in adult, with ca 15 pairs of slender opisthogastric setae flanked by 5–10 pairs of thicker R -marginal setae of similar lengths (40–47), the more posterolateral elements of them thicker, blunter. Gnathosoma. Gnathotectum, chelicerae, palpal structures, and subcapitulum much as in female, except setal pair pc simple. Cheliceral shaft, excluding basal section, 132–138 long; movable digit 28–35 long. Subcapitulum with 11 to 13 rows of deutosternal denticles formed as in female; subcapitular setae hp1 (45–50) and slightly longer hp3 (52–58) similarly attenuated, hp2 (13–15) and capitular setae pc (15–18) similarly short. Corniculi (25–30) formed as in female, exceeded by elongated tubular salivary styli (45–50) and even longer, slender, finely fimbriated internal malae (70–75). Palpus length 127–133; palptrochanter setae v1 12–15, v2 30–34; palpgenual seta d (22–23) slightly spine-like. Legs. Leg lengths, excluding pretarsi, I (325–350), II (278–312), III (287–315), IV (335–360). Leg setation with normal deutonymphal complement of setae as given for adult female, with occasional variabilities. Tarsi II to IV with ventroapical process and pretarsal structures formed as in female; apical dorsosetal processes (d-1) relatively short (9–10, about 0.6 as long as pretarsus to base of claws 13–15) as in female; form and relative lengths of setae much as in female, with particular setae enlarged, spine-like: I Fe ad-1 (45–50), pd-1 (30–33); I Ge ad-1 (38–45), pd-1 (37–43); II Fe ad-1 (50–53), pd-1 (32–33); II Ge ad-1 (35–38), pd-1 (34–35); III Fe ad-1 (45–50); III Ge ad-1 (42–43), pd-1 (37–40); IV Fe ad-1 (47–52); IV Ge ad-1 (45–20), pd-1 (40–48); IV Ti ad-1 (40–47), pd-1 (36–42). Tarsus IV with asymmetry in lengths of setae (v-1) as in female. Claws of all legs strongly formed (length ca 13), projecting freely from pulvillus (as in Fig. 19E), in contrast to adult and protonymph. Protonymph. Idiosomatic dorsum (Fig. 12A). Idiosoma 370 long, 268 wide at widest level of setae r3, r5 (n = 1), with weak

Berlesia vorontsovi new species (Figures 13–17) Diagnosis Adult: dorsal shield faintly reticulated posteriorly, smooth elsewhere, with irregular indication of lateral incisions at level of setae s6; setae j1, mid-dorsals (j4–j6, z5, J1, J3) very short; dorsal shield setae j3 and z4 relatively long, about as long or slightly longer than diagonal interval between them, and longer than transverse interval between j3; setae z1, J2 and J4 absent. Dorsolateral regions of idiosoma with setae of r -R -and S -series sparse, not hypertrichous. Soft ventral cuticle with 6 and 4 pairs of opisthogastric setae on female and male, respectively. Leg I genu and tibia each with 10–11 setae, pd-3, av-2 and sometimes ad-3 absent. Tarsus IV with seta pv-1 asymmetrically shorter, but similar in attenuated form to av-1. Peritremes embellished with internal cellular structure. Description Adult: Idiosomatic dorsum. Adult female (Fig. 13A). Idiosoma 741–791 long, 432–451 wide between insertions of setae r3; dorsal shield 580 long, greatest width 306–324 at level between insertions of s4 (n=4), ornamentation not apparent, shield with irregular indication of lateral incisions at level of setae s6 and irregular indentations more posteriorly. Dorsal shield with usually 9 or 10 pairs (7 or 8 podonotal, 2 opisthonotal) of smooth setae of which j1 may be asymmetrically on/off shield margin and J4 always absent; setae j3 (131–140), z4 (133–137), s4 (135–140) of similarly long lengths; other shield setae (j4–j6, z5, J1, J3) tiny (ca 5), occasionally asymmetrically vestigial. Soft dorsal peripheral cuticle with tiny setae j1 (10–12) anteriorly (occasionally on shield), z2 (110–119), s6 (110–126) and r2, r3, r5 (103–105) anterolaterally, and Z2 (108–119), Z4 (133–142), S2 (85–89), S3 (60–89), S4 (96–140), S5 (128) and R1 (78–92) posterolaterally, Z5 (92–101) slightly off shield posteriorly. Dorsal shield pore-like structures difficult to discern (maximum observed complement shown in Fig. 13A). Adult male (Fig. 15A). Idiosoma 672 long, 409–432 wide between insertions of setae r3; dorsal shield 570 long, greatest width 321–335 at level between insertions of r5 (n=3), without distinct ornamentation and with shallow lateral incisions, setae j1 off shield anteriorly, z2 and r2–r5, Z2, Z4, S2–S4 on lateral subcutaneous extensions. Dorsal shield with 9 or 10 pairs (7 podonotal, 2–3 opisthonotal) of smooth setae; setae j1 (7–8), j4–j6, z5 (3), J1, J3 (3) occasionally asymmetrically vestigial or absent, J4 absent; other setal lengths j3 (92–108), z2 (41–46), z4 (92–117), s4 (105–108), s6 (96–103), r2 (64–76), r3 (71–76), r5 (73–78), Z2 (108–115), Z4 (108–115), Z5 (92–96), S2 (60–64), S3 (53–76), S4 (92–110), S5 (119–133), R1 (55–73). Dorsal shield with complement of ca 15 pairs of discernible pore-like structures (4 podonotal, 11 opisthonotal), many of which are difficult to discern whether secretory (gland pores) or non-secretory (poroids). Idiosomatic venter. Adult female (Fig. 13B). Tritosternum base 25–30 long and 28–34 wide at its base, with paired ciliated laciniae fused for about 8% of entire length (92–103) (Fig. 13F). Presternal region without platelets and lineation. Sternal shield mid-length (96–98) much less than narrowest width (158–172) between legs II, unornamented; sternal setae st1 (96–110), st2 (69–87), st3 (87–92), st4 (62–69) attenuated, with poroid iv3 on soft cuticle. Endopodal strips absent alongside coxae III–IV. Epigynal shield smooth, its rounded hyaline anterior margin reaching or slightly overlapping posterior edge of sternal shield, its lateral margins widened at insertions of attenuated setae st5 (73–82), its posterior margin slightly convex, occasionally indented or with fragmented platelets posteriorly; shield 185–218 long, 46–48 at narrowest width between legs IV, 68–76 at widest level at setae st5. Opisthosomatic venter with several irregularly spaced platelets widely separated at level between metapodal plates, and with two pairs of small ovoid metapodal platelets, anterior pair 14–16 long, 5–7 wide, posterior pair 25–30 long, 14–21 wide. Anal shield subrhomboidal, weakly reticulate posteriorly, without paranal gland pores; shield greatest width (126–130) slightly less than length (137–149, including cribrum), cribrum U-shaped, width 55–64; postanal seta (69–73) longer than paranal setae (37–53) inserted in line with posterior margin of anus. Soft cuticle with maximum of 6 pairs of opisthogastric setae flanking anal shield, JV1 (53–60), JV2 (46–48), ZV2 (44–46) anteriorly and JV3 (39–48), ZV3 (46–53), JV5 (60–66) posterolaterally, ZV1 and JV4 setae always absent, JV2, JV3, JV5, ZV2 sometimes unilaterally or bilaterally absent; one pair of pore-like structures observed on soft cuticle. Peritrematal shield and peritreme extending to posterior margin of coxae III; peritreme structure with embellished internal chambers (Fig. 13E), its length 60–62, including stigmatal diameter of ca 11; peritrematal shield continuous posteriorly with short exopodal strip extending partly around posterior margin of coxa IV. Exopodal elements absent alongside coxae II–IV. Spermathecal apparatus with simple, elongate (80–119) tubulus leading from solenostome in soft cuticle between bases of legs III–IV to an unpaired sacculus (Fig. 13G). Adult male (Fig. 15B). Tritosternum formed as in female, with laciniae fused for only about one-tenth of entire length (98–100). Presternal region with a pair of weakly developed platelets abutting genital opening, one or both platelets sometimes fused with sternitigenital shield. Sternitigenital shield length 252–263, narrowest widths 124–134 at level between coxae II, 124–131 between coxae III; posterior margin width 62–69; shield with scalelike ornamentation anterior to setae st4, narrowly connected to anterior edge of endopodal strips beside coxae IV; alveoli of setae st5 barely contiguous with rounded posterolateral corners of shield; setae st1–st5 attenuated as in female, st4 (34–37) clearly shorter than others, st1 (76), st2 (66–73), st3 (66–69), st5 (57–69). Paragenital poroids on soft cuticle flanking posterolateral corners of sternitigenital shield. Postgenital groove not developed. Inguinal region with six or seven pairs of small platelets or sigilla as in B. hospitabilis. Anal shield rhomboid with a variously formed anterior extension, weak ornamentation present laterally; shield longer (144–156, including cribrum ca 34–60 long) than wide (112–119, cribrum width 64–73); postanal seta (73–78) about 1.5 times as long as paranal setae (48–50). Soft cuticle with four pairs of opisthogastric setae flanking anal shield; other than JV5, these setae shorter than in female, and ZV1, ZV3, JV3, JV4 absent; JV1 (46–48), JV2 (34–41), JV5 (37–48), ZV2 (25–30); pore-like structures on soft cuticle difficult to discern. Peritremes, and peritrematal shield with brief poststigmatal exopodal extension, as in female. Gnathosoma. Adult female. Gnathotectum anterior margin acutely triangular with very weakly serrate edges (Figs 13H). Cheliceral shaft, excluding basal section, 160–165 long; fixed digit with truncated apex and minute subapical tooth, other structures not observed; movable digit (23–25) with three coarse teeth flared laterally (Figs 13C, D). Subcapitulum with 11 relatively narrow rows of deutosternal denticles, varying somewhat in width and consisting from a single denticle to multidentate (Fig. 13J); subcapitular setae hp1 (62–82) and hp3 (82–103) attenuated, hp2 (30–44) and capitular setae pc (39–41) similarly short, but pc with strongly bulbiform base and laterally positioned flagellum. Corniculi somewhat hyaline, blade-like, narrowly rounded apically (39–44), but exceeded by elongated tubular salivary styli (60–64) and slender, finely fimbriated internal malae (57–62). Palp length 183–188; palptrochanter setae dissimilar in length, v1 46–78, v2 41–46; palpus with femoral seta al-1 acicular, genual setae al-1 spatulate, al-2 acicular; palpfemoral seta d (60–64) spine-like, typically longer than femur (Fig. 13J); palptarsal apotele two-tined (Fig. 13K). Adult male. Gnathotectum slightly serrate anteriorly as in female, sides almost parallel posteriorly, leading to single median projection (Fig. 15D). Cheliceral shaft, excluding short basal section 92–94 long, including fixed digit projection; fixed digit straight, more slender and pointed than in female; movable digit reduced, except for greatly elongated, sinuous spermatodactyl 404–444 long, its base extending anteroventrally beyond apex of fixed digit, gradually narrowing distally, abruptly hooked apically (Fig. 15C). Unlike female, corniculi shorter (25–30), more truncated apically, greatly exceeded by salivary styli; capitular setae pc with bulbous base as in female (Fig. 15E). Form and length of other gnathosomatic structures much as in female: attenuated salivary styli (44), internal malae (89–108); setae hp1 (57–69) and hp3 (78–92) much longer than hp2 and pc (21); deutosternum with 11–12 rows of denticles, basal 4 rows multidentate, apical rows with 1–2 denticles (if two, then one is always distinctly larger). Palp (Figs. 15F, G) length 179–185. Form and lengths of palpal setae as in female. Legs. Adult female (Figs 14 A–F). Legs I (561–589) distinctly shorter than legs IV (644–672), and only slightly shorter than dorsal shield; other leg lengths (excluding pretarsi): II (524–552), III (543–561). Leg I pretarsus (65–89) to apex of pulvillus, tarsus (169–176) about 1.9–2.1 longer than each of the similarly long femur (85–92), genu (80–85), and tibia (87–92). Legs II–IV with tarsus/tibia length ratios about 1.7–2.1. Complement of setae on leg segments holotrichous for femora I-II-III-IV (12-11-6-6) as presented for the genus; chaetotactic formulae for genua I-II-III-IV: 10–11 (2-2 or 3/1, 2/1-2), 11 (2-3/1, 2/1-2), 9 (2-2/1, 2/1-1), 10 (2-2/1, 3/1-1); tibiae: 10–11 (2-2 or 3/1, 2/1-2), 9–10 (2-1 or 2/1, 2/1-2), 7–8 (1 or 2-1/1, 2/1-1), 10 (2-1/1, 3/1-2); genu I and tibia I without pd-3, av-2, sometimes without ad-3; tibia II sometimes without ad-2; tibia III without pl-2, sometimes without al-2. Tarsus I with seta md (103–110) longer than other setae on basal half of segment (Fig. 14B); dorsal subapical cluster of sensory structures with a row of three tightly adjacent, tiny, pointed setae surrounded by three solenidia, one of which biramous (Fig. 14B). Tarsi II–IV with ventroapical ridge, serrations not discernible, short, rounded process not discernible, and with setae (d-1) (ca 14–16) less than half as long as pretarsus to base of claws (34–41). Coxal seta av on leg I and pv on legs II, III strongly bulbiform basally like subcapitular pc (Figs 14A, C, D); coxa IV seta v setiform (Fig. 14F). Trochanters II–IV with basal seta pv-2 attenuated (71–94), and III–IV ad slightly spinelike. Femur IV with seta v large, somewhat spine-like (49). Tarsus IV with apical seta pv-1 asymmetrically shorter, but not thicker than av-1 (Fig. 14E). Legs I–IV each with particular dorsal setae enlarged, spinelike: I Fe ad-1 (103– 126), pd-1 (50–85); I Ge ad-1 (87–92), pd-1 (112–128), ad-2 (48–50); II Fe ad-1 (115–121), pd-1 (80–82); II Ge ad-1 (69–76), pd-1 (119–121), ad-2 (69–71); II Ti pd-1 (69–80); III Fe pd-1 (115–124); III Ge ad-1 (92–117), pd-1 (92–110); III Ti ad (85–96), pd-1 (69–73); IV Fe ad-1 (114–135); IV Ge ad-1 (124–133), pd-1 (92–121); IV Ti ad-1 (92–105), pd-1 (92–103). Note: all long setae on leg IV in the holotype are stunted on one side, their measurements not included above. Relative lengths of other leg setae as shown in Figures 14 A–F. Adult male (Figs 16 A–E). Legs similar in relative dimensions to those of female, including legs IV distinctively longest. Lengths I 478–488, II 423–442, III 442–460, IV 561–598. Legs without noticeable male dimorphism in thickness of segments, with same setation as female; legs I–III with form and relative sizes of setae similar to those of female, including dorsal spine-like setae, but ventral setae more spine-like on femora, genua, tibiae. Basal trochanter seta pv-2 slender, attenuated as on female, though less so on legs III and IV, which also have pv-1 shorter, thicker than on female. Leg IV segments with nearly all dorsal and ventral setae spine-like; genu, tibia and basal half of tarsus with pd- setae normally formed (not as a column of blunt spine-like setae as in male B. hospitabilis); femur IV with seta v slightly smaller (37–45) than on female. Lengths of elongated dorsal spine-like setae: I Fe ad-1 (92–96), pd-1 (34–41); I Ge ad-1 (71–87), pd-1 (92–94); II Fe ad-1 (92–110), pd-1 (66–73); II Ge ad-1 (48–60), pd-1 (92–96); III Fe ad-1 (94–98); III Ge ad-1 (80–89), pd-1 (79–89); III Ti ad (50–64), pd-1 (48–55); IV Fe ad-1 (92–137); IV Ge ad-1 (105–115), pd-1 (98–110); IV Ti ad-1 (80–92), pd-1 (85–94). Tarsi II and III with setae (v-1), and mv bulbiform; tarsus IV with setae (d-2) and basitarsal (d-3) not thickened, spine-like. Deutonymph. Unknown. Protonymph. Idiosomatic dorsum. Idiosoma 495–543 long, 381–437 wide at widest level of setae r5 (n = 4), with weakly sclerotized dorsal shields (Fig. 17A); podonotal shield 294–312 long and 308–321 wide at level between setae s4, strongly ornamented with a network of lines, with 12 pairs of setae (j3–j6, z2, z4, z5, s4, s6, r2, r 3,r5 ); pygidial shield typically narrowly fused with anterior lateral sclerites, rarely slightly separated, without obvious ornamentation, 188–197 long and 266–280 at widest level between setae R1, with five pairs of setae (J1, J3, Z2, Z3, Z5), J4 absent. Soft cuticle with only one pair of podonotal setae (j1) and two pairs of opisthonotal setae (S2, R1); S3-S5 absent. Form and relative lengths of dorsal setae as in adult, all smooth. Setal lengths on podonotum: j1 (6–7), j3 (69–76), j4–j6 (6), z2 (30–37), z4 (78–87), z5 (6), s4 (80–87), s6 (55–57), r2 (18–25), r3 (30–37), r5 (32–37); on opisthonotum: J1 (7), J3 (7), Z2 (62–64), Z3 (44–46), Z5 (62–69), S2 (32), R1 (27–32). Peritreme (46 including stigma), extending anteriorly to posterior edge of coxae III. Idiosomatic venter. Tritosternum formed much as in adult, with laciniae fused for 11–16 of total length (73–89). Presternal area devoid of ornamentation. Sternal shield with weak but distinct outline, strongly narrowing between setae st2 and st3, length 144, with setae st1–st2 (48–60) and poroids iv1, iv2; setae st3 (48–53) on soft cuticle; setae st5 and paragenital poroids not discernible (Fig. 17B). Anal shield smooth, subrhomboid, broadest between paranal and postanal setae, length 92–94 including cribrum, width 78–80 (cribrum strip width 39–44); postanal seta (39–50) only slightly longer than paranal setae (39–44). Soft opisthogastric integument with four pairs of setae: JV1 (34–41), JV2 (18–25), ZV2 (16–21), JV5 longer (48–55), ZV2 unilaterally absent in 3 of 4 specimens. Gnathosoma. Gnathotectum with anterior margin acutely triangular, tapering to single median projection, apparently smooth (Fig. 17D). Cheliceral shaft, excluding short basal section 119–124 long; fixed digit straight, more bluntly pointed than in female; movable digit (18–21) formed much as in female, with three teeth flared laterally (Fig. 17C). Subcapitulum with 13 rows of deutosternal denticles, 5 basal rows slightly wider, multidentate, remaining rows uni-or bidentate; subcapitular setae hp1 and hp3 similar in attenuated lengths (32–48), hp2 (21–22) and capitular setae pc (26–30) similar in short simple form. Corniculi hyaline, blade-like, bluntly pointed apically, moderately short (24–25), greatly exceeded by elongated tubular salivary styli (46). Palpus length 137–142, with normal protonymphal complement of setae; single palptrochanter seta 9–11; palpfemoral seta d (30–32) slightly thickened. Legs. Leg lengths, excluding pretarsi, I 206, II 183, III 181–183, IV 211–215. Leg setation of protonymph typical of general holotrichous pattern presented for free-living Gamasina by Evans (1963): coxae, 2-2-2-1; trochanters, 4-4-4-4; femora, 10 (2 2/1 2/1 2)-8 (1 2/1 2/1 1)-5 (1 2/1 1/0 0)-4 (1 2/0 1/0 0); genua, 8 (1 2/1 2/1 1)-6 (1 2/0 2/0 1)-6 (1 2/0 2/0 1)-5 (1 2/0 2/0 0); tibiae, 8 (1 2/1 2/1 1)-7 (1 1/1 2/1 1)-7 (1 1/1 2/1 1)-7 (1 1/1 2/1 1). Legs I–IV each with particular setae enlarged, spine-like, with similar pattern as for adults. Tarsi II–IV with ventroapical process, pretarsal structures and weakly developed claws formed as in female. Tarsus IV with asymmetry in lengths of setae (v-1) as in female. Larva. No material available (adult females anticipated to be nymphiparous). Type material. Type specimens: all collected by B.M. OConnor (BMOC 84-1800-039) from one adult female c

Key to genera and species of Katydiseiinae, adult females 1. Dorsal vertex setae j1 absent (not to be confused with j3, present slightly behind level of poroids id-1); sternal shield reduced, bearing only setae st1, without endopodal extensions between bases of legs I–II and II–III; anal shield with paranal setae inserted at mid-level of anus; subcapitulum with hypostome prolonged anteriorly by two paramedian processes reaching level of palptibia and enveloping extremities of internal malae and labrum; coxa of legs I–IV with all setae setiform; genua of legs I-II-III with 8-6-6 setae, each lacking al-2, pl-2......................................................................................................... Genus Katydiseius, monobasic, K. nadchatrami Fain & Lukoschus, 1983 - Dorsal vertex setae j1 present (inserted slightly anteromediad poroids id-1, closely anterior to j3); sternal shield well developed, bearing setae st1, st2, with endopodal extensions between bases of legs I–II and II–III; anal shield with paranal setae inserted at level of, or behind, posterior margin of anus; subcapitulum with sclerotized extensions of hypostome extended only as corniculi to level of palptrochanter, but with flexible mouthpart structures (salivary styli, internal malae, labrum) extending freely to level of palpgenu; coxa of legs I–III and sometimes IV each with one seta strongly bulbiform basally; genua of legs I-II-III with minimally 11-10-9 setae, each with al-2, pl-2... Genus Berlesia Canestrini, 1884............................... 2 2. Coxal seta of leg IV conspicuously bulbiform, similar to one of coxal setae on each of legs I–III and subcapitular setae pc; anal shield lateral margins nearly parallel; tarsi II–IV each with one of apical setae distinctively longer than adjacent setae and as long as entire pretarsus.............................................................. B. rapax Canestrini, 1884 - Coxal seta of leg IV setiform, in contrast to a bulbiform coxal seta on each of legs I–III and subcapitulum; anal shield subrhomboidal, lateral margins angular; tarsi II–IV each with apical setae similar in length, none as long as entire pretarsus........ 3 3. Vertex setae j1 moderately long, z1 present; dorsal shield with mid–dorsal setae (j4–j6, z5, J1–J3) also moderately long (J2 present), e.g., j4 nearly as long as interval j4–j5; dorsal shield flanked by about 25 pairs of setae, mostly of the hypertrichous r–R and S series; soft opisthogastric cuticle also hypertrichous, with about 15 pairs of setae........... B. multisetosa n. sp. - Vertex setae j1 short, z1 absent; dorsal shield with mid-dorsal setae also short (J2 absent), e.g., j4 ca one-fourth as long as interval j4–j5; dorsal shield flanked by at most 15 pairs of setae mostly of the r–R and S series; soft opisthogastric cuticle with at most 7 pairs of setae................................................................................... 4 4. Dorsal shield with setae j3 and z4 very long, longer than diagonal interval j3–z4 and twice as long as transverse interval j3–j3, and with setae J4 absent; metapodal plates ovoid, nearly as wide as long; dorsal palpfemoral seta d extending beyond apex of palpgenu; leg I genu and tibia each with 10 or 11 setae, third ventral (av-2) absent.................... B. vorontsovi n. sp. - Dorsal shield with setae j3 and z4 moderately long, shorter than interval j3–z4 and no longer than interval j3–j3, and with minute setae J4 present; metapodal plates slender/baculiform, three times as long as wide; dorsal palpfemoral seta d extending to about mid-level of palpgenu; leg I genu and tibia each with 12 setae, third ventral (av-2) present....... B. hospitabilis n. sp.

Lindquist, Evert E., Oconnor, Barry M., Shaw, Matthew D., Sidorchuk, Ekaterina A. (2020): Review of the genera Berlesia Canestrini, 1884, and Katydiseius Fain & Lukoschus 1983, the subfamily Katydiseiinae Fain & Lukoschus, 1983, and their family group relationships (Acari: Mesostigmata: Gamasina), with description of three new species parasitic on gryllacridid crickets (Orthoptera). Zootaxa 4857 (1): 5-70, DOI: https://doi.org/10.11646/zootaxa.4857.1.4

Key to genera and species of Katydiseiinae, adult females 1. Dorsal vertex setae j1 absent (not to be confused with j3, present slightly behind level of poroids id-1); sternal shield reduced, bearing only setae st1, without endopodal extensions between bases of legs I–II and II–III; anal shield with paranal setae inserted at mid-level of anus; subcapitulum with hypostome prolonged anteriorly by two paramedian processes reaching level of palptibia and enveloping extremities of internal malae and labrum; coxa of legs I–IV with all setae setiform; genua of legs I-II-III with 8-6-6 setae, each lacking al-2, pl-2......................................................................................................... Genus Katydiseius, monobasic, K. nadchatrami Fain & Lukoschus, 1983 - Dorsal vertex setae j1 present (inserted slightly anteromediad poroids id-1, closely anterior to j3); sternal shield well developed, bearing setae st1, st2, with endopodal extensions between bases of legs I–II and II–III; anal shield with paranal setae inserted at level of, or behind, posterior margin of anus; subcapitulum with sclerotized extensions of hypostome extended only as corniculi to level of palptrochanter, but with flexible mouthpart structures (salivary styli, internal malae, labrum) extending freely to level of palpgenu; coxa of legs I–III and sometimes IV each with one seta strongly bulbiform basally; genua of legs I-II-III with minimally 11-10-9 setae, each with al-2, pl-2... Genus Berlesia Canestrini, 1884............................... 2 2. Coxal seta of leg IV conspicuously bulbiform, similar to one of coxal setae on each of legs I–III and subcapitular setae pc; anal shield lateral margins nearly parallel; tarsi II–IV each with one of apical setae distinctively longer than adjacent setae and as long as entire pretarsus.............................................................. B. rapax Canestrini, 1884 - Coxal seta of leg IV setiform, in contrast to a bulbiform coxal seta on each of legs I–III and subcapitulum; anal shield subrhomboidal, lateral margins angular; tarsi II–IV each with apical setae similar in length, none as long as entire pretarsus........ 3 3. Vertex setae j1 moderately long, z1 present; dorsal shield with mid–dorsal setae (j4–j6, z5, J1–J3) also moderately long (J2 present), e.g., j4 nearly as long as interval j4–j5; dorsal shield flanked by about 25 pairs of setae, mostly of the hypertrichous r–R and S series; soft opisthogastric cuticle also hypertrichous, with about 15 pairs of setae........... B. multisetosa n. sp. - Vertex setae j1 short, z1 absent; dorsal shield with mid-dorsal setae also short (J2 absent), e.g., j4 ca one-fourth as long as interval j4–j5; dorsal shield flanked by at most 15 pairs of setae mostly of the r–R and S series; soft opisthogastric cuticle with at most 7 pairs of setae................................................................................... 4 4. Dorsal shield with setae j3 and z4 very long, longer than diagonal interval j3–z4 and twice as long as transverse interval j3–j3, and with setae J4 absent; metapodal plates ovoid, nearly as wide as long; dorsal palpfemoral seta d extending beyond apex of palpgenu; leg I genu and tibia each with 10 or 11 setae, third ventral (av-2) absent.................... B. vorontsovi n. sp. - Dorsal shield with setae j3 and z4 moderately long, shorter than interval j3–z4 and no longer than interval j3–j3, and with minute setae J4 present; metapodal plates slender/baculiform, three times as long as wide; dorsal palpfemoral seta d extending to about mid-level of palpgenu; leg I genu and tibia each with 12 setae, third ventral (av-2) present....... B. hospitabilis n. sp., Published as part of Lindquist, Evert E., Oconnor, Barry M., Shaw, Matthew D. & Sidorchuk, Ekaterina A., 2020, Review of the genera Berlesia Canestrini, 1884, and Katydiseius Fain & Lukoschus 1983, the subfamily Katydiseiinae Fain & Lukoschus, 1983, and their family group relationships (Acari: Mesostigmata: Gamasina), with description of three new species parasitic on gryllacridid crickets (Orthoptera), pp. 5-70 in Zootaxa 4857 (1) on page 44, DOI: 10.11646/zootaxa.4857.1.4, http://zenodo.org/record/4396474, {"references":["Fain, A. & Lukoschus, F. S. (1983) Katydiseius nadchatrami n. g., n. sp. (Acari: Otopheidomenidae) from the tracheae of a Malaysian katydid Chloracris brullei Pictet & Saussure, 1892 (Orthoptera, Pseudophyllidae). International Journal of Acarology, 9, 173 - 178. https: // doi. org / 10.1080 / 01647958308683333","Canestrini, G. (1884) Acari nuovi o poco noti. Atti del Reale Istituto Veneto di Scienze, Lettere ed Arti, Serie VI, Tomo II, 693 - 724, pls. VI-IX."]}

Berlesia and Katydiseius as separate genera The above descriptions, presenting morphological similarities as well as distinctions between Berlesia and Katydiseius, may indicate that the latter taxon is a derivative subset of the former, such that it could be considered as either a subgenus or as fully congeneric with it. A critical missing piece of the puzzle is any knowledge of the male of the monotypic Katydiseius. However, Fain & Lukoschus (1983) described the type species as having a free-living larva with chelicerae nearly as fully developed as in the protonymph, whose chelicerae in turn were not indicated to differ from the deutonymph. A fully developmental ontogenetic sequence of instars may in turn suggest other differences in behavior, including copulation. The life history of species of Berlesia is compacted/shortened, by way of nymphipary and by male copulation with females not yet eclosed from their deutonymphal skins, which may be facilitated by an extremely elongated spermatodactyl of the male. Based on the presence of a free-living and apparently functional larva, such life history compaction is not indicated in Katydiseius nadchatrami. Indeed, the parasitic association of Berlesia species with gryllacridid crickets may differ considerably from that of K. nadchatrami with pseudophylline tettigoniid crickets, such that perhaps males with extremely elongated spermatodactyls adapted for mating with pharate females should not be assumed for the latter. Note also that the deutonymph of K. nadchatrami has an entire dorsal shield like that of the female, rather than the unusual, apomorphic condition of that shield being divided in the deutonymph but entire in the adult of Berlesia species. Unlike that of Berlesia spp., the deutonymph of K. nadchatrami was not noted to have more distinctly developed claws than in other instars; this is notable as it may indicate a difference in phoresy, or even more interestingly in copulatory behavior (see ontogenetic considerations, below). The bulbiform subcapitular and coxal setae in female Berlesia are also apomorphic with respect to the simple form of those setae in Katydiseius. Each genus is diagnosed above with unique apomorphies, with the presence of other, shared apomorphies from which we infer a sister relationship., Published as part of Lindquist, Evert E., Oconnor, Barry M., Shaw, Matthew D. & Sidorchuk, Ekaterina A., 2020, Review of the genera Berlesia Canestrini, 1884, and Katydiseius Fain & Lukoschus 1983, the subfamily Katydiseiinae Fain & Lukoschus, 1983, and their family group relationships (Acari: Mesostigmata: Gamasina), with description of three new species parasitic on gryllacridid crickets (Orthoptera), pp. 5-70 in Zootaxa 4857 (1) on page 35, DOI: 10.11646/zootaxa.4857.1.4, http://zenodo.org/record/4396474, {"references":["Fain, A. & Lukoschus, F. S. (1983) Katydiseius nadchatrami n. g., n. sp. (Acari: Otopheidomenidae) from the tracheae of a Malaysian katydid Chloracris brullei Pictet & Saussure, 1892 (Orthoptera, Pseudophyllidae). International Journal of Acarology, 9, 173 - 178. https: // doi. org / 10.1080 / 01647958308683333"]}

Berlesia multisetosa new species (Figures 7–12) Diagnosis Adult: dorsal shield entirely smooth, female’s with irregular indication of lateral incisions at level of setae s6; vertex setae j1, mid-dorsals (j4–j6, z5, J1–J3) and J4 of moderate length (e.g., j4 nearly as long as interval between j4 and j5); dorsal shield setae j3 and z4 slightly longer, but shorter than diagonal interval between them, and shorter than transverse interval between j3; setae z1 and J2 present. Dorsolateral areas of idiosoma with setae of r–R and S series hypertrichous. Soft ventral cuticle hypertrichous, with about 15 pairs of opisthogastric setae. Leg I genu and tibia each with 11 setae, pd-3 and av-2 absent. Tarsus IV with seta pv-1 bluntly spine-like and asymmetrically shorter than attenuated av-1. Peritremes embellished with internal cellular structures. Description Idiosomatic dorsum. Adult female (Fig. 7A). Idiosoma 625–700 long, 412–432 wide between insertions of setae r3; dorsal shield 520–530 long, greatest width 290–320 at level between insertions of s4 (n=2), unornamented except for subdermal sigillar patterns; shield with irregularly formed lateral incisions at level of setae s6. Dorsal shield with usually 10 to 14 pairs (5 to 8 podonotal, 4 to 6 opisthonotal) of smooth setae of which any of z2, s5, J4 or Z4 asymmetrically on/off shield margin; of shield setae, j3 (60–65), z4 (62–75), s4 (62–75), s5 (60–70) of similar lengths; j4 (45–65), j5 (40–60), j6 (31–53), z5 (33–52), J1–J4 somewhat shorter but not tiny (22–40), with z4 and s4 (62–75), s5 (60–67), Z1 (57–70) and Z4 (60–72) similarly longer. Soft dorsal peripheral cuticle with setae j1 (47–52), z1 (40–43), z2 (54) anteriorly, s6 (60–65) and a hypertrichy of 12–15 r -setae (50–62) anterolaterally, and Z2 (57–65), S1–S4 (55–70), S5 (75) and hypertrichy of 10–15 R -setae (52–65) posterolaterally. Dorsal shield with complement of nine pairs of discernible pore-like structures (four podonotal, five opisthonotal), of which three pairs (one podonotal, two opisthonotal) superficially appear secretory (gland pores) and six pairs (three podonotal, three opisthonotal) non-secretory (poroids). Adult male (Fig. 9A). Idiosoma 580–612 long, 370–437 wide between insertions of setae r3; dorsal shield 520– 532 long, greatest width 337–365 at level between insertions of s6 (n=3), unornamented, with irregular but entire lateral margins, which widened to hold setae j1, z1, s3 anteriorly and Z2, S2 laterally. Dorsal shield with usually 23 to 25 pairs (14 or 15 podonotal, 9 or 10 opisthonotal) of smooth setae of which s6, Z2, Z4 often asymmetrically off shield margin; relative setal lengths much as in female, except J1–J4 shorter (20–30) compared to j4–j6 (37–57); other setal lengths j1 (35–42), j3 (58–68), z1 (25–32), z2 (55–62), z4 (60–65), s4–s6 (57–67), about eight r -setae (40–52), Z1–Z2 (60–65), Z3 (60–72), Z4–Z5 (65–77), S1–S3 (62–68), S4 (65–72), S5 (68–85), about ten R -setae (52–68). Dorsal shield with complement of eleven pairs of discernible pore-like structures (4 podonotal, 7 opisthonotal), most of which are difficult to discern whether secretory (gland pores) or non-secretory (poroids). Idiosomatic venter. Adult female (Fig. 7B). Tritosternum base 23–25 long and 23–26 wide at its base, with paired ciliate laciniae fused for about 5–7 of entire length (ca 100). Presternal area with a delineated, lightly sclerotized, streaky region without clear platelets. Sternal shield mid-length (98–100) much less than narrowest width (148–155) between legs II, faintly scabrous; sternal setae st1, st2 attenuated, of similar length (55–67). Sternal setae st3, st4 similarly attenuated (50–60), with poroid iv3 on soft cuticle. Endopodal strips rudimentary alongside coxae III–IV. Epigynal shield smooth, its rounded hyaline anterior margin reaching or slightly overlapping posterior edge of sternal shield, its lateral margins slightly widened at insertions of attenuated setae st5 (57–65), its posterior margin slightly convex; shield 205–223 long, 60–63 at narrowest width between legs IV, 77–85 at widest level at setae st5. Postgenital groove undeveloped, without elements of platelets. Opisthosomatic venter with two pairs of weak metapodal platelets, lateral pair 30–32 x 8–12, inner pair 17–20 x 7–10. Anal shield subrhomboidal, lightly lineated; shield greatest width (128) slightly less than length (132–147, including cribrum 20), cribrum width 75; postanal seta (60–70) about 1.5 longer than paranal setae (37–50) inserted in line with posterior margin of anus. Soft cuticle with ca 15 pairs of opisthogastric setae of similar lengths (43–57) flanking anal shield, JV5 not distinguishable posterolaterally; of four or five pairs of pore-like structures on soft cuticle, paranal gland pores well separated from lateral margins of anal shield. Peritrematal shield and peritreme extending barely to mid-level of coxae III; peritreme with embellished internal chambers (Fig. 7F), its length 45, including stigmatal diameter of ca 22; peritrematal shield continuous posteriorly with short exopodal strip extending partly around posterior margin of coxa IV. Exopodal elements absent alongside coxae II–IV. Spermathecal apparatus with simple, elongate (90–95) tubulus leading from solenostome between bases of legs III–IV to an unpaired sacculus (Fig. 7G). Adult male (Fig. 9B). Tritosternum formed as in female, with laciniae fused for only about one-tenth of entire length (ca 100). Presternal region with two pairs of small platelets anterior to genital opening. Sternitigenital shield length 245–255, narrowest widths 115–125 at level between coxae II, 100–107 between coxae III; posterior margin width 62–80; shield unornamented, narrowly separated from anterior edge of endopodal strips beside coxae IV; alveoli of setae st5 on corners of irregularly shaped posterior margin of shield; setae st1–st5 similarly attenuated as in female, st1 and st5 (53–67) slightly longer than st2–st4 (50–65). Paragenital poroids on soft cuticle flanking posterolateral corners of sternitigenital shield. Postgenital groove not developed. Inguinal region with five or six pairs of sigilla indicating unusual muscular development connected with deeply retractable bases of chelicerae. Anal shield similar in rhomboid form, slight ornamentation as in female, its anterior and lateral apices more irregular; shield longer (130–135, including cribrum ca 15–20 long) than wide (103–112, cribrum width 60–62); postanal seta (57–62) ca 1.5 to nearly twice as long as paranal setae (30–40). Soft cuticle with about 15 pairs of opisthogastric setae flanking anal shield; these setae gradually slightly lengthening posteriorly, those in JV1-JV2-ZV1-ZV2 area 43–48, those mid-posteriorly 45–62, and posterior-most JV5 (62–75); of five pairs of pore-like structures evident on soft cuticle, paranal gland pores well separated from lateral margins of anal shield. Short but embellished peritremes (40–45), and peritrematal shield with brief poststigmatal exopodal extension as in female (Figs 9J, 20B). Gnathosoma. Adult female. Gnathotectum anterior margin obtusely smoothly triangular, with a blunt, medial anterior projection (Figs 7H, J). Cheliceral shaft, excluding basal section, 155–158 long; fixed digit with blunt apex, vestige of pilus dentilis, and a vestigial tooth midway on paraxial face (Fig. 7E); movable digit (35) with three coarse teeth flared laterally (Figs 7D, E). Subcapitulum with 11 to 13 rows of deutosternal denticles, each similarly narrow, with a single denticle, or any row occasionally with two contiguous denticles (Fig. 7C); subcapitular setae hp1 (60–63) and slightly longer hp3 (75–80) similarly attenuated, hp2 (20–25) and capitular setae pc (22–27) similarly short, but pc strongly bulbiform basally. Corniculi somewhat hyaline, blade-like, bluntly pointed apically, moderately short (35–38), closely adjacent to similarly short tubular salivary styli (ca 37–40) and longer, slender, finely fimbriated internal malae (87–90). Palp length 150–162; palptrochanter setae dissimilar in length, v1 22–25, v2 38–52; palpus with femoral seta al-1 and genual setae al-1, al-2 blunt, not spatulate; palpfemoral seta d (30–32) slightly spine-like, shorter than femur length; palptarsal apotele two-tined. Adult male. Gnathotectum with simple, triangular anterior margin much as in female, but sides acutely angular, tapering to single median projection (Fig. 9E). Cheliceral shaft, excluding short basal section 120–125 long; fixed digit straight, short (30–42) more slender and pointed than in female; movable digit bluntly pointed beyond apex of fixed digit, reduced, amalgamated into spermatodactyl base which extends anteroventrally; spermatodactyl greatly elongated, sinuous, gradually tapering, 425–465 long, without elaborations (Figs 9C, D). Unlike female, corniculi shorter (22–25), more truncated apically, greatly exceeded by elongated salivary styli (63–70), and capitular setae pc nearly setiform, slightly bulbous basally. Form and length of other gnathosomatic structures much as in female: internal malae (120–130); setae hp1 (62–63) and hp3 (65–77) much longer than hp2 (18–20) and pc (23–25); deutosternum with 11–12 rows of mostly contiguously doubled denticles (Fig. 9F). Palp length 160–165; palpfemoral seta d (25–30) similar in form as in female. Legs. Adult female (Figs 8 A–H). Legs I (425–450) nearly as long as legs IV (460–470), and about 0.8 as long as dorsal shield; other leg lengths (excluding pretarsi): II (375–380), III (385–400). Leg I pretarsus (37–38) to apex of pulvillus, tarsus (115–125) about 1.7–2.1 longer than each of the similarly long femur (67–70), genu (53–63), and tibia (60–65). Legs II–IV with tarsus/tibia length ratios about 1.8–2.3. Complement of setae on leg segments holotrichous for femora II-III-IV (11-6-6) as presented for the genus, but femur I with 11 setae, lacking pv-2; chaetotactic formulae for genua I-II-III-IV: 11 (2-3/1, 2/1-2), 11 (2-3/1, 2/1-2), 9 (2-2/1, 2/1-1), 10 (2-2/1, 3/1-1); tibiae: 11 (2-3/1, 2/1-2), 10 (2-2/1, 2/1-2), 8 (2-1/1, 2/1-1), 10 (2-1/1, 3/1-2); genu I and tibia I without pd-3 and av-2; tibia III without pl-2. Tarsus I with seta md (55–60) longer than other setae on basal half of segment (Fig. 8B); dorsal subapical cluster of sensory structures with a cluster of three, tightly adjacent, tiny, pointed setae surrounded by three solenidia, one of which biramous (Fig. 8C). Tarsi II–IV with a finely serrated ventroapical ridge basal to a short, rounded process (Figs. 8E, H), and with dorsoapical setae (d-1) (12–15) nearly half as long as pretarsus to base of claws (ca 25). Coxal seta av on leg I and pv on legs II, III strongly bulbiform basally, like subcapitular pc; coxa IV seta v setiform. Trochanters I–IV with basal seta pv-2 attenuated (50–55), and III–IV ad slightly spinelike. Tarsus IV with apical seta pv-1 asymmetrically shorter, bluntly spine-like, than attenuated av-1 (Fig. 8H). Legs I–IV each with particular setae enlarged, spine-like: I Fe ad-1 (65), pd-1 (45); I Ge ad-1 (50–60), pd-1 (55–58); II Fe ad-1 (57–63), pd-1 (47–48); II Ge ad-1 (50–52), pd- 1 (47–48); II Ti pd-1 not differentiated; III Fe ad-1 (52–54); III Ge ad-1 (57–62), pd-1 (50–52); III Ti ad (42), pd-1 (30); IV Fe ad-1 (57–62), pv not differentiated; IV Ge ad-1 (66–68), pd-1 (62–65); IV Ti ad-1 (53–60), pd-1 (55–60). Relative lengths of other leg setae as shown in Figs 8A, D, F–H. Adult male (Figs 10 A–D). Legs similar in relative dimensions to those of female; lengths I 415–430, II 360– 378, III 375–400, IV 440–470. Legs I and IV without dimorphism in thickness of segments, legs II and III thickersegmented than in female. Leg I with form and relative sizes of setae similar to those of female, except femur I with 12 setae, including pv-2, and with av slightly bulbiform. Legs I–IV retain attenuated form of basal seta pv-2 of trochanter. Legs II–IV with particular setae dimorphically modified: leg II Fe av spine-like; Ge av, Ti av, and tarsus mv, av-2, av-1, pv-1 bulbiform; leg III coxal pv simple; Tr ad markedly arched; Fe al less spine-like, and tarsus mv, av-2, pv-2, av-1, pv-1 bulbiform; leg IV Tr ad markedly arched, av, pv-1 thickened, more spine-like; Fe pd, pl more spine-like; Ge pd-2, pd-3 and Ti pl-1 larger, more spine-like; tarsus av-2, pv-2, av-1, pv-1 bulbiform. Lengths of spine-like setae similarly enlarged as in female: I Fe ad-1 (48–62), pd-1 (37–45); I Ge ad-1 (45–52), pd-1 (52–60); II Fe ad-1 (55–65), pd-1 (42–49); II Ge ad-1 (38–45), pd-1 (50–57); II Ti pd-1 not differentiated; III Fe ad-1 (40–50); III Ge ad-1 (43–53), pd-1 (47–50); III Ti ad (35), pd-1 (27); IV Fe ad-1 (57–70), pv not differentiated; IV Ge ad-1 (67–70), pd-1 (67–75); IV Ti ad-1 (55–65), pd-1 (60–67). Deutonymph. Idiosomatic dorsum (Fig. 11A). Idiosoma 475–500 long, 362–365 wide between insertions of setae r3 (n=3); dorsal shields weakly sclerotized, unornamented, faintly divided by transverse strip of soft cuticle between setae j6 and J1. Podonotal shield 210–225 long, ca 215–225 wide at its greatest width at level of setae s5, with ca 12 pairs of setae (j1, j3–j6, z1, z2, z4. z5, s4–s6); opisthonotal shield ca 175–195 long and 175–205 at widest level near setae Z2, with nine pairs of setae (J1–J4, Z1–Z5). Soft cuticle with ca 25 pairs of dorsolateral setae, including uncertain homologies of the s -and S -series among hypertrichous elements of the marginal r–R -series (Fig. 11A). Form and relative lengths of dorsal setae more homologous than in adult, many of them slightly barbed as shown in Fig. 11A; other than z1 clearly shortest (18–28), setal lengths range from j1 (35–40) to Z5 longest (65). Peritreme similarly short (length 37–38, including stigma diameter 15–17) and embellished as in adult (Fig. 11D). Idiosomatic venter (Fig. 11B). Tritosternum similar in form to that of adult, with laciniae fused for only about 5–10 of entire length (87–92 excluding base). Presternal area devoid of ornamentation. Intercoxal shield 182–200 long, weakly sclerotized, bearing five pairs of setae and three pairs of poroids, seta st5 sometimes barely off shield edge; setae st1–st4 (43–50) slightly shorter than st5 (52–57); paragenital poroids on soft cuticle flanking posterior edge of shield. Opisthogaster with oboval, smooth anal shield; length 95–100, greatest width 75–80; shield with postanal seta (45–50) about 1.3 longer and stouter than paranal setae (35–37). Soft cuticle with hypertrichous complement and relative lengths of setae flanking anal shield much as in adult, with ca 15 pairs of slender opisthogastric setae flanked by 5–10 pairs of thicker R -marginal setae of similar lengths (40–47), the more posterolateral elements of them thicker, blunter. Gnathosoma. Gnathotectum, chelicerae, palpal structures, and subcapitulum much as in female, except setal pair pc simple. Cheliceral shaft, excluding basal section, 132–138 long; movable digit 28–35 long. Subcapitulum with 11 to 13 rows of deutosternal denticles formed as in female; subcapitular setae hp1 (45–50) and slightly longer hp3 (52–58) similarly attenuated, hp2 (13–15) and capitular setae pc (15–18) similarly short. Corniculi (25–30) formed as in female, exceeded by elongated tubular salivary styli (45–50) and even longer, slender, finely fimbriated internal malae (70–75). Palpus length 127–133; palptrochanter setae v1 12–15, v2 30–34; palpgenual seta d (22–23) slightly spine-like. Legs. Leg lengths, excluding pretarsi, I (325–350), II (278–312), III (287–315), IV (335–360). Leg setation with normal deutonymphal complement of setae as given for adult female, with occasional variabilities. Tarsi II to IV with ventroapical process and pretarsal structures formed as in female; apical dorsosetal processes (d-1) relatively short (9–10, about 0.6 as long as pretarsus to base of claws 13–15) as in female; form and relative lengths of setae much as in female, with particular setae enlarged, spine-like: I Fe ad-1 (45–50), pd-1 (30–33); I Ge ad-1 (38–45), pd-1 (37–43); II Fe ad-1 (50–53), pd-1 (32–33); II Ge ad-1 (35–38), pd-1 (34–35); III Fe ad-1 (45–50); III Ge ad-1 (42–43), pd-1 (37–40); IV Fe ad-1 (47–52); IV Ge ad-1 (45–20), pd-1 (40–48); IV Ti ad-1 (40–47), pd-1 (36–42). Tarsus IV with asymmetry in lengths of setae (v-1) as in female. Claws of all legs strongly formed (length ca 13), projecting freely from pulvillus (as in Fig. 19E), in contrast to adult and protonymph. Protonymph. Idiosomatic dorsum (Fig. 12A). Idiosoma 370 long, 268 wide at widest level of setae r3, r5 (n = 1), with weak, Published as part of Lindquist, Evert E., Oconnor, Barry M., Shaw, Matthew D. & Sidorchuk, Ekaterina A., 2020, Review of the genera Berlesia Canestrini, 1884, and Katydiseius Fain & Lukoschus 1983, the subfamily Katydiseiinae Fain & Lukoschus, 1983, and their family group relationships (Acari: Mesostigmata: Gamasina), with description of three new species parasitic on gryllacridid crickets (Orthoptera), pp. 5-70 in Zootaxa 4857 (1) on pages 18-26, DOI: 10.11646/zootaxa.4857.1.4, http://zenodo.org/record/4396474

Berlesia hospitabilis new species (Figures 1–6, 18, 19, 21) Berlesia, undescribed species, host of Podapolipus gryllacridi Lindquist & Sidorchuk, 2019. Diagnosis Adult: dorsal shield faintly reticulated posteriorly, smooth elsewhere, with irregular indication of lateral incisions at level of setae s6; setae j1, mid-dorsals (j4–j6, z5, J1–J3) and J4 very short (e.g., j4 about one-fourth as long as interval between j4 and j5); dorsal shield setae j3 and z4 moderately long, but shorter than diagonal interval between them, and shorter than transverse interval between j3; setae z1 and J2 absent. Dorsolateral margins of idiosoma with setae of r-R and S series sparse, not hypertrichous. Soft ventral cuticle with 7 and 6 pairs of opisthogastric setae on female and male, respectively. Leg I genu and tibia each with 12 setae, pd-3 absent, av-2 present. Tarsus IV with seta pv-1 asymmetrically shorter but similar in attenuated form to av-1. Peritremes simple, not elaborated with internal cellular structure. Description Adult: Idiosomatic dorsum. Adult female (Fig. 1A). Idiosoma 550–730 long, 288–377 wide between insertions of setae r3; dorsal shield 507–550 long, greatest width 250–287 at level between insertions of s4 (n=4), faintly ornamented with subdermal reticula outlined by sparse puncta (mostly not evident in micrographs of surface structure); shield with irregular indication of lateral incisions at level of setae s6. Dorsal shield with usually 10 or 11 pairs (7 podonotal, 3 or 4 opisthonotal) of smooth setae of which s4 or Z5 asymmetrically on/off shield margin and J4 present or absent; setae j3 (58–75), z4 (67–85), s4 (70–85), Z5 (70–75) of similarly long lengths; other shield setae (j4–j6, z5, J1, J3, J4) tiny (5–12), occasionally asymmetrically rudimentary. Soft dorsal peripheral cuticle with tiny setae j1 (10–15) anteriorly, z2 (40–50), s6 (77–83) and r2, r3, r5 (50–70) anterolaterally, and Z2 (70–83), Z4 (80–85), S2 (55–67), S3 (62–68), S4 (72–83), S5 (75–90) and R1 (48–62) posterolaterally. Dorsal shield with complement of 15 pairs of discernible pore-like structures (5 podonotal, 10 opisthonotal), of which five pairs (two podonotal, three opisthonotal) superficially appear secretory (gland pores) and 10 pairs (three podonotal, seven opisthonotal) non-secretory (poroids). Adult male (Fig. 3A). Idiosoma 566–570 long, 322–330 wide between insertions of setae r3; dorsal shield 512–524 long, greatest width 350–358 at level between insertions of r5 (n=2), faintly ornamented and with irregular lateral incisions as in female, margins widened to hold setae j1 anteriorly and r2–r5, Z2, Z4, S2–S4 on lateral subcutaneous extensions. Dorsal shield with 21 or 22 pairs (13 podonotal, 8 or 9 opisthonotal) of smooth setae of which S3 asymmetrically present or absent; setae j1 (10–12), j4–j6, z5 (7–9), J1, J3, J4 (5–7) tiny as in female, occasionally asymmetrically rudimentary; other setal lengths j3 (30–37), z2 (14–18), z4 (47–50), s4 (65–71), s6 (55–60), r2 (25–28), r3 (28–30), r5 (43–45), Z2 (70–72), Z4 (62–67), Z5 (62–63), S2 (44–50), S3 (50–53), S4 (80–82). Soft dorsal peripheral cuticle with setae R1 (37–39) laterally and S5 (68–78) posteriorly. Dorsal shield with complement of ca 22 pairs of discernible pore-like structures (9 podonotal, 13 opisthonotal), many of which are difficult to discern whether secretory (gland pores) or non-secretory (poroids). Idiosomatic venter. Adult female (Fig. 1B). Tritosternum base 27–32 long and 22–27 wide at its base, with paired, shortly ciliated laciniae fused for about 8–15 of entire length (150–162) (Fig. 1F). Presternal region without platelets. Sternal shield mid-length (75–82) much less than narrowest width (125–145) between legs II, unornamented; sternal setae st1, st2 attenuated, of similar length (53–62). Sternal setae st3, st4 similarly attenuated (45–57), with poroid iv3 on soft cuticle. Endopodal strips absent alongside coxae III–IV. Epigynal shield smooth, its rounded hyaline anterior margin reaching or slightly overlapping posterior edge of sternal shield, its lateral margins slightly widened at insertions of attenuated setae st5 (55–62), its posterior margin slightly convex; shield 198–220 long, 55–63 at narrowest width between legs IV, 68–75 at widest level at setae st5. Postgenital groove with elements of one pair of platelets widely separated at level behind paragenital poroids. Opisthosomatic venter with two pairs of small baculiform platelets, longitudinal metapodal platelets 27–34 long, 5–7 wide and nearly transversal postgenital platelets. Anal shield subrhomboidal, unornamented; shield greatest width (118–135) slightly less than length (128–160, including cribrum 12–15), cribrum width 43–50; postanal seta (52–65) about 1.5 longer than paranal setae (35–40) inserted in line with posterior margin of anus. Soft cuticle with seven pairs of opisthogastric setae flanking anal shield, JV1 (40–43), JV2 (38–45), ZV1 (35–40), ZV2 (32–40) anteriorly and JV3 (32–38), ZV3 (27–32), JV5 (52–60) posterolaterally; of three pairs of pore-like structures on soft cuticle, paranal gland pores well separated from lateral margins of anal shield. Peritrematal shield and peritreme extending barely to mid-level of coxae III; peritreme structure simple, without embellished internal chambers (Fig. 1J), its length 52–55, including stigmatal diameter of ca 15; peritrematal shield continuous posteriorly with short exopodal strip extending partly around posterior margin of coxa IV. Exopodal elements absent alongside coxae II–IV. Spermathecal apparatus with simple, elongate (65–75) tubulus leading from solenostome in soft cuticle between bases of legs III–IV to an unpaired sacculus (Fig. 1K). Adult male (Fig. 3C). Tritosternum formed as in female, with laciniae fused for only about one-tenth of entire length (125–140). Presternal region with a pair of platelets preceding genital opening. Sternitigenital shield length 255, narrowest widths 115 at level between coxae II, 100 between coxae III; posterior margin width 63; shield unornamented, narrowly connected to anterior edge of endopodal strips beside coxae IV; alveoli of setae st5 barely contiguous with rounded posterolateral corners of shield; setae st1–st5 attenuated as in female, st4 (26–30) clearly shorter than others (42–50). Paragenital poroids on soft cuticle flanking posterolateral corners of sternitigenital shield. Postgenital groove not developed. Inguinal region with five or six pairs of sigilla indicating unusual muscular development connected with deeply retractable bases of chelicerae.Anal shield similar in rhomboid form, lack of ornamentation as in female, its anterior and lateral apices more irregular; shield longer (136–138, including cribrum ca 12 long) than wide (118–120, cribrum width 45–50); postanal seta (62–65) nearly twice as long as paranal setae (28–37). Soft cuticle with six pairs of opisthogastric setae flanking anal shield, ZV1 absent (present on female); other than JV5, these setae much shorter than in female; JV1 (23–30), JV2 (18–25), JV3 (29–32), ZV2 and ZV3 tiny (7–8), JV5 (49–57); six pairs of pore-like structures evident on soft cuticle, paranal gland pores well separated from lateral margins of anal shield. Peritremes, and peritrematal shield with brief poststigmatal exopodal extension, as in female. Gnathosoma. Adult female. Gnathotectum anterior margin with a single smooth, medial anterior projection extending from a convex margin which smooth or slightly serrate at base of projection (Figs 1G, H). Cheliceral shaft, excluding basal section, 175–178 long; fixed digit with truncated apex bearing two or three tiny serrae, alveolar remnant of pilus dentilis, and minute subapical tooth; movable digit (33–35) with three coarse teeth flared laterally (Figs 1C, D). Subcapitulum with 11 to 13 rows of deutosternal denticles, each similarly narrow, with a single denticle (Fig. 1E); subcapitular setae hp1 (76–80) and hp3 (82–90) similar in attenuated lengths, hp2 (22–27) and capitular setae pc (20) similarly short, but pc strongly bulbiform basally. Corniculi somewhat hyaline, blade-like, narrowly rounded apically, moderately long (50–52), but exceeded by elongated tubular salivary stylets (70–80) and even longer, slender, finely fimbriated internal malae (80–90). Palp length 155–162; palptrochanter setae dissimilar in length, v1 53–63, v2 17–20; palpus with femoral seta al-1 and genual setae al-1, al-2 blunt, not spatulate; palpfemoral seta d (31–35) slightly spine-like, nearly as long as femur; palptarsal apotele two-tined (Figs 2 L–N). Adult male. Gnathotectum with simple anterior margin as in female, but sides more angular, leading to single median projection (Fig. 3E). Cheliceral shaft, excluding short basal section ca 130 long, including fixed digit projection; fixed digit straight, more slender and pointed than in female; movable digit reduced, except for greatly elongated, sinuous spermatodactyl 375–430 long, its base extending anteroventrally beyond apex of fixed digit, gradually narrowing apically, without elaborations (Figs 3B, 18A, 19B). Unlike female, corniculi shorter (25–26), more truncated apically, greatly exceeded by elongated salivary styli (96–100), and capitular setae pc setiform (Fig. 3D). Form and length of other gnathosomatic structures much as in female: internal malae (115–134); setae hp1 (ca 50) and hp3 (70–76) much longer than hp2 and pc (16–18); deutosternum with 11–12 rows of single or rarely doubled denticles. Palp length 150–152. Form and length of palpal setae as in female. Legs. Adult female (Figs 2 A–K). Legs I (462–500) nearly as long as legs IV (495–537), and about 0.9 as long as dorsal shield; other leg lengths (excluding pretarsi): II (395–427), III (405–440). Leg I pretarsus (40–45) to apex of pulvillus, tarsus (125–137) about 1.7–2.1 longer than each of the similarly long femur (73–80), genu (60–65), and tibia (65–70). Legs II–IV with tarsus/tibia length ratios about 1.8–2.1. Complement of setae on leg segments holotrichous for femora I-II-III-IV (12-11-6-6) as presented for the genus; chaetotactic formulae for genua I-II-III-IV: 12 (2-3/2, 2/1-2), 11 (2-3/1, 2/1-2), 9 (2-2/1, 2/1-1), 10 (2-2/1, 3/1-1); tibiae: 12 (2-3/2, 2/1-2), 10 (2-2/1, 2/1-2), 8 (2-1/1, 2/1-1), 10 (2-1/1, 3/1-2); genu I and tibia I without pd-3; tibia III without pl-2. Tarsus I with seta md (65–70) longer than other setae on basal half of segment (Fig. 2G); dorsal subapical cluster of sensory structures with a row of four, tightly adjacent, tiny, pointed setae surrounded by three solenidia, one of which biramous (Figs 2J, K). Tarsi II–IV with a finely serrated ventroapical ridge basal to a short, rounded process (Figs 2E, F), and with setae (d-1) (10–15) nearly half as long as pretarsus to base of claws (22–30). Coxal seta av on leg I and pv on legs II, III strongly bulbiform like subcapitular pc (Figs 2 A–C); coxa IV seta v setiform (Fig. 2D). Trochanters II–IV with basal seta pv-2 attenuated (50–55), and III–IV ad slightly spinelike. Femur IV with seta v large, spine-like (40). Tarsus IV with apical seta pv-1 asymmetrically shorter, but not thicker than av-1 (Fig. 2E). Legs I–IV each with particular dorsal setae enlarged, spinelike: I Fe ad-1 (55–62), pd-1 (37–42); I Ge ad-1 (50–52), pd-1 (55–57); II Fe ad-1 (60–65), pd- 1 (55–60); II Ge ad-1 (34–35), pd-1 (55–60); II Ti pd-1 (29); III Fe ad-1 (45–52); III Ge ad-1 (55–60), pd-1 (60–63); III Ti ad (55), pd-1 (61); IV Fe ad-1 (50–66); IV Ge ad-1 (62–70), pd-1 (65–70); IV Ti ad-1 (52–55), pd-1 (57–60). Relative lengths of other leg setae as shown in Figures 2 A–G. Adult male (Figs 4 A–E). Legs similar in relative dimensions to those of female. Lengths I 440–452, II 367–400, III 392–415, IV 494–495. Legs without noticeable male dimorphism in thickness of segments, with same chaetotaxy as female; legs I–III with form and relative sizes of setae somewhat similar to those of female, including attenuated form of basal seta pv-2 of trochanter, but particular dorsal spine-like setae generally shorter and ventral setae more spine-like on femora, genua, tibiae. Other than trochanter, leg IV segments with nearly all dorsal and ventral setae spine-like; tibia and basal half of tarsus with pd- setae formed as a column of five blunt spine-like setae; femur IV with spine-like seta v distinctly smaller (20) than on female. Lengths of elongated dorsal spine-like setae: I Fe ad-1 (56), pd-1 (20); I Ge ad-1 (27–32), pd-1 (34–37); II Fe ad-1 (65), pd-1 (45); II Ge ad-1 (23), pd-1 (44); III Fe ad-1 (23); III Ge ad-1 (52), pd-1 (41); III Ti ad (37), pd-1 (33); IV Fe ad-1 (67); IV Ge ad-1 (75), pd-1 (50); IV Ti ad-1 (60). Tarsi II and III with setae (v-1), and mv bulbiform; tarsus IV with setae (d-2) and basitarsal (d-3) thickened, spine-like. Deutonymph. Idiosomatic dorsum (Fig. 5A). Idiosoma ca 520 long, ca 290 wide between insertions of setae r3 (n=2); dorsal shields weakly sclerotized, unornamented, completely divided by transverse strip of soft cuticle between setae j6 and J1. Podonotal shield 245–285 long, ca 260–290 wide at its greatest width at level of setae r5, with 10 or 11 pairs of (j1, j3–j6, z2, z4, z5, s4, s6, sometimes r5); opisthonotal shield ca 175–240 long and 210–225 at widest level anteriad setae Z2, with five or six pairs of setae (J1 J3, J4, Z2, Z5, sometimes Z3). Soft cuticle with two or three pairs of podonotal setae (r2, r3, sometimes r5) and five or six pairs of opisthonotal setae (S2–S5, R1, sometimes Z3). Form and relative lengths of dorsal setae as in adult female, all smooth. Setal lengths on podonotum: j1 (11), j3 (35–40), j4 (10–12), j5–j6 (6–7), z2 (17–20), z4 (ca 60), z5 (5–7), s4 (ca 60), s6 (57), r2 (ca 30), r3 (37), r5 (32–35); on opisthonotum: J1 (5–7), J3–J4 (3–5), Z2, Z4 (50–60), Z5 (50–60), S2–S3 (40–42), S4 (45–48), S5 (68–70), R1 (25). Peritreme short (50 including stigma), extending anteriorly to mid-level of coxae III, similar in form to adult. Idiosomatic venter. Tritosternum similar in form to that of adult, with laciniae fused for about 22–29 of entire length (98–112 excluding base). Presternal area devoid of ornamentation. Intercoxal area length 210–225, weakly sclerotized (lateral margins not clearly discerned), bearing five pairs of setae and three pairs of poroids (Fig. 5C); setae st1 (50), st2–st3 (42–43) longer than st4 (27) and st5 (32–35); paragenital poroid on soft cuticle flanking setae st5. Opisthogaster with oboval, smooth anal shield (Fig. 5C); length ca 90, greatest width ca 70; shield with postanal seta (49) stouter and about 1.4 longer than paranal setae (35). Soft opisthogastric integument with seven pairs of setae; JV1, JV2 (30–32), ZV1 (8–10), ZV2 (15), JV3, ZV3 (10–12), JV5 thicker (33–36). Paranal gland pores on anal shield laterally. Gnathosoma. Gnathotectum elongate-angular as in male (Fig. 5B); chelicerae, palpal structures, and subcapitulum much as in female, except setal pair pc simple. Cheliceral shaft, excluding basal section, 145 long; movable digit 28–33 long. Subcapitulum with 11 or 12 rows of deutosternal denticles formed as in female; subcapitular setae hp1 (60–69) and slightly longer hp3 (65–70) similarly attenuated, hp2 (18–22) and capitular setae pc (20–21) similarly short. Corniculi (55) with slightly narrower apices than in female, exceeded by elongated tubular salivary styli (80) and even longer, slender, finely fimbriated internal malae (ca 100). Palpus length 135; palptrochanter setae v1 10, v2 28; palpgenual seta d (10) slightly spine-like. Legs. Leg lengths, excluding pretarsi, I (430–490), II (346–372), III (385–414), IV (450–481). Leg setation with normal deutonymphal complement of setae as given for adult female. Tarsi II to IV with ventroapical process and pretarsal structures formed as in female; dorsoapical setal processes (d-1) relatively short (12–13, about 0.6 as long as pretarsus to base of claws 20–25) as in female; form and relative lengths of setae much as in female, with particular setae enlarged, spine-like: I Fe ad-1 (45–48), pd-1 (31–33); I Ge ad-1 (40–43), pd-1 (47–50); II Fe ad-1 (57), pd-1 (38–42); II Ge ad-1 (36), pd-1 (46–50); III Fe ad-1 (53); III Ge ad-1 (46–50), pd-1 (40); IV Fe ad-1 (56); IV Ge ad-1 (67), pd-1 (61); IV Ti ad-1 (ca 50), pd-1 (ca 50). Tarsus IV with asymmetry in lengths of setae (v-1) as in female. Claws of all legs strongly formed (length ca 15), projecting freely from pulvillus (Fig. 19E), in contrast to adult and protonymph. Protonymph. Idiosomatic dorsum. Idiosoma 405 long, 250 wide at widest level of setae r5 (n=1), with weakly sclerotized dorsal shields (Fig. 6A); podonotal shield 208 long and 225 at level between setae s4

Berlesia vorontsovi new species (Figures 13–17) Diagnosis Adult: dorsal shield faintly reticulated posteriorly, smooth elsewhere, with irregular indication of lateral incisions at level of setae s6; setae j1, mid-dorsals (j4–j6, z5, J1, J3) very short; dorsal shield setae j3 and z4 relatively long, about as long or slightly longer than diagonal interval between them, and longer than transverse interval between j3; setae z1, J2 and J4 absent. Dorsolateral regions of idiosoma with setae of r -R -and S -series sparse, not hypertrichous. Soft ventral cuticle with 6 and 4 pairs of opisthogastric setae on female and male, respectively. Leg I genu and tibia each with 10–11 setae, pd-3, av-2 and sometimes ad-3 absent. Tarsus IV with seta pv-1 asymmetrically shorter, but similar in attenuated form to av-1. Peritremes embellished with internal cellular structure. Description Adult: Idiosomatic dorsum. Adult female (Fig. 13A). Idiosoma 741–791 long, 432–451 wide between insertions of setae r3; dorsal shield 580 long, greatest width 306–324 at level between insertions of s4 (n=4), ornamentation not apparent, shield with irregular indication of lateral incisions at level of setae s6 and irregular indentations more posteriorly. Dorsal shield with usually 9 or 10 pairs (7 or 8 podonotal, 2 opisthonotal) of smooth setae of which j1 may be asymmetrically on/off shield margin and J4 always absent; setae j3 (131–140), z4 (133–137), s4 (135–140) of similarly long lengths; other shield setae (j4–j6, z5, J1, J3) tiny (ca 5), occasionally asymmetrically vestigial. Soft dorsal peripheral cuticle with tiny setae j1 (10–12) anteriorly (occasionally on shield), z2 (110–119), s6 (110–126) and r2, r3, r5 (103–105) anterolaterally, and Z2 (108–119), Z4 (133–142), S2 (85–89), S3 (60–89), S4 (96–140), S5 (128) and R1 (78–92) posterolaterally, Z5 (92–101) slightly off shield posteriorly. Dorsal shield pore-like structures difficult to discern (maximum observed complement shown in Fig. 13A). Adult male (Fig. 15A). Idiosoma 672 long, 409–432 wide between insertions of setae r3; dorsal shield 570 long, greatest width 321–335 at level between insertions of r5 (n=3), without distinct ornamentation and with shallow lateral incisions, setae j1 off shield anteriorly, z2 and r2–r5, Z2, Z4, S2–S4 on lateral subcutaneous extensions. Dorsal shield with 9 or 10 pairs (7 podonotal, 2–3 opisthonotal) of smooth setae; setae j1 (7–8), j4–j6, z5 (3), J1, J3 (3) occasionally asymmetrically vestigial or absent, J4 absent; other setal lengths j3 (92–108), z2 (41–46), z4 (92–117), s4 (105–108), s6 (96–103), r2 (64–76), r3 (71–76), r5 (73–78), Z2 (108–115), Z4 (108–115), Z5 (92–96), S2 (60–64), S3 (53–76), S4 (92–110), S5 (119–133), R1 (55–73). Dorsal shield with complement of ca 15 pairs of discernible pore-like structures (4 podonotal, 11 opisthonotal), many of which are difficult to discern whether secretory (gland pores) or non-secretory (poroids). Idiosomatic venter. Adult female (Fig. 13B). Tritosternum base 25–30 long and 28–34 wide at its base, with paired ciliated laciniae fused for about 8% of entire length (92–103) (Fig. 13F). Presternal region without platelets and lineation. Sternal shield mid-length (96–98) much less than narrowest width (158–172) between legs II, unornamented; sternal setae st1 (96–110), st2 (69–87), st3 (87–92), st4 (62–69) attenuated, with poroid iv3 on soft cuticle. Endopodal strips absent alongside coxae III–IV. Epigynal shield smooth, its rounded hyaline anterior margin reaching or slightly overlapping posterior edge of sternal shield, its lateral margins widened at insertions of attenuated setae st5 (73–82), its posterior margin slightly convex, occasionally indented or with fragmented platelets posteriorly; shield 185–218 long, 46–48 at narrowest width between legs IV, 68–76 at widest level at setae st5. Opisthosomatic venter with several irregularly spaced platelets widely separated at level between metapodal plates, and with two pairs of small ovoid metapodal platelets, anterior pair 14–16 long, 5–7 wide, posterior pair 25–30 long, 14–21 wide. Anal shield subrhomboidal, weakly reticulate posteriorly, without paranal gland pores; shield greatest width (126–130) slightly less than length (137–149, including cribrum), cribrum U-shaped, width 55–64; postanal seta (69–73) longer than paranal setae (37–53) inserted in line with posterior margin of anus. Soft cuticle with maximum of 6 pairs of opisthogastric setae flanking anal shield, JV1 (53–60), JV2 (46–48), ZV2 (44–46) anteriorly and JV3 (39–48), ZV3 (46–53), JV5 (60–66) posterolaterally, ZV1 and JV4 setae always absent, JV2, JV3, JV5, ZV2 sometimes unilaterally or bilaterally absent; one pair of pore-like structures observed on soft cuticle. Peritrematal shield and peritreme extending to posterior margin of coxae III; peritreme structure with embellished internal chambers (Fig. 13E), its length 60–62, including stigmatal diameter of ca 11; peritrematal shield continuous posteriorly with short exopodal strip extending partly around posterior margin of coxa IV. Exopodal elements absent alongside coxae II–IV. Spermathecal apparatus with simple, elongate (80–119) tubulus leading from solenostome in soft cuticle between bases of legs III–IV to an unpaired sacculus (Fig. 13G). Adult male (Fig. 15B). Tritosternum formed as in female, with laciniae fused for only about one-tenth of entire length (98–100). Presternal region with a pair of weakly developed platelets abutting genital opening, one or both platelets sometimes fused with sternitigenital shield. Sternitigenital shield length 252–263, narrowest widths 124–134 at level between coxae II, 124–131 between coxae III; posterior margin width 62–69; shield with scalelike ornamentation anterior to setae st4, narrowly connected to anterior edge of endopodal strips beside coxae IV; alveoli of setae st5 barely contiguous with rounded posterolateral corners of shield; setae st1–st5 attenuated as in female, st4 (34–37) clearly shorter than others, st1 (76), st2 (66–73), st3 (66–69), st5 (57–69). Paragenital poroids on soft cuticle flanking posterolateral corners of sternitigenital shield. Postgenital groove not developed. Inguinal region with six or seven pairs of small platelets or sigilla as in B. hospitabilis. Anal shield rhomboid with a variously formed anterior extension, weak ornamentation present laterally; shield longer (144–156, including cribrum ca 34–60 long) than wide (112–119, cribrum width 64–73); postanal seta (73–78) about 1.5 times as long as paranal setae (48–50). Soft cuticle with four pairs of opisthogastric setae flanking anal shield; other than JV5, these setae shorter than in female, and ZV1, ZV3, JV3, JV4 absent; JV1 (46–48), JV2 (34–41), JV5 (37–48), ZV2 (25–30); pore-like structures on soft cuticle difficult to discern. Peritremes, and peritrematal shield with brief poststigmatal exopodal extension, as in female. Gnathosoma. Adult female. Gnathotectum anterior margin acutely triangular with very weakly serrate edges (Figs 13H). Cheliceral shaft, excluding basal section, 160–165 long; fixed digit with truncated apex and minute subapical tooth, other structures not observed; movable digit (23–25) with three coarse teeth flared laterally (Figs 13C, D). Subcapitulum with 11 relatively narrow rows of deutosternal denticles, varying somewhat in width and consisting from a single denticle to multidentate (Fig. 13J); subcapitular setae hp1 (62–82) and hp3 (82–103) attenuated, hp2 (30–44) and capitular setae pc (39–41) similarly short, but pc with strongly bulbiform base and laterally positioned flagellum. Corniculi somewhat hyaline, blade-like, narrowly rounded apically (39–44), but exceeded by elongated tubular salivary styli (60–64) and slender, finely fimbriated internal malae (57–62). Palp length 183–188; palptrochanter setae dissimilar in length, v1 46–78, v2 41–46; palpus with femoral seta al-1 acicular, genual setae al-1 spatulate, al-2 acicular; palpfemoral seta d (60–64) spine-like, typically longer than femur (Fig. 13J); palptarsal apotele two-tined (Fig. 13K). Adult male. Gnathotectum slightly serrate anteriorly as in female, sides almost parallel posteriorly, leading to single median projection (Fig. 15D). Cheliceral shaft, excluding short basal section 92–94 long, including fixed digit projection; fixed digit straight, more slender and pointed than in female; movable digit reduced, except for greatly elongated, sinuous spermatodactyl 404–444 long, its base extending anteroventrally beyond apex of fixed digit, gradually narrowing distally, abruptly hooked apically (Fig. 15C). Unlike female, corniculi shorter (25–30), more truncated apically, greatly exceeded by salivary styli; capitular setae pc with bulbous base as in female (Fig. 15E). Form and length of other gnathosomatic structures much as in female: attenuated salivary styli (44), internal malae (89–108); setae hp1 (57–69) and hp3 (78–92) much longer than hp2 and pc (21); deutosternum with 11–12 rows of denticles, basal 4 rows multidentate, apical rows with 1–2 denticles (if two, then one is always distinctly larger). Palp (Figs. 15F, G) length 179–185. Form and lengths of palpal setae as in female. Legs. Adult female (Figs 14 A–F). Legs I (561–589) distinctly shorter than legs IV (644–672), and only slightly shorter than dorsal shield; other leg lengths (excluding pretarsi): II (524–552), III (543–561). Leg I pretarsus (65–89) to apex of pulvillus, tarsus (169–176) about 1.9–2.1 longer than each of the similarly long femur (85–92), genu (80–85), and tibia (87–92). Legs II–IV with tarsus/tibia length ratios about 1.7–2.1. Complement of setae on leg segments holotrichous for femora I-II-III-IV (12-11-6-6) as presented for the genus; chaetotactic formulae for genua I-II-III-IV: 10–11 (2-2 or 3/1, 2/1-2), 11 (2-3/1, 2/1-2), 9 (2-2/1, 2/1-1), 10 (2-2/1, 3/1-1); tibiae: 10–11 (2-2 or 3/1, 2/1-2), 9–10 (2-1 or 2/1, 2/1-2), 7–8 (1 or 2-1/1, 2/1-1), 10 (2-1/1, 3/1-2); genu I and tibia I without pd-3, av-2, sometimes without ad-3; tibia II sometimes without ad-2; tibia III without pl-2, sometimes without al-2. Tarsus I with seta md (103–110) longer than other setae on basal half of segment (Fig. 14B); dorsal subapical cluster of sensory structures with a row of three tightly adjacent, tiny, pointed setae surrounded by three solenidia, one of which biramous (Fig. 14B). Tarsi II–IV with ventroapical ridge, serrations not discernible, short, rounded process not discernible, and with setae (d-1) (ca 14–16) less than half as long as pretarsus to base of claws (34–41). Coxal seta av on leg I and pv on legs II, III strongly bulbiform basally like subcapitular pc (Figs 14A, C, D); coxa IV seta v setiform (Fig. 14F). Trochanters II–IV with basal seta pv-2 attenuated (71–94), and III–IV ad slightly spinelike. Femur IV with seta v large, somewhat spine-like (49). Tarsus IV with apical seta pv-1 asymmetrically shorter, but not thicker than av-1 (Fig. 14E). Legs I–IV each with particular dorsal setae enlarged, spinelike: I Fe ad-1 (103– 126), pd-1 (50–85); I Ge ad-1 (87–92), pd-1 (112–128), ad-2 (48–50); II Fe ad-1 (115–121), pd-1 (80–82); II Ge ad-1 (69–76), pd-1 (119–121), ad-2 (69–71); II Ti pd-1 (69–80); III Fe pd-1 (115–124); III Ge ad-1 (92–117), pd-1 (92–110); III Ti ad (85–96), pd-1 (69–73); IV Fe ad-1 (114–135); IV Ge ad-1 (124–133), pd-1 (92–121); IV Ti ad-1 (92–105), pd-1 (92–103). Note: all long setae on leg IV in the holotype are stunted on one side, their measurements not included above. Relative lengths of other leg setae as shown in Figures 14 A–F. Adult male (Figs 16 A–E). Legs similar in relative dimensions to those of female, including legs IV distinctively longest. Lengths I 478–488, II 423–442, III 442–460, IV 561–598. Legs without noticeable male dimorphism in thickness of segments, with same setation as female; legs I–III with form and relative sizes of setae similar to those of female, including dorsal spine-like setae, but ventral setae more spine-like on femora, genua, tibiae. Basal trochanter seta pv-2 slender, attenuated as on female, though less so on legs III and IV, which also have pv-1 shorter, thicker than on female. Leg IV segments with nearly all dorsal and ventral setae spine-like; genu, tibia and basal half of tarsus with pd- setae normally formed (not as a column of blunt spine-like setae as in male B. hospitabilis); femur IV with seta v slightly smaller (37–45) than on female. Lengths of elongated dorsal spine-like setae: I Fe ad-1 (92–96), pd-1 (34–41); I Ge ad-1 (71–87), pd-1 (92–94); II Fe ad-1 (92–110), pd-1 (66–73); II Ge ad-1 (48–60), pd-1 (92–96); III Fe ad-1 (94–98); III Ge ad-1 (80–89), pd-1 (79–89); III Ti ad (50–64), pd-1 (48–55); IV Fe ad-1 (92–137); IV Ge ad-1 (105–115), pd-1 (98–110); IV Ti ad-1 (80–92), pd-1 (85–94). Tarsi II and III with setae (v-1), and mv bulbiform; tarsus IV with setae (d-2) and basitarsal (d-3) not thickened, spine-like. Deutonymph. Unknown. Protonymph. Idiosomatic dorsum. Idiosoma 495–543 long, 381–437 wide at widest level of setae r5 (n = 4), with weakly sclerotized dorsal shields (Fig. 17A); podonotal shield 294–312 long and 308–321 wide at level between setae s4, strongly ornamented with a network of lines, with 12 pairs of setae (j3–j6, z2, z4, z5, s4, s6, r2, r 3,r5 ); pygidial shield typically narrowly fused with anterior lateral sclerites, rarely slightly separated, without obvious ornamentation, 188–197 long and 266–280 at widest level between setae R1, with five pairs of setae (J1, J3, Z2, Z3, Z5), J4 absent. Soft cuticle with only one pair of podonotal setae (j1) and two pairs of opisthonotal setae (S2, R1); S3-S5 absent. Form and relative lengths of dorsal setae as in adult, all smooth. Setal lengths on podonotum: j1 (6–7), j3 (69–76), j4–j6 (6), z2 (30–37), z4 (78–87), z5 (6), s4 (80–87), s6 (55–57), r2 (18–25), r3 (30–37), r5 (32–37); on opisthonotum: J1 (7), J3 (7), Z2 (62–64), Z3 (44–46), Z5 (62–69), S2 (32), R1 (27–32). Peritreme (46 including stigma), extending anteriorly to posterior edge of coxae III. Idiosomatic venter. Tritosternum formed much as in adult, with laciniae fused for 11–16 of total length (73–89). Presternal area devoid of ornamentation. Sternal shield with weak but distinct outline, strongly narrowing between setae st2 and st3, length 144, with setae st1–st2 (48–60) and poroids iv1, iv2; setae st3 (48–53) on soft cuticle; setae st5 and paragenital poroids not discernible (Fig. 17B). Anal shield smooth, subrhomboid, broadest between paranal and postanal setae, length 92–94 including cribrum, width 78–80 (cribrum strip width 39–44); postanal seta (39–50) only slightly longer than paranal setae (39–44). Soft opisthogastric integument with four pairs of setae: JV1 (34–41), JV2 (18–25), ZV2 (16–21), JV5 longer (48–55), ZV2 unilaterally absent in 3 of 4 specimens. Gnathosoma. Gnathotectum with anterior margin acutely triangular, tapering to single median projection, apparently smooth (Fig. 17D). Cheliceral shaft, excluding short basal section 119–124 long; fixed digit straight, more bluntly pointed than in female; movable digit (18–21) formed much as in female, with three teeth flared laterally (Fig. 17C). Subcapitulum with 13 rows of deutosternal denticles, 5 basal rows slightly wider, multidentate, remaining rows uni-or bidentate; subcapitular setae hp1 and hp3 similar in attenuated lengths (32–48), hp2 (21–22) and capitular setae pc (26–30) similar in short simple form. Corniculi hyaline, blade-like, bluntly pointed apically, moderately short (24–25), greatly exceeded by elongated tubular salivary styli (46). Palpus length 137–142, with normal protonymphal complement of setae; single palptrochanter seta 9–11; palpfemoral seta d (30–32) slightly thickened. Legs. Leg lengths, excluding pretarsi, I 206, II 183, III 181–183, IV 211–215. Leg setation of protonymph typical of general holotrichous pattern presented for free-living Gamasina by Evans (1963): coxae, 2-2-2-1; trochanters, 4-4-4-4; femora, 10 (2 2/1 2/1 2)-8 (1 2/1 2/1 1)-5 (1 2/1 1/0 0)-4 (1 2/0 1/0 0); genua, 8 (1 2/1 2/1 1)-6 (1 2/0 2/0 1)-6 (1 2/0 2/0 1)-5 (1 2/0 2/0 0); tibiae, 8 (1 2/1 2/1 1)-7 (1 1/1 2/1 1)-7 (1 1/1 2/1 1)-7 (1 1/1 2/1 1). Legs I–IV each with particular setae enlarged, spine-like, with similar pattern as for adults. Tarsi II–IV with ventroapical process, pretarsal structures and weakly developed claws formed as in female. Tarsus IV with asymmetry in lengths of setae (v-1) as in female. Larva. No material available (adult females anticipated to be nymphiparous). Type material. Type specimens: all collected by B.M. OConnor (BMOC 84-1800-039) from one adult female c, Published as part of Lindquist, Evert E., Oconnor, Barry M., Shaw, Matthew D. & Sidorchuk, Ekaterina A., 2020, Review of the genera Berlesia Canestrini, 1884, and Katydiseius Fain & Lukoschus 1983, the subfamily Katydiseiinae Fain & Lukoschus, 1983, and their family group relationships (Acari: Mesostigmata: Gamasina), with description of three new species parasitic on gryllacridid crickets (Orthoptera), pp. 5-70 in Zootaxa 4857 (1) on pages 26-33, DOI: 10.11646/zootaxa.4857.1.4, http://zenodo.org/record/4396474

Mucroseius insolitus sp. nov. (Figures 1 – 5) Type material Holotype female (slide No 24 – 07 – 2017 /1)ı Russiaı Altai Republicı Mayminsky Districtı vicinity of Manzherokı 51°49 ʹ Nı 85°48 ʹ Eı from the metathoracic spiracular atria of the Black pine sawyer beetleı Monochamus galloprovincialis (Olivierı 1795) ı collected from the surface of felled Scots pine (Pinus sylvestris)ı 24 July 2017ı V. A. Trach coll. Paratypes: three femalesı with same date as holotype; two femalesı Russiaı Tyumen Provinceı Nizhnetavdinsky Regionı vicinity of lake Kuchakı 57°21 ʹ Nı 66°03 ʹ Eı from the metathoracic spiracular atria of the Black pine sawyer beetle collected from trunk of living of Scots pineı 31 July 2018ı V. A. Khaustov coll.; one femaleı Austriaı Tirolı under elytra of the Small white-marmorated longhorned beetleı Monochamus sutor (Linnaeusı 1758) ı Kraatz coll. Type deposition Holotype at ZMUO; paratypes at ONUDZı TUMZı and CNCI. Diagnosis Dorsal shield smooth over most of surface but lightly reticulated along margins; with 39 – 41 pairs of setae (j1–j6, z1–z6, s1–s6, r2–r6, J1–J5, Z1–Z5, S1–S5, R1–R3) (setae r5, r6, R2, R3 sometimes asymmetrically off shield on one side); length of dorsal shield setae 35 – 50 mostly. Sternal shield with posterior margin weakly concaveı arched to level of bases of setae st3. Epigynal shield reticula outline about 10 elongate cells between bases of setae st5. Epistome clearly triramous. Distal flangelike process (mucro) of the fixed chela absent. Tarsus IV with two erect macrosetaeı ad2 and pd2; tibia III with eight setae (seta pl2 absent). Description Female (n = 4) Idiosomal dorsum (Figures 1 and 5 (a)). Dorsal shield oval; 444 (456 – 502) long and 248 (251 – 260) maximum width at r4 level; smooth over most of surface but lightly reticulated along marginsı midlateral incisions on lateral margins absent; with 39 – 41 pairs of setae (usually j1–j6, z1–z6, s1–s6, r2–r6, J1–J5, Z1–Z5, S1–S5, R 1– R 3 ı but any of j2, z1, z2, s2, J5 sometimes asymmetrically lacking on one sideı or any of r5, r6, R 2, R 3 sometimes asymmetrically off shield on one side) and 23 pairs of distinguishable pore-like structures (seven gland pores and 16 poroids). Soft cuticle with setae R 4– R 6 (usually visible in ventrolateral regions) and poroids Rp. Setae Z5 sparsely barbedı other dorsal setae simpleı needle-like; measurements of setae: j1 33 – 34 (36 – 44)ı j2 35 (36 – 46)ı j3 40 – 41 (42 – 46)ı j4 38 – 41 (40 – 50)ı j5 37 (38 – 48)ı j6 40 – 42 (42 – 55)ı z1 22 (23 – 29)ı z2 38 (46 – 50)ı z3 42 – 46 (48 – 55)ı z4 50 (53 – 67)ı z5 47 (48 – 61)ı z6 38 – 44 (44 – 55)ı s1 33 (32 – 38)ı s2 35 – 38 (36 – 40)ı s3 44 – 51 (46 – 63)ı s4 56 – 57 (48 – 69)ı s5 48 – 56 (59 – 69)ı s6 44 – 50 (44 – 61)ı r2 42 – 43 (40 – 50)ı r3 50 – 51 (46 – 61)ı r4 41 – 44 (50 – 65)ı r5 52 – 53 (50 – 61)ı r6 36 – 38 (40 – 51)ı J1 40 – 41 (40 – 55)ı J2 42 – 43 (42 – 50)ı J3 39 (38 – 48)ı J4 39 – 40 (40 – 57)ı J5 17 (19 – 27)ı Z1 43 – 44 (38 – 55)ı Z2 44 – 49 (40 – 53)ı Z3 39 – 40 (36 – 53)ı Z4 37 – 39 (36 – 57)ı Z5 72 – 73 (63 – 95)ı S1 39 – 41 (42 – 55)ı S2 42 – 44 (44 – 63)ı S3 41 – 43 (33 – 34)ı S4 42 (42 – 59)ı S5 31 – 35 (34 – 50)ı R 1 32 – 34 (36 – 46)ı R 2 36 – 37 (36 – 46)ı R 3 35 (38 – 48)ı R 4 36 – 35 (36 – 55)ı R 5 35 – 37 (36 – 44)ı R 6 32 – 35 (36 – 46)ı UR1–UR3 32 – 35 (34 – 48). Idiosomal venter (Figures 2 and 5 (b)). Tritosternum with trapezoidal baseı 15 (15 – 19) longı 15 (13 – 15) wide at baseı laciniae piloseı fused for about fourth of total length 17 (17 – 19)ı their free parts 45 (44 – 48) long. Pre-sternal area transversely lineateı without platelets. Sternal shield fused with endopodal strips of coxae I/II and coxae II/III; 108 (103 – 109) long along midlineı 172 (168 – 189) wide at level of endopodal projections between coxae I/IIı 157 (158 – 164) wide at level of endopodal projections between coxae II/IIIı 95 (97 – 91) wide at narrowest interval about mid-level of coxae II; with three pairs of setae (st1–st3) (37 – 41 (36 – 46) long)ı two pairs of poroids (iv1, iv2) and gland pores gvb; poroids iv1 positioned posteriad seta st1 ı poroids iv2 positioned between setae st2 and st3 ı gland pores gvb positioned on apex of endopodal projections between coxae I and II; posterior margin weakly concaveı arched to level of bases of setae st3; reticulated over all of surface except posteriorly where overlapped by hyaline flap of epigynal shield. Setae st4 (38 – 40 (38 – 42) long) located on tiny metasternal platelets with greatest dimension only about twice diameter of setal alveolus; poroids iv3 absent. Anterior margin of epigynal shield broadly roundedı overlapping posterior sternal shield areaı extending to level of setae st2 ı posterior margin of shield slightly rounded; epigynal shield reticula outline about 10 elongate cells between bases of setae st5; epigynal shield 180 (181 – 185) longı with greatest width of hyaline anterior part 102 (105 – 118)ı greatest width of posterior part 77 (76 – 84); length st5 34 (34 – 38); poroids iv5 placed on soft cuticle closely beside shield. Postgenital platelets absent. Free endopodal plates strongly formed between coxae III and IV. Anal shield ovalı slightly reticulateı with anterior margin roundedı usually indented mediallyı posterior margin nearly truncate; 91 (90 – 105) long and 76 (76 – 82) wide; anus located slightly anterior to mid-level of shieldı anal opening 33 (34 – 40) long; cribrum well-developed; one pair of gland pores present (gv3); length of para-anal setae 21 (21 – 23)ı length of post-anal seta 47 (48 – 61). Posteriad coxae IV two pairs of elongate metapodal platelets present; the larger platelet 21 (21 – 23) longı 7 (6 – 8) wide; the smaller platelet 11 (6 – 11) longı 5 (3 – 5) wide. Soft cuticle around anal shield with 12 – 13 pairs of setae (JV1–JV5, ZV1–ZV5 ı flanked by two or three pairs of UR setae) and five pairs of distinguishable poroids. All ventral setae simple; length of setae of JV-, ZV - and UR -series 27 – 36 (27 – 44) (excluding JV5 60 (61 – 84)). Exopodal platelets of coxae II – IIIı coxae III – IV and platelet enveloping coxa IV posteriorly fused into a single stripı bearing gland pores gv2 at posteromedial extremity. Peritrematal shields fused with dorsal shield at level of setae s1; with four pairs of distinguishable pore-like structures (poroids ip2, ip3 ı and gland pores gp2, gp3); peritreme extending slightly anteriad of setae s1. Spermathecal apparatus not distinguishable. Gnathosoma (Figure 3 (a–f) and 5(c)). Epistome triramousı with medial tine slightly serrated apicallyı longer than lateral tinesı these serrated along their outer margins (Figure 3 (a – c) and 5(c)). Subcapitulum 99 (101 – 109) wide at widest level. Deutosternum with seven rows of denticlesı rows 1 – 5 connected; anterior four rows each with three to eight denticlesı 5th row widenedı concave mediallyı with 10 – 15 denticlesı 6th row freeı widenedı with 10 – 17 denticlesı 7th row freeı with four to eight denticles (Figure 3 (d)). Hypostome with four pairs of simple setae; palpcoxal seta (pc) 19 – 20 (17 – 21)ı hp1 28 – 29 (27 – 32)ı hp2 20 (15 – 16)ı hp3 35 – 37 (34 – 38). Corniculi 29 – 34 longı 8 – 11 wideı symmetricalı horn-likeı sclerotisedı with paraxial process near base; internal malae slenderı extending to apex of corniculi; salivary styli with blunt apices not reaching tips of corniculi. Palp length from trochanter to tarsus apex 148 – 150 (145 – 158); setal formula normal: 2 – 5 – 6 – 15 – 16; palpfemoral seta al ı palpgenual setae al1 and al2 spatulateı palptarsal apotele 2-tined (Figure 3 (dı e)). Second cheliceral segment length 88 – 90 (90 – 101)ı movable digit length 36 (36 – 42). Fixed cheliceral digit with 17 (16 – 20) similar small teethı offset subapical tooth in addition to apical hookı and membranous lobeı distal process absentı antiaxial lyrifissure and dorsal seta distinct; movable digit with three teeth in addition to apical hookı and ventral projection (Figure 3 (f)). Legs (Figures 4 (a–d) and 5(d)). Lengths: I 410 – 420 (419 – 446)ı II 332 – 339 (353 – 409)ı III 364 – 373 (372 – 428)ı IV 469 – 476 (476 – 530). Leg chaetotactic formulae normal for genus: leg I: coxa 2ı trochanter 6 (1 0/1 1/2 1)ı femur 12 (2 3/1 2/2 2)ı genu 13 (2 3/2 3/1 2)ı tibia 13 (2 3/2 3/ 1 2); leg II: coxa 2ı trochanter 5 (1 0/1 0/2 1)ı femur 11 (2 3/1 2/2 1)ı genu 11 (2 3/1 2/1 2)ı tibia 10 (2 2/1 2/1 2); leg III: coxa 2ı trochanter 5 (1 0/1 0/2 1)ı femur 6 (1 2/1 1/0 1)ı genu 9 (2 2/1 2/1 1)ı tibia 8 (2 1/1 2/1 1) lacking pl2; leg IV: coxa 1ı trochanter 5 (1 0/1 0/2 1)ı femur 6 (1 2/1 1/0 1)ı genu 9 (2 2/1 3/0 1)ı tibia 10 (2 1/1 3/1 2). Tarsi II-IV: 18 (3 3/2 1/1 3/2 3). Tarsus I with erect middorsal macroseta 48 (53 – 61) long; tarsus IV with two erect macrosetaeı ad2 70 – 71 (74 – 101) and pd2 79 – 85 (99 – 122); all setae smooth. Etymology. The specific name of the new species is derived from the Latin word insolitus ı meaning ‘unusual‘ ı and refers to the absence of the distal flangelike process of the fixed chela of its female. Differential diagnosis Mucroseius insolitus sp. nov. differs from all known congeners by the absence of the distal flangelike process of the female fixed chela. The following keyı modified from that of Lindquist and Wu (1991) ı further distinguishes the new species from others in the genus. Key to adult females of Mucroseius (based on Lindquist and Wu 1991) 1. Tarsus IV with 2 macrosetaeı pd2 and ad2 ı distinguished by nearly perpendicular orientation and greater length from other tarsal setae........................................................ 2 – Tarsus IV with 1 macrosetaı only pd2 readily distinguished as such............................ 7 2. Fixed chela with broadly rounded distal flange; tibia III with 9 setae (pl2 present); dorsal shield ornamented with conspicuous little ridges and reticula over entire surface................................................................................................................................ M. aciculatus – Fixed chela with slenderı bluntly pointed distal flangeı or flange absent; tibia III with 8 setae (pl2 absent); dorsal shield smooth or reticulateı not carinulate...................... 3 3. Fixed chela without distal flange............................................................ M. insolitus sp. nov. – Fixed chela with slender distal flange.......................................................................................... 4 4. Epistome weakly triramousı its median process broadı untapered and conspicuously serratedı and its lateral processes weakly projected and conspicuously serrated; dorsal shield with setae s4 nearly as long or longer than caudal setae Z5; leg IV with anterior macroseta subequallyı or within 15 μm as long ası posterior macroseta.. M. disparisetus – Epistome clearly triramousı its medial process tapering to a slightly serrated or smooth pointı and its lateral processes well projected and sparsely serrated; dorsal shield with setae s4 usually not more than 0.8 as long as caudal setae Z5 (some M. algonquian excepted); leg IV with anterior macroseta clearly (by 20 – 30 μm) shorter than posterior macroseta........................................................................................ 5 5. Dorsal shield smooth over most of surfaceı with 7 – 9 pairs of marginal setae (r2–r6, R 1– R 2 ı sometimes R 3– R 4) inserted on its lateral edges; epigynal shield reticula outline 12 – 16 finely elongate cells between bases of setae st5; sternal shield with posterior margin deeply excavated to level of setae st2............................... M. nipponensis – Dorsal shield smooth or reticulated over most of surfaceı with only 2 or 3 pairs of marginal setae (r2–r3 ı sometimes r4) inserted on its lateral edges; epigynal shield reticula outline 9 – 11 elongate cells between bases of setae st5; sternal shield with posterior margin deeply excavated or not................................................................................ 6 6. Dorsal shield conspicuously reticulate over most or all of surface (sometimes smooth medially from setae j4 to j6 ı and from J1 to J4); sternal shield with posterior margin often deeply excavated to level of setae st2 or poroids iv2 ı occasionally this margin remaining weakly evident at level of setae st3.................................................. M. algonquian – Dorsal shield mostly unornamentedı smooth over most of surface (lightly reticulated along anterolateral marginsı and sometimes midlaterally between setae j6–J2 and z6–Z2 and caudally); sternal shield with posterior margin gently concave to level of setae st3........................................................................................................................ M. monochami 7. Dorsal shield strongly ornamentedı squamateı over entire surface; 8 or 9 pairs of marginal setae (r2–r6, R 1– R 3 ı sometimes R 4) inserted on lateral edges of dorsal shield; sternal shield with posterior margin deeply excavated to level of setae st2............................ M. squamosus – Dorsal shield unornamented over most of surface; 3 – 6 pairs of marginal setae (r2–r4 ı sometimes r5–r6 ı rarely R 1 or R 2) inserted on lateral edges of dorsal shield; sternal shield with posterior margin gently concave to level of setae st3................................ 8 8. Anal shield with anterior margin flattened medially; dorsal shield usually lacking one or both setae of pair z3 ı and with setae J1 and J2 extending slightly beyond bases of J2 and J3 ı respectively.................................................................................................. M. californicus – Anal shield with anterior margin smoothly rounded medially; dorsal shield with both setae of pair z3 presentı and with setae J1 and J2 barely or not quite extending to bases of J2 and J3 ı respectively............................................................................ M. mexicanus Remarks Table 1 presents a list of species of the genus Mucroseius with their distribution and hosts. Key to adult females of Mucroseius (based on Lindquist and Wu 1991) 1. Tarsus IV with 2 macrosetaeı pd2 and ad2 ı distinguished by nearly perpendicular orientation and greater length from other tarsal setae........................................................ 2 – Tarsus IV with 1 macrosetaı only pd2 readily distinguished as such............................ 7 2. Fixed chela with broadly rounded distal flange; tibia III with 9 setae (pl2 present); dorsal shield ornamented with conspicuous little ridges and reticula over entire surface................................................................................................................................ M. aciculatus

Key to adult females of Mucroseius (based on Lindquist and Wu 1991) 1. Tarsus IV with 2 macrosetaeı pd2 and ad2 ı distinguished by nearly perpendicular orientation and greater length from other tarsal setae........................................................ 2 ��� Tarsus IV with 1 macrosetaı only pd2 readily distinguished as such............................ 7 2. Fixed chela with broadly rounded distal flange; tibia III with 9 setae (pl2 present); dorsal shield ornamented with conspicuous little ridges and reticula over entire surface................................................................................................................................ M. aciculatus ��� Fixed chela with slenderı bluntly pointed distal flangeı or flange absent; tibia III with 8 setae (pl2 absent); dorsal shield smooth or reticulateı not carinulate...................... 3 3. Fixed chela without distal flange............................................................ M. insolitus sp. nov. ��� Fixed chela with slender distal flange.......................................................................................... 4 4. Epistome weakly triramousı its median process broadı untapered and conspicuously serratedı and its lateral processes weakly projected and conspicuously serrated; dorsal shield with setae s4 nearly as long or longer than caudal setae Z5; leg IV with anterior macroseta subequallyı or within 15 ��m as long ası posterior macroseta.. M. disparisetus ��� Epistome clearly triramousı its medial process tapering to a slightly serrated or smooth pointı and its lateral processes well projected and sparsely serrated; dorsal shield with setae s4 usually not more than 0.8 as long as caudal setae Z5 (some M. algonquian excepted); leg IV with anterior macroseta clearly (by 20 ��� 30 ��m) shorter than posterior macroseta........................................................................................ 5 5. Dorsal shield smooth over most of surfaceı with 7 ��� 9 pairs of marginal setae (r2���r6, R 1��� R 2 ı sometimes R 3��� R 4) inserted on its lateral edges; epigynal shield reticula outline 12 ��� 16 finely elongate cells between bases of setae st5; sternal shield with posterior margin deeply excavated to level of setae st2............................... M. nipponensis ��� Dorsal shield smooth or reticulated over most of surfaceı with only 2 or 3 pairs of marginal setae (r2���r3 ı sometimes r4) inserted on its lateral edges; epigynal shield reticula outline 9 ��� 11 elongate cells between bases of setae st5; sternal shield with posterior margin deeply excavated or not................................................................................ 6 6. Dorsal shield conspicuously reticulate over most or all of surface (sometimes smooth medially from setae j4 to j6 ı and from J1 to J4); sternal shield with posterior margin often deeply excavated to level of setae st2 or poroids iv2 ı occasionally this margin remaining weakly evident at level of setae st3.................................................. M. algonquian ��� Dorsal shield mostly unornamentedı smooth over most of surface (lightly reticulated along anterolateral marginsı and sometimes midlaterally between setae j6���J2 and z6���Z2 and caudally); sternal shield with posterior margin gently concave to level of setae st3........................................................................................................................ M. monochami 7. Dorsal shield strongly ornamentedı squamateı over entire surface; 8 or 9 pairs of marginal setae (r2���r6, R 1��� R 3 ı sometimes R 4) inserted on lateral edges of dorsal shield; sternal shield with posterior margin deeply excavated to level of setae st2............................ M. squamosus ��� Dorsal shield unornamented over most of surface; 3 ��� 6 pairs of marginal setae (r2���r4 ı sometimes r5���r6 ı rarely R 1 or R 2) inserted on lateral edges of dorsal shield; sternal shield with posterior margin gently concave to level of setae st3................................ 8 8. Anal shield with anterior margin flattened medially; dorsal shield usually lacking one or both setae of pair z3 ı and with setae J1 and J2 extending slightly beyond bases of J2 and J3 ı respectively.................................................................................................. M. californicus ��� Anal shield with anterior margin smoothly rounded medially; dorsal shield with both setae of pair z3 presentı and with setae J1 and J2 barely or not quite extending to bases of J2 and J3 ı respectively............................................................................ M. mexicanus, Published as part of Trach, Viacheslav A., Khaustov, Alexander A. & Lindquist, Evert E., 2019, A new unique species of Mucroseius Lindquist�� 1962 (Acari: Mesostigmata: Melicharidae) mites associated with sawyer beetles (Cerambycidae: Monochamus Dejean�� 1821) from the Palaearctic Region, pp. 761-778 in Journal of Natural History 53 (13) on pages 767-769, DOI: 10.1080/00222933.2019.1606954, http://zenodo.org/record/3675600, {"references":["Lindquist EEi Wu KW. 1991. Review of mites of the genus Mucroseius (Acari: Mesostigmata: Ascidae) associated with sawyer beetles (Cerambycidae: Monochamus and Mecynippus) and pine wood nematodes [Aphelenchoididae: Bursaphelenchus xylophilus (Steiner and Buhrer) Nickle] i with descriptions of six new species from Japan and North Americai and notes on their previous misidentification. Can Entomol. 123: 875 - 927."]}

Trach, Viacheslav A., Khaustov, Alexander A., Lindquist, Evert E. (2019): A new unique species of Mucroseius Lindquistı 1962 (Acari: Mesostigmata: Melicharidae) mites associated with sawyer beetles (Cerambycidae: Monochamus Dejeanı 1821) from the Palaearctic Region. Journal of Natural History 53 (13): 761-778, DOI: 10.1080/00222933.2019.1606954

Mucroseius insolitus sp. nov. (Figures 1 ��� 5) Type material Holotype female (slide No 24 ��� 07 ��� 2017 /1)ı Russiaı Altai Republicı Mayminsky Districtı vicinity of Manzherokı 51��49 �� Nı 85��48 �� Eı from the metathoracic spiracular atria of the Black pine sawyer beetleı Monochamus galloprovincialis (Olivierı 1795) ı collected from the surface of felled Scots pine (Pinus sylvestris)ı 24 July 2017ı V. A. Trach coll. Paratypes: three femalesı with same date as holotype; two femalesı Russiaı Tyumen Provinceı Nizhnetavdinsky Regionı vicinity of lake Kuchakı 57��21 �� Nı 66��03 �� Eı from the metathoracic spiracular atria of the Black pine sawyer beetle collected from trunk of living of Scots pineı 31 July 2018ı V. A. Khaustov coll.; one femaleı Austriaı Tirolı under elytra of the Small white-marmorated longhorned beetleı Monochamus sutor (Linnaeusı 1758) ı Kraatz coll. Type deposition Holotype at ZMUO; paratypes at ONUDZı TUMZı and CNCI. Diagnosis Dorsal shield smooth over most of surface but lightly reticulated along margins; with 39 ��� 41 pairs of setae (j1���j6, z1���z6, s1���s6, r2���r6, J1���J5, Z1���Z5, S1���S5, R1���R3) (setae r5, r6, R2, R3 sometimes asymmetrically off shield on one side); length of dorsal shield setae 35 ��� 50 mostly. Sternal shield with posterior margin weakly concaveı arched to level of bases of setae st3. Epigynal shield reticula outline about 10 elongate cells between bases of setae st5. Epistome clearly triramous. Distal flangelike process (mucro) of the fixed chela absent. Tarsus IV with two erect macrosetaeı ad2 and pd2; tibia III with eight setae (seta pl2 absent). Description Female (n = 4) Idiosomal dorsum (Figures 1 and 5 (a)). Dorsal shield oval; 444 (456 ��� 502) long and 248 (251 ��� 260) maximum width at r4 level; smooth over most of surface but lightly reticulated along marginsı midlateral incisions on lateral margins absent; with 39 ��� 41 pairs of setae (usually j1���j6, z1���z6, s1���s6, r2���r6, J1���J5, Z1���Z5, S1���S5, R 1��� R 3 ı but any of j2, z1, z2, s2, J5 sometimes asymmetrically lacking on one sideı or any of r5, r6, R 2, R 3 sometimes asymmetrically off shield on one side) and 23 pairs of distinguishable pore-like structures (seven gland pores and 16 poroids). Soft cuticle with setae R 4��� R 6 (usually visible in ventrolateral regions) and poroids Rp. Setae Z5 sparsely barbedı other dorsal setae simpleı needle-like; measurements of setae: j1 33 ��� 34 (36 ��� 44)ı j2 35 (36 ��� 46)ı j3 40 ��� 41 (42 ��� 46)ı j4 38 ��� 41 (40 ��� 50)ı j5 37 (38 ��� 48)ı j6 40 ��� 42 (42 ��� 55)ı z1 22 (23 ��� 29)ı z2 38 (46 ��� 50)ı z3 42 ��� 46 (48 ��� 55)ı z4 50 (53 ��� 67)ı z5 47 (48 ��� 61)ı z6 38 ��� 44 (44 ��� 55)ı s1 33 (32 ��� 38)ı s2 35 ��� 38 (36 ��� 40)ı s3 44 ��� 51 (46 ��� 63)ı s4 56 ��� 57 (48 ��� 69)ı s5 48 ��� 56 (59 ��� 69)ı s6 44 ��� 50 (44 ��� 61)ı r2 42 ��� 43 (40 ��� 50)ı r3 50 ��� 51 (46 ��� 61)ı r4 41 ��� 44 (50 ��� 65)ı r5 52 ��� 53 (50 ��� 61)ı r6 36 ��� 38 (40 ��� 51)ı J1 40 ��� 41 (40 ��� 55)ı J2 42 ��� 43 (42 ��� 50)ı J3 39 (38 ��� 48)ı J4 39 ��� 40 (40 ��� 57)ı J5 17 (19 ��� 27)ı Z1 43 ��� 44 (38 ��� 55)ı Z2 44 ��� 49 (40 ��� 53)ı Z3 39 ��� 40 (36 ��� 53)ı Z4 37 ��� 39 (36 ��� 57)ı Z5 72 ��� 73 (63 ��� 95)ı S1 39 ��� 41 (42 ��� 55)ı S2 42 ��� 44 (44 ��� 63)ı S3 41 ��� 43 (33 ��� 34)ı S4 42 (42 ��� 59)ı S5 31 ��� 35 (34 ��� 50)ı R 1 32 ��� 34 (36 ��� 46)ı R 2 36 ��� 37 (36 ��� 46)ı R 3 35 (38 ��� 48)ı R 4 36 ��� 35 (36 ��� 55)ı R 5 35 ��� 37 (36 ��� 44)ı R 6 32 ��� 35 (36 ��� 46)ı UR1���UR3 32 ��� 35 (34 ��� 48). Idiosomal venter (Figures 2 and 5 (b)). Tritosternum with trapezoidal baseı 15 (15 ��� 19) longı 15 (13 ��� 15) wide at baseı laciniae piloseı fused for about fourth of total length 17 (17 ��� 19)ı their free parts 45 (44 ��� 48) long. Pre-sternal area transversely lineateı without platelets. Sternal shield fused with endopodal strips of coxae I/II and coxae II/III; 108 (103 ��� 109) long along midlineı 172 (168 ��� 189) wide at level of endopodal projections between coxae I/IIı 157 (158 ��� 164) wide at level of endopodal projections between coxae II/IIIı 95 (97 ��� 91) wide at narrowest interval about mid-level of coxae II; with three pairs of setae (st1���st3) (37 ��� 41 (36 ��� 46) long)ı two pairs of poroids (iv1, iv2) and gland pores gvb; poroids iv1 positioned posteriad seta st1 ı poroids iv2 positioned between setae st2 and st3 ı gland pores gvb positioned on apex of endopodal projections between coxae I and II; posterior margin weakly concaveı arched to level of bases of setae st3; reticulated over all of surface except posteriorly where overlapped by hyaline flap of epigynal shield. Setae st4 (38 ��� 40 (38 ��� 42) long) located on tiny metasternal platelets with greatest dimension only about twice diameter of setal alveolus; poroids iv3 absent. Anterior margin of epigynal shield broadly roundedı overlapping posterior sternal shield areaı extending to level of setae st2 ı posterior margin of shield slightly rounded; epigynal shield reticula outline about 10 elongate cells between bases of setae st5; epigynal shield 180 (181 ��� 185) longı with greatest width of hyaline anterior part 102 (105 ��� 118)ı greatest width of posterior part 77 (76 ��� 84); length st5 34 (34 ��� 38); poroids iv5 placed on soft cuticle closely beside shield. Postgenital platelets absent. Free endopodal plates strongly formed between coxae III and IV. Anal shield ovalı slightly reticulateı with anterior margin roundedı usually indented mediallyı posterior margin nearly truncate; 91 (90 ��� 105) long and 76 (76 ��� 82) wide; anus located slightly anterior to mid-level of shieldı anal opening 33 (34 ��� 40) long; cribrum well-developed; one pair of gland pores present (gv3); length of para-anal setae 21 (21 ��� 23)ı length of post-anal seta 47 (48 ��� 61). Posteriad coxae IV two pairs of elongate metapodal platelets present; the larger platelet 21 (21 ��� 23) longı 7 (6 ��� 8) wide; the smaller platelet 11 (6 ��� 11) longı 5 (3 ��� 5) wide. Soft cuticle around anal shield with 12 ��� 13 pairs of setae (JV1���JV5, ZV1���ZV5 ı flanked by two or three pairs of UR setae) and five pairs of distinguishable poroids. All ventral setae simple; length of setae of JV-, ZV - and UR -series 27 ��� 36 (27 ��� 44) (excluding JV5 60 (61 ��� 84)). Exopodal platelets of coxae II ��� IIIı coxae III ��� IV and platelet enveloping coxa IV posteriorly fused into a single stripı bearing gland pores gv2 at posteromedial extremity. Peritrematal shields fused with dorsal shield at level of setae s1; with four pairs of distinguishable pore-like structures (poroids ip2, ip3 ı and gland pores gp2, gp3); peritreme extending slightly anteriad of setae s1. Spermathecal apparatus not distinguishable. Gnathosoma (Figure 3 (a���f) and 5(c)). Epistome triramousı with medial tine slightly serrated apicallyı longer than lateral tinesı these serrated along their outer margins (Figure 3 (a ��� c) and 5(c)). Subcapitulum 99 (101 ��� 109) wide at widest level. Deutosternum with seven rows of denticlesı rows 1 ��� 5 connected; anterior four rows each with three to eight denticlesı 5th row widenedı concave mediallyı with 10 ��� 15 denticlesı 6th row freeı widenedı with 10 ��� 17 denticlesı 7th row freeı with four to eight denticles (Figure 3 (d)). Hypostome with four pairs of simple setae; palpcoxal seta (pc) 19 ��� 20 (17 ��� 21)ı hp1 28 ��� 29 (27 ��� 32)ı hp2 20 (15 ��� 16)ı hp3 35 ��� 37 (34 ��� 38). Corniculi 29 ��� 34 longı 8 ��� 11 wideı symmetricalı horn-likeı sclerotisedı with paraxial process near base; internal malae slenderı extending to apex of corniculi; salivary styli with blunt apices not reaching tips of corniculi. Palp length from trochanter to tarsus apex 148 ��� 150 (145 ��� 158); setal formula normal: 2 ��� 5 ��� 6 ��� 15 ��� 16; palpfemoral seta al ı palpgenual setae al1 and al2 spatulateı palptarsal apotele 2-tined (Figure 3 (dı e)). Second cheliceral segment length 88 ��� 90 (90 ��� 101)ı movable digit length 36 (36 ��� 42). Fixed cheliceral digit with 17 (16 ��� 20) similar small teethı offset subapical tooth in addition to apical hookı and membranous lobeı distal process absentı antiaxial lyrifissure and dorsal seta distinct; movable digit with three teeth in addition to apical hookı and ventral projection (Figure 3 (f)). Legs (Figures 4 (a���d) and 5(d)). Lengths: I 410 ��� 420 (419 ��� 446)ı II 332 ��� 339 (353 ��� 409)ı III 364 ��� 373 (372 ��� 428)ı IV 469 ��� 476 (476 ��� 530). Leg chaetotactic formulae normal for genus: leg I: coxa 2ı trochanter 6 (1 0/1 1/2 1)ı femur 12 (2 3/1 2/2 2)ı genu 13 (2 3/2 3/1 2)ı tibia 13 (2 3/2 3/ 1 2); leg II: coxa 2ı trochanter 5 (1 0/1 0/2 1)ı femur 11 (2 3/1 2/2 1)ı genu 11 (2 3/1 2/1 2)ı tibia 10 (2 2/1 2/1 2); leg III: coxa 2ı trochanter 5 (1 0/1 0/2 1)ı femur 6 (1 2/1 1/0 1)ı genu 9 (2 2/1 2/1 1)ı tibia 8 (2 1/1 2/1 1) lacking pl2; leg IV: coxa 1ı trochanter 5 (1 0/1 0/2 1)ı femur 6 (1 2/1 1/0 1)ı genu 9 (2 2/1 3/0 1)ı tibia 10 (2 1/1 3/1 2). Tarsi II-IV: 18 (3 3/2 1/1 3/2 3). Tarsus I with erect middorsal macroseta 48 (53 ��� 61) long; tarsus IV with two erect macrosetaeı ad2 70 ��� 71 (74 ��� 101) and pd2 79 ��� 85 (99 ��� 122); all setae smooth. Etymology. The specific name of the new species is derived from the Latin word insolitus ı meaning ���unusual��� ı and refers to the absence of the distal flangelike process of the fixed chela of its female. Differential diagnosis Mucroseius insolitus sp. nov. differs from all known congeners by the absence of the distal flangelike process of the female fixed chela. The following keyı modified from that of Lindquist and Wu (1991) ı further distinguishes the new species from others in the genus. Key to adult females of Mucroseius (based on Lindquist and Wu 1991) 1. Tarsus IV with 2 macrosetaeı pd2 and ad2 ı distinguished by nearly perpendicular orientation and greater length from other tarsal setae........................................................ 2 ��� Tarsus IV with 1 macrosetaı only pd2 readily distinguished as such............................ 7 2. Fixed chela with broadly rounded distal flange; tibia III with 9 setae (pl2 present); dorsal shield ornamented with conspicuous little ridges and reticula over entire surface................................................................................................................................ M. aciculatus ��� Fixed chela with slenderı bluntly pointed distal flangeı or flange absent; tibia III with 8 setae (pl2 absent); dorsal shield smooth or reticulateı not carinulate...................... 3 3. Fixed chela without distal flange............................................................ M. insolitus sp. nov. ��� Fixed chela with slender distal flange.......................................................................................... 4 4. Epistome weakly triramousı its median process broadı untapered and conspicuously serratedı and its lateral processes weakly projected and conspicuously serrated; dorsal shield with setae s4 nearly as long or longer than caudal setae Z5; leg IV with anterior macroseta subequallyı or within 15 ��m as long ası posterior macroseta.. M. disparisetus ��� Epistome clearly triramousı its medial process tapering to a slightly serrated or smooth pointı and its lateral processes well projected and sparsely serrated; dorsal shield with setae s4 usually not more than 0.8 as long as caudal setae Z5 (some M. algonquian excepted); leg IV with anterior macroseta clearly (by 20 ��� 30 ��m) shorter than posterior macroseta........................................................................................ 5 5. Dorsal shield smooth over most of surfaceı with 7 ��� 9 pairs of marginal setae (r2���r6, R 1��� R 2 ı sometimes R 3��� R 4) inserted on its lateral edges; epigynal shield reticula outline 12 ��� 16 finely elongate cells between bases of setae st5; sternal shield with posterior margin deeply excavated to level of setae st2............................... M. nipponensis ��� Dorsal shield smooth or reticulated over most of surfaceı with only 2 or 3 pairs of marginal setae (r2���r3 ı sometimes r4) inserted on its lateral edges; epigynal shield reticula outline 9 ��� 11 elongate cells between bases of setae st5; sternal shield with posterior margin deeply excavated or not................................................................................ 6 6. Dorsal shield conspicuously reticulate over most or all of surface (sometimes smooth medially from setae j4 to j6 ı and from J1 to J4); sternal shield with posterior margin often deeply excavated to level of setae st2 or poroids iv2 ı occasionally this margin remaining weakly evident at level of setae st3.................................................. M. algonquian ��� Dorsal shield mostly unornamentedı smooth over most of surface (lightly reticulated along anterolateral marginsı and sometimes midlaterally between setae j6���J2 and z6���Z2 and caudally); sternal shield with posterior margin gently concave to level of setae st3........................................................................................................................ M. monochami 7. Dorsal shield strongly ornamentedı squamateı over entire surface; 8 or 9 pairs of marginal setae (r2���r6, R 1��� R 3 ı sometimes R 4) inserted on lateral edges of dorsal shield; sternal shield with posterior margin deeply excavated to level of setae st2............................ M. squamosus ��� Dorsal shield unornamented over most of surface; 3 ��� 6 pairs of marginal setae (r2���r4 ı sometimes r5���r6 ı rarely R 1 or R 2) inserted on lateral edges of dorsal shield; sternal shield with posterior margin gently concave to level of setae st3................................ 8 8. Anal shield with anterior margin flattened medially; dorsal shield usually lacking one or both setae of pair z3 ı and with setae J1 and J2 extending slightly beyond bases of J2 and J3 ı respectively.................................................................................................. M. californicus ��� Anal shield with anterior margin smoothly rounded medially; dorsal shield with both setae of pair z3 presentı and with setae J1 and J2 barely or not quite extending to bases of J2 and J3 ı respectively............................................................................ M. mexicanus Remarks Table 1 presents a list of species of the genus Mucroseius with their distribution and hosts. Key to adult females of Mucroseius (based on Lindquist and Wu 1991) 1. Tarsus IV with 2 macrosetaeı pd2 and ad2 ı distinguished by nearly perpendicular orientation and greater length from other tarsal setae........................................................ 2 ��� Tarsus IV with 1 macrosetaı only pd2 readily distinguished as such............................ 7 2. Fixed chela with broadly rounded distal flange; tibia III with 9 setae (pl2 present); dorsal shield ornamented with conspicuous little ridges and reticula over entire surface................................................................................................................................ M. aciculatus, Published as part of Trach, Viacheslav A., Khaustov, Alexander A. & Lindquist, Evert E., 2019, A new unique species of Mucroseius Lindquist�� 1962 (Acari: Mesostigmata: Melicharidae) mites associated with sawyer beetles (Cerambycidae: Monochamus Dejean�� 1821) from the Palaearctic Region, pp. 761-778 in Journal of Natural History 53 (13) on pages 764-769, DOI: 10.1080/00222933.2019.1606954, http://zenodo.org/record/3675600, {"references":["Lindquist EEi Wu KW. 1991. Review of mites of the genus Mucroseius (Acari: Mesostigmata: Ascidae) associated with sawyer beetles (Cerambycidae: Monochamus and Mecynippus) and pine wood nematodes [Aphelenchoididae: Bursaphelenchus xylophilus (Steiner and Buhrer) Nickle] i with descriptions of six new species from Japan and North Americai and notes on their previous misidentification. Can Entomol. 123: 875 - 927."]}

Supplemental Material for Have policy process scholars embraced causal mechanisms? A review of five popular frameworks by Jeroen van der Heijden, Johanna Kuhlmann, Evert Lindquist and Adam Wellstead in Public Policy and Administration

Podotarsonemus zuluensis Seeman, Lindquist & Husband sp. nov. Figures 25���27 Type material. Holotype. Adult female. SOUTH AFRICA, Natal, Ntumeni, D. Otte, 7 Mar. 1965, ex probable Pelusca sp. (Tetrigidae), specimen # 1. RWH 10VIII2010 -12. In UMMZ. Paratypes. 7 females, 1 male, 5 larvae, as follows. 4 females, 2 larvae, same data as holotype. 4 females, 1 male, 4 larvae, same data as holotype except host specimen #2. In UMMZ except 1 female, 1 larva in each of CNCI and QM. Diagnosis. Female and male: Palps with 2 setae (1 minute); chelicerae without teeth; prodorsal plate entirely punctate; femora I���II without setae d, l��; tibia I seta l��� short (3���5); genua II���III bare; tarsus II with 4 setae, seta u���� absent; tarsus III with 5 setae, seta u���� present; trochanter IV with seta. Female: Setae sc1 present; vestigial setae v2 expressed as anterolateral invaginations; coxisternal plates III���IV separated by longitudinal striae; setae h short, 5��� 7 long; tracheal atria separate, post-atrial sac lightly sclerotized, not punctate. Male: Genital capsule without copulatory flanges; setae h1 rudimentary, like h2; tibia IV with solenidion, without setae; tarsus IV rudimentary, subterminal. Male and larva: Leg I with tibia and tarsus fused. Larva: setae h1-2 and ps1-2 short, 5���12 long leg I without claws. Description. ADULT FEMALE (Fig. 25, n = 7 measured) Gnathosoma. Length 36���45, width 40���41. Palps 6���7, with 2 setae (2���3 and 1 long). Cheliceral stylets 18���22, without teeth. Pharynx length 25���27, width 11���14. Dorsal setae ch 19���21, ventral setae su 13���19, palpcoxal setae e 6���8. Idiosoma. Length 205���260, width 145���165. Tracheal atria separate, post-atrial sac lightly sclerotized, not punctate, with two small prongs indicating separate terminal tracheae. Prodorsal plate with punctate sculpturing, 76���90 long, 93���104 wide; with 3 pairs of setae: v 1 14���16, thickened, blunt, with barbs, sc 1 11���14, sc2 51���60, and vestigial v2 expressed as anterolateral invaginations. Distance between setae v1-v 1 24���27, v2-v2 32 ���34, sc1-sc1 37���39, sc2-sc2 70��� 81, v1 -sc 2 17���22. Plates C, D, EF and H with punctate sculpturing. Setae c1, d, e, f blunt, with small barbs, c 1 13���15, c2 37���42, d 13���15, e 7���10, f 10���11. Distance between setae c1-c1 56���61, c2-c2 107���115, c1-c 2 22���26, d-d 26���33, e-e 40���50, f-f 16���23. Setae h short, 5���7. Distance between setae h-h 22���25. Cupules ia, im, ih present. Setae ps attenuate> 60, distance between setae ps-ps 11���13. Coxisternal setae short to attenuate, 1a 13���19, 2a 13���25, 3a > 80, 3b > 45. Distance between setae 1a-1a 15���18, 2a-2a 31���39, 3a-3a 13���17, 3b-3b 25���34; alveolar vestiges of setae 1b present, 2b not evident. Coxisternal plates III���IV separated by longitudinal striae; coxisternal plates III convex anteriorly, sometimes just overlapping posterior margin of coxisterna II. Legs. Leg lengths: I 45 ���46, II 41 ���45, III 41 ���45. Leg IV Fe-Ge length 17���20, Ti-Ta length 15���17. Leg III, femur and genu fused. Femur I l���� rudimentary, v���� 18���24; genu I l�� 3, l���� 2���3, v�� 4���5, v���� 4���5; tibia and tarsus I fused, tibial setae d 24��� 33, l�� 7���8, l���� 3���5, v�� 5���9, v���� 8���11, solenidion �� 3, tarsal setae tc�� 12���13, tc���� 11���12, pv�� 8���9, pv���� 5���6, s spine-like, 6, solenidion �� 7���8. Femur II v���� 11���14; tibia II d 15���31, l�� 6���8, v�� 15���19, v���� 17���19; tarsus II tc�� 16���17, tc���� 18���23, pv�� 14���19, u�� spine-like, 5���6, solenidion �� 3���4. Tibia III d 23���25, l�� 7���11, v�� 18���23, v���� 17���22; tarsus III tc�� 17���20, tc���� 22���27, pv�� 16���18, u�� spine-like, 5���6, u���� 3���4. Trochanter IV with ventral seta 10���13; femorogenu IV bare; tibiotarsus IV with ventral tibial seta 14���18 and terminal tarsal seta> 240. ADULT MALE (Fig. 26, n = 1 measured) Gnathosoma. Damaged. Cheliceral stylets 18, without teeth. Pharynx length 27, width 10. Dorsal setae 13, ventral setae 6, palpcoxal setae 8. Idiosoma. Length 139, width 108. Prodorsal plate punctate, 57 long, 81 wide; with 3 pairs of barbed setae (v1 8, sc1 6, sc2 21) and vestigial v2. Distance between setae v1-v1 25, v2-v2 31, sc1-sc1 57, sc2-sc2 26, v1 -sc2 32. Plate CD punctate. Setae with small barbs, setae c1, 8, c2 9, d 8; setae c 1 in transverse alignment with setae c2. Plate EF punctate, e 5, f 6. Distance between setae c1-c1 53, c2-c2 96, c1-c2 21, d-d 8, e-e 21, f-f 8. Cupules ia, im present. Genital capsule 26 long, 28 wide, bearing alveolar remnants of h1, h2; ventral flanges absent. Coxisternal setae small, 1a 3, 2a 8, 3a 13, 3b 10. Distance between setae 1a-1a 14, 2a-2a 36, 3a-3a 19, 3b-3b 20; setae 1a just posterior to ap1, setae 2b far posterior to ap2. Apodemes: ap2 almost reaching medially obsolete ap sj; ap5 small, extend 12 from base, not fused medially. Legs. Leg lengths: I 36, II 35, III 32, IV 21. Leg III, femur and genu separate. Femur I, l���� rudimentary, v���� 12; genu I l�� 3, l���� 3, v�� 2, v���� 2; tibia and tarsus I fused, tibial setae d 21, l�� 3, l���� 2, v�� 2, v���� 5, solenidion �� 2, tarsal setae ft�� 9, ft���� 8, tc�� 10, tc���� 9, pv�� 6, pv���� 5, s spine-like, 5, solenidion �� 6. Femur II v���� 13; tibia II d 15, l�� 3, v�� 12, v���� 13; tarsus II tc�� 7, tc���� 14, pv�� 7, u�� spine-like, 5, solenidion �� 6. Tibia III d 14, l�� 4, v�� 10, v���� 8; tarsus III tc�� 15, tc���� 14, pv�� 12, u�� spine-like, 5, u���� 3. Trochanter IV with ventral seta 3 long; femorogenu IV bare; tibia IV solenidion �� 7; tarsus IV rudimentary, subterminal, with vestigial seta. LARVA (Fig. 27, n = 4 measured) Gnathosoma. Length 36���44, width 39���46. Palps 5. Cheliceral stylets 19���22, without teeth. Pharynx length 20���26, width 10���12. Dorsal setae 10���11, ventral setae 6���8. Idiosoma. Length 190���195, width 115���120. Prodorsal plate smooth, 55 long, split in all specimens; with 3 pairs of weakly barbed setae (v1 5���7, sc1 5���6, sc 2 6���10) and vestigial v2. Distance between setae v1-v 1 13���25, v2- v2 24, sc1-sc1 59, sc2-sc2 39, v1 -sc 2 24���25. Plates C, D, EF punctate. Setae tapering, barbed, c 1 8���10, c2 8, d 9��� 10, e 8���9, f 8. Distance between setae c1-c1 51���54, c2-c2 96���100, c1-c 2 24���26, d-d 22���27, e-e 32���34, f-f 14���16. Plate H punctate, setae h1 short, 7���8, h2 ventral, 5���12 (unpaired h 3 in one specimen). Distance between setae h 1 17���21. Cupules ia, im, ih present. Setae ps1 fine, 8���10, ps2 fine, 10. Coxisternal setae 1a 4���5, 2a 4���7, 3a attenuate, 10���17, 3b 5���10. Distance between setae 1a-1a 13���26, 2a-2a 27���32, 3a-3a 34, 3b-3b 26. Legs. Femur I, l���� rudimentary, v���� 7���10; genu I l�� 1, l���� 1���2, v�� 2���3, v���� 2���3; tibia and tarsus I fused, tibial setae d 10���11, l�� 2���3, l���� 2, v�� 2���3, v���� 5���6, solenidion �� 2���4, tarsal setae tc�� 7���8, tc���� 8, pv�� 5, pv���� 3���4, s spine-like, 5, solenidion �� 3���4. Femur II v���� 3���4; tibia II d 7���8, l�� 1���2, v�� 3���5, v���� 9; tarsus II tc�� 4���7, tc���� 9���10, p�� 1, u�� spine-like, 5, u���� 1, solenidion �� 2. Tibia III d 7���8, l�� 2���3, v�� 6���8, v���� 9���10; tarsus III tc�� 7���8, tc���� 12���15, p�� 1, u�� spine-like, 5, u���� 1. Leg I without claws. Etymology. The species name zuluensis refers to the presence of this species in Zululand, which is now a part of KwaZulu-Natal, South Africa. Remarks. Podotarsonemus zuluensis is most similar to P. indicus but differs by several character states as noted in the Differential diagnosis for P. indicus, and also detailed in the key below., Published as part of Seeman, Owen D., Lindquist, Evert E. & Husband, Robert W., 2018, A new tribe of tarsonemid mites (Trombidiformes: Heterostigmatina) parasitic on tetrigid grasshoppers (Orthoptera), pp. 1-54 in Zootaxa 4418 (1) on pages 42-47, DOI: 10.11646/zootaxa.4418.1.1, http://zenodo.org/record/1242435

Podotarsonemus indicus Seeman, Lindquist & Husband sp. nov. Figures 16���19 Type material. Holotype. Adult female. INDIA, Madras, Dist. Coimbatore, Coimbatore, P.S. Nathan, 5 Jul. 1946, ex Scelimena kempi (Hancock), RWH VII2010 (slide 1). In UMMZ. Paratypes. 15 females, 11 males, 2 larvae, as follows. 11 females, 11 males, 1 larva same data as holotype. 3 females, 1 larva, same data as holotype except 26 Nov. 1945. 1 female, same data as holotype except 3 Nov. 1947. In UMMZ, except 1 female, 1 male, 1 larva in CNCI and 1 female, 1 male in QM. Diagnosis. Female and male: Palps with 2 setae; chelicerae without teeth; prodorsal plate with punctate sculpturing; femora I���II without setae l��, d; genua II���III bare; tarsi II���III with 5 setae, u���� present; trochanter IV with 1 seta. Female: Setae sc1 present; vestigial setae v2 expressed as alveoli within prodorsal shield; coxisternal plates III���IV separated by longitudinal striae; setae h not attenuate, h1 rudimentary, like h2; leg I with tibia and tarsus fused; femorogenu IV bare; tibia IV with solenidion and attenuate terminal seta; tarsus IV rudimentary, subterminal. Larva: Setae h1-2, ps2 attenuate, setae ps1 short; leg I with separate tibia and tarsus, with 2 rudimentary claws. Description. ADULT FEMALE (Figures 16���17, n = 9 measured) Gnathosoma. Length 57���67, width 68���71. Palps 9���13, with 2 setae (8 and 3 long). Cheliceral stylets 28���31, without teeth. Pharynx length 41���50, width 20���23. Dorsal setae ch 35���42, ventral setae su 27���32, palpcoxal setae e 7���8. Idiosoma. Length 242���300 (physogastric 600), width 226���240. Tracheal atria fused, post-atrial sac heavily sclerotized, punctate, with single long terminal extension of trachea. Prodorsal plate with punctate sculpturing, 103���110 long, 148���174 wide; with 3 pairs of setae: v1 21 ���26, thickened, with small barbs, sc 1 17���21, sc2 67���78, and vestigial v2. Distance between setae v1-v1 63 ���75, v2-v2 68 ���82, sc1-sc1 124���140, sc2-sc2 83���106, v1 -sc2 33��� 38. Plates C, D, EF with punctate sculpturing. Setae c1, d, e, f thickened, with small barbs, c 1 18���20, c2 51���65, d 17���20, e 15���21, f 15���20. Distance between setae c1-c1 122���128, c2-c2 201���209, c1-c2 35���38, d-d 64���76, e-e 73��� 80, f-f 28���34. Plate H variably covered by EF, setae h not attenuate, 11���26. Distance between setae h-h 31���37. Cupules ia, im, ih present. Setae ps attenuate> 300, distance between setae ps-ps 9���12. Coxisternal setae attenuate, 1a 80 (often broken), 2a > 120 (often broken), 3a > 100, 3b > 100. Distance between setae 1a-1a 26���31, 2a-2a 59��� 63, 3a-3a 15���18, 3b-3b 35���43; alveolar vestiges of setae 1b and 2b present. Coxisternal plates III���IV separated by longitudinal striae; coxisternal plates III convex anteriorly, sometimes just reaching posterior margin of coxisterna II. Legs. Leg lengths: I 71 ���73, II 75 ���79, III 70 ���73. Leg IV Fe-Ge length 33���34, Ti-Ta length 20���22. Leg III, femur and genu fused, line of fusion apparent and one specimen with asymmetric separation of femur and genu. Femur I l���� 5���6, v���� 15���28; genu I l�� 9���11, l���� 9���10, v�� 3���5, v���� 4���5; tibia and tarsus I fused, tibial setae d 31���34, l�� 9���12, l���� 9���11, v�� 6���7, v���� 11���13, solenidion �� 3���4, tarsal setae tc�� 12���14, tc���� 12���13, pv�� 10���13, pv���� 8���10, s spinelike, 6���7, solenidion �� 7���8. Femur II v���� 15���24; tibia II d 24���30, l�� 6���11, v�� 13���21, v���� 15���25; tarsus II tc�� 16���17, tc���� 18, pv�� 13���15, u�� spinelike, 7, u���� 6���8, solenidion �� 6���7. Tibia III d 24���32, l�� 14���20, v�� 18���22, v���� 17���23; tarsus III tc�� 17���22, tc���� 26���32, pv�� 15���17, u�� spine-like, 8, u���� 5��� 6. Trochanter IV with seta 11���13; tibiotarsus with ventral tibial seta 45���60, terminal tarsal seta> 400. ADULT MALE (Fig. 18, n = 5 measured) Gnathosoma. Similar to female. Length 61, width 66. Palps 11���12. Cheliceral stylets 2 5���26, without teeth. Pharynx length 34���35, width 18. Dorsal setae 39���41, ventral setae 7, palpcoxal setae 3. Idiosoma. Length 170���238, width 168���179. Prodorsal plate punctate, 74 long, 104���105 wide; with 3 pairs of setae (v1 14 ���18, sc 1 17���18, sc2 75���78) and vestigial v2. Distance between setae v1-v1 34 ���38, v2-v2 44 ���50, sc1-sc1 87���90, sc2-sc2 64��� 65, v1 -sc 2 29���32. Plate CD punctate. Setae c1 tapering, with few barbs, 7, c2 attenuate, 70���84, d with few barbs, 19���20; setae c 1 in nearly transverse alignment with c2. Plate EF punctate, e 18���19, f 16, with few barbs. Distance between setae c1-c1 64, c2-c2 134, c1-c 2 21���25, d-d 33, e-e 47���51, f-f 16. Cupules ia, im present. Genital capsule 43 long, 51 wide, bearing alveolar remnants of h1, h2; ventral flanges absent. Coxisternal setae attenuate 1a 30���36, 2a 28���30, 3a > 190, 3b > 30. Distance between setae 1a-1a 33���35, 2a-2a 56���59, 3a-3a 19���23, 3b-3b 29���31; setae 1a at level of or just posterior to ap1, setae 2b usually just posterior to ap2 (one specimen with asymmetrical position of setae 2b far posterior to ap2). Apodemes: ap2 not reaching medially obsolete ap sj; ap5 obsolete. Legs. Leg lengths: I 70 ���72, II 83 ���89, III 83 ���85, IV 52 ���54. Leg III, femur and genu separate. Femur I, l���� 5���8, v���� 30; genu I l�� 8���12, l���� 7���8, v�� 4, v���� 3���4; tibia and tarsus I fused, tibial setae d 44���50, l�� 7���10, l���� 10, v�� 7, v���� 15, solenidion �� 5, tarsal setae ft�� 16, ft���� 15, tc�� 15���17, tc���� 15���17, pv�� 12, pv���� 14, s spine-like, 7, solenidion �� 7���8. Femur II v���� 25; tibia II d 38���42, l�� 27���29, v�� 10���18, v���� 31���32; tarsus II pv�� 6���9, tc�� 18���20, tc���� 18���20, u���� 5���6, u�� spine-like, 6, solenidion �� 8. Tibia III d 32���35, l�� 27���30, v�� 26���32, v���� 23���25; tarsus III pv�� 17���20, tc�� 27���28, tc���� 25���29, u�� spine-like, 7, u���� 5��� 6. Trochanter IV with seta, 8; femorogenu IV bare; tibia IV �� 11, terminal seta attenuate, 150; tarsus IV rudimentary, subterminal. LARVA (Fig. 19, n = 1 measured) Gnathosoma. Length 46, width not measurable. Palp length 5, with 2 setae (lengths 6 and minute). Cheliceral stylets 26, without teeth. Pharynx length 29, width 18. Dorsal setae 18, ventral setae 9. Idiosoma. Length 225, width 160. Prodorsal plate smooth, 60 long, 95 wide; with 3 pairs of tapering, weakly barbed setae (v1 10, sc1 8, sc2 17) and vestigial v2. Distance between setae v1-v1 28, v2-v2 49, sc1-sc1 not measurable as sc1 missing on left hand side, sc2-sc2 58, v1 -sc2 41. Plates C, D, EF smooth. Setae tapering, weakly barbed, c1 17, c2 14, d 14, e 12, f 15. Distance between setae c1-c1 76, c2-c2 130, c1-c2 30, d-d 28, e-e 41, f-f 13. Plate H smooth, setae h1 long, 65, h2 ventral,> 120. Distance between setae h1 24. Cupules ia, im, ih present. Setae ps1 fine, 8, ps2 attenuate 60. Coxisternal setae attenuate, 1a 36, 2a 41, 3a > 40, 3b 30. Distance between setae 1a-1a 36, 2a-2a 41, 3a-3a 37, 3b-3b 35. Longitudinal striae between coxisterna III���IV. Legs. Femur I, l���� rudimentary, v���� 10; genu I l��, l���� rudimentary, v�� 3, v���� 2; tibia and tarsus I separate, tibial setae d 15, l�� 4, l���� 4, v�� 3, v���� 6, solenidion �� 3, tarsal setae tc�� 9, tc���� 9, pv�� 3, pv���� 6, s spine-like, 5, solenidion �� 4. Femur II v���� 7; tibia II d 9, l�� 5, v�� 11, v���� 11; tarsus II tc�� 7, tc���� 12, p�� 4, u�� spine-like, 5, u���� 2, solenidion �� 3. Tibia III d 9, l�� 6, v�� 10, v���� 10; tarsus III tc�� 6, tc���� 13, p�� 4, u�� spine-like, 5, u���� 3. Leg I with 2 rudimentary claws. Etymology. The species name indicus refers to the country where the type specimens were collected. Differential diagnosis. Podotarsonemus indicus belongs to the australiensis species group (P. australiensis, P. indicus, P. zuluensis) whose females retain a seta on trochanter IV, have longitudinal striae between coxisterna III��� IV and have setae h not attenuated. Of these species, P. indicus most closely resembles P. zuluensis but is distinguished by having: females with tracheal atria fused (free in other Podotarsonemus), females with alveolar remnants of setae v2 within the prodorsal shield (anterolateral invaginations of prodorsal shield in P. zuluensis), females and males retaining seta u��� on tarsus II (absent in P. zuluensis), males with setae c2 much longer (> ��4) than c1 (P. zuluensis) and tibia IV having an attenuate terminal seta and subterminal solenidion (lacking terminal seta and solenidion terminal in P. zuluensis), and larvae with setae h1-2 and ps2 attenuate (short in P. zuluensis)., Published as part of Seeman, Owen D., Lindquist, Evert E. & Husband, Robert W., 2018, A new tribe of tarsonemid mites (Trombidiformes: Heterostigmatina) parasitic on tetrigid grasshoppers (Orthoptera), pp. 1-54 in Zootaxa 4418 (1) on pages 30-35, DOI: 10.11646/zootaxa.4418.1.1, http://zenodo.org/record/1242435

Podotarsonemus nipponicus Seeman, Lindquist & Husband sp. nov. Figures 20–22 Type material. Holotype. Adult female. JAPAN, Maesato, Ishigaki-jimi Is., Okinawa Pref., T. Takahashi, 22 Aug. 1975, ex hind wings of Euparatettix tricarinatus (Bolívar) (Tetrigidae), Kurosa No. 4880-1. In NMST. Paratypes. 16 females, 7 males, 32 larvae, as follows. 3 females, 3 larvae, same data as holotype (slides 2–7). 2 females, 2 larvae, same data as holotype except 17 Jun. 1975, Kurosa No. 4877 (slides 1–4). 3 females, 1 male, 4 larvae, same data as holotype except 17 Jun. 1975, Kurosa No. 4879 (slides 1, 3, 4, 5, 6). 1 female, 3 males, 4 larvae, same data as holotype except 17 Jun. 1975, Kurosa No. 4881 (slides 1, 2, 3, 4–8). 3 females, 1 male, 3 larvae, same data as holotype except 17 Jun. 1975, Kurosa No. 4882 (slides 1–6, 8). 2 larvae, same data as holotype except Kurosa No. 4883 (slides 1–2). 4 females, 2 males, 14 larvae, same data as holotype except Kurosa No. 4884 (slides 1–18, 20–24). In NMST except 1 female, 1 male, 1 larva in each of CNCI, QM, UMMZ. Diagnosis. Female and male: Palps with 2 setae; chelicerae with 1 tooth (female) or without teeth (male); prodorsal plate with punctate sculpturing anteriorly, imbricate sculpturing behind sc2; femora I–II without seta lʹ; genua II–III bare; tarsi II–III with 5 setae, seta uʹʹ present; trochanter IV without seta. Female: Setae sc1 present; vestigial setae v2 expressed as alveoli within prodorsal shield; coxisternal plates III–IV separated by soft cuticle; setae h attenuate,> 100 long; femorogenu IV with 2 setae; tracheal atria separate, post-atrial sac lightly sclerotized, not punctate. Male: Genital capsule without copulatory flanges; setae h1 rudimentary, like h2; femorogenu IV with 1 seta; tibia IV with solenidion and rudimentary seta; tarsus IV absent. Male and larva: Leg I with tibia and tarsus fused. Larva: Setae h1-2, ps1-2 attenuate; leg I without claws. Description. ADULT FEMALE (Fig. 20, n = 15 measured) Gnathosoma. Length 41–42, width 43–45. Palps 10, with 2 setae (3–5 and 1–4 long). Cheliceral stylets 1 7–20, each with 1 tooth. Pharynx length 23–29, width 22–24. Dorsal setae ch 20–24, ventral setae su 14–23, palpcoxal setae e 8–13. Idiosoma. Length 166–180 (physogastric 275–500), width 135–139. Tracheal atria separate, post-atrial sac lightly sclerotized, not punctate, with two small distal prongs indicating separate terminal tracheae. Prodorsal plate with punctate sculpturing anteriorly, imbricate posterior to sc1, 72–78 long, 110–111 wide; with 3 pairs of setae: v 1 16–20, thickened, blunt, with small barbs, sc 1 6–12, sc2 58–61, and vestigial v2. Distance between setae v1-v1 34 – 38, v2-v2 42 –50, sc1-sc1 90–95, sc2-sc2 54– 62, v1 -sc 2 22–26. Plates C, D with punctate sculpturing anteriorly, imbricate posteriorly, EF and H with punctate sculpturing. Setae c1, d, e, f thickened, with small barbs, c 1 15–17, c2 40–47, d 15–19, e 15–17, f 17–20. Distance between setae c1-c1 70–77, c2-c2 127–137, c1-c 2 23–28, d-d 42– 46, e-e 41–46, f-f 20–26. Setae h attenuate,> 100. Distance between setae h-h 22–25. Cupules ia, im present, ih indiscernible. Setae ps attenuate 220, distance between setae ps-ps 9–11. Coxisternal setae short to attenuate, 1a 9– 11, 2a 19–25, 3a > 200, 3b 30. Distance between setae 1a-1a 15–21, 2a-2a 21–25, 3a-3a 5–13, 3b-3b 24–35; alveolar vestiges of setae 1b and 2b present. Legs. Leg lengths: I 49 –51, II 48 –50, III 50. Leg IV Fe-Ge length 19–21, Ti-Ta length 17–19. Leg III, femur and genu fused. Femur I d 1–2, lʹʹ 3–12, vʹʹ 22–27; genu I lʹ 9–15, lʹʹ 7–11, vʹ 10–13, vʹʹ 9–12; tibia and tarsus I fused, tibial setae d 23, lʹ 11–12, lʹʹ 4–6, vʹ 9–11, vʹʹ 12–13, solenidion φ 3–4, tarsal setae tcʹ 13–14, tcʹʹ 11–12, pvʹ 14–17, pvʹʹ 10–12, s spine-like, 7–8, solenidion ω 7–8. Femur II vʹʹ 12–19; tibia II d 21–23, lʹ 11–17, vʹ 16–22, vʹʹ 18–22; tarsus II tcʹ 15–20, tcʹʹ 20–23, pvʹ 16–20, uʹ spine-like, 7–8, uʹʹ 2–4, solenidion ω 6. Tibia III d 18–21, lʹ 17–23, vʹ 16–20, vʹʹ 16–21; tarsus III pvʹ 22–25, tcʹ 21–26, tcʹʹ 22–24, uʹ spine-like, 5–6, uʹʹ 6–8. Femorogenu IV with proximal femoral seta 13–18, distal genual seta 26–28; tibiotarsus with attenuate ventral tibial seta> 80 and terminal tarsal seta> 180. ADULT MALE (Fig. 21, n = 5 measured) Gnathosoma. Length 30–31, width 32–36. Palps 6–7. Cheliceral stylets 12–13, without teeth. Pharynx length 20–21, width 11–12. Dorsal setae 9–16, ventral setae 11–13, palpcoxal setae 6–8. Idiosoma. Length 127–138, width 99–108. Prodorsal plate punctate, 59–62 long, 83–94 wide; with 3 pairs of setae (v 1 10–11, sc 1 7–10, sc 2 28–40) and vestigial v2. Distance between setae v1-v 1 21–25, v2-v 2 27–30, sc1-sc1 56–59, sc2-sc 2 30–33, v1-sc2 33–39. Plate CD punctate. Setae c1 tapering, with small barbs, 11–13, c2 fine, 16– 21, d with small barbs, 10–12; setae c1 obliquely aligned between setae c2 and d. Plate EF punctate, e 5–7, f 6–8, with small barbs. Distance between setae c1-c1 51–59, c2-c2 85–92, c1-c 2 23–26, d-d 15–18, e-e 19–21, f-f 6–8. Cupules ia, im present. Genital capsule 27–30 long, 24–27 wide, bearing alveolar remnants of h1, h2; ventral flanges absent. Coxisternal setae attenuate 1a 9–11, 2a 13–17, 3a 80–100, 3b 16–20. Distance between setae 1a-1a 10–13, 2a-2a 23–25, 3a-3a 14–16, 3b-3b 14–17; setae 1a at level of or just posterior to ap1, setae 2b at level of ap2. Apodemes: ap2 almost reaching medially obsolete ap sj; ap5 small, extend 11–13 from base, not fused medially. Legs. Leg lengths: I 38 –39, II 40, III 38, IV 2 6–29. Leg III, femur and genu fused. Femur I, d 1–2, lʹʹ 3–4, vʹʹ 10–15; genu I lʹ 3–4, lʹʹ 3–4, vʹ 3–4, vʹʹ 3–4; tibia and tarsus I fused, tibial setae d 14– 19, lʹ 3–4, lʹʹ 2, vʹ 3–5, vʹʹ 6–8, solenidion φ 2, tarsal setae ftʹ 7–9, ftʹʹ 7–9, tcʹ 11–13, tcʹʹ 9–10, pvʹ 10–11, pvʹʹ 7–8, s spine-like, 6, solenidion ω 5–6. Femur II vʹʹ 6–7; tibia II d 14–17, lʹ 7–8, vʹ 10–11, vʹʹ 11–14; tarsus II tcʹ 10–12, tcʹʹ 11, pvʹ 7–9, uʹ spine-like, 6, uʹʹ 2, solenidion ω 8. Tibia III d 15–20, lʹ 7–10, vʹ 11–15, vʹʹ 13–15; tarsus III tcʹ 11–16, tcʹʹ 11–15, pvʹ 7–10, uʹ spine-like, 6, uʹʹ 5–6. Trochanter IV bare; femorogenu IV with genual seta 3–6; tibia IV solenidion φ 7, vʹ 1; tarsus IV absent. LARVA (Fig. 22, n = 30 measured) Gnathosoma. Length 37–41, width 33–39. Palps 6–7. Cheliceral stylets 11–14, without teeth. Pharynx length 15–17, width 11–12. Dorsal setae 11–14, ventral setae 8–10. Idiosoma. Length 167–250, width 114–155. Prodorsal plate smooth, 47–52 long, 89–99 wide; with 3 pairs of weakly barbed setae (v 1 8–10, sc1 6, sc 2 14–17) and vestigial v2. Distance between setae v1-v 1 25–29, v2-v2 37 – 44, sc1-sc1 59–65, sc2-sc2 41– 44, v1 -sc 2 16–21. Plates C, D, EF smooth. Setae weakly barbed, c 1 11–13, c 2 9–11, d 11–13, e 8–10, f 11–12. Distance between setae c1-c1 53–59, c2-c2 90–105, c1-c 2 20–24, d-d 28–33, e-e 28–30, f-f 11–15. Plate H smooth, setae h1 attenuate,> 70, h2 ventral, > 100. Distance between setae h 16–21. Cupules ia, im, ih present. Setae ps1-2 attenuate, ps1 ca. 40, ps2 > 40. Coxisternal setae 1a 4–8, 2a 8–10, 3a attenuate,> 110, 3b 10–14. Distance between setae 1a-1a 15–16, 2a-2a 30–31, 3a-3a 34–43, 3b-3b 37–48. Legs. Femur I, lʹʹ, d rudimentary, vʹʹ 6–10; genu I lʹ 1–2, lʹʹ 1–2, vʹ 1–2, vʹʹ 1–2; tibia and tarsus I fused, tibial setae d 10–16, lʹ 2, lʹʹ 1–2, vʹ 2–3, vʹʹ 6–7, solenidion φ 2, tarsal setae tcʹ 9–10, tcʹʹ 8, pvʹ 7–8, pʹʹ 2–3, s spine-like, 5– 6, solenidion ω 3–5. Femur II vʹʹ 3; tibia II d 7–10, lʹ 3–5, vʹ 7, vʹʹ 8–9; tarsus II tcʹ 10–13, tcʹʹ 9–11, pʹ 5–6, uʹ spine-like, 5–6, uʹʹ 7– 8, solenidion ω 3–5. Tibia III d 7, lʹ 3, vʹ 7–8, vʹʹ 7–9; tarsus III tcʹ 11–12, tcʹʹ 10–11, pʹ 4–5, uʹ spine-like, 5–6, uʹʹ 7–11. Leg I without claws. Etymology. The species name nipponicus refers to the country where the type specimens were collected. Differential diagnosis. Podotarsonemus nipponicus belongs to the boliviensis species group (P. boliviensis, P. ecuadorensis, P. nipponicus, and P. queenslandensis) whose females lack a seta on trochanter IV, lack longitudinal striae between coxisterna III–IV, and have setae h attenuated. Of these species, P. nipponicus is most similar to P. queenslandensis in having no more than three setae on femur I and just one seta on femur II. Podotarsonemus nipponicus is distinguished from P. queenslandensis, which is known only from the female, by having setae sc1 (absent in P. queenslandensis), retaining a rudimentary seta d on femur I (absent in P. queenslandensis) and having the prodorsal, C and D plates with some imbricate sculpturing (punctate in P. queenslandensis). Remarks. One specimen (4881-4) is a pharate male, demonstrating that both sexes have active, sexually monomorphic larvae. This is the plesiomorphic condition in the Tarsonemidae and other Acariformes, but within the Podapolipidae, only the genera Chrysomelobia and Tarsopolipus retain an active, monomorphic male (Husband 1997; Seeman & Nahrung 2005). Most other genera of Podapolipidae have free-living larval and adult females, but adult males that emerge directly from their eggs as sexually mature paedomorphic forms (i.e. larviform males). The only exception is Scarabapolipus which have active larvae in both sexes, but they are dimorphic (Husband & Kurosa 1994).

Key to species of Podotarsonemus 1. Female and male: with seta on trochanter IV (e.g., Figs 2, 4); female: coxisternal plates III–IV separate, longitudinal striae between coxisterna (Fig. 2); setae h small to moderate length, not attenuate (length australiensis species group................................................................... 2 - Female and male: without seta on trochanter IV; female: coxisternal plates III–IV not separate, lacking longitudinal striae between coxisterna (Fig. 8); setae h attenuate (length> 70); male: tarsus IV absent … boliviensis species group........... 4 2. Female: setae sc1 absent (Fig. 1); genua II–III with setae (l ′, l′′, v ′), femur I with four setae (l ′, l′′, d, v′′), femorogenu IV with two setae; male: tibia and tarsus I separate; femorogenu IV with seta, tarsus IV developed, with claw and setae (Fig. 3)............................................................................ Podotarsonemus australiensis sp. nov. - Female: setae sc1 present (Figs 16, 25); genua II–III without setae, femur I with two setae (l″, v ″), femorogenu IV bare; male: tibia and tarsus I fused; femorogenu IV without seta, tarsus IV rudimentary, without claw and setae (Figs. 18, 26)......... 3 3. Female: vestigial v2 formed as alveoli separate from lateral edges of prodorsal shield (Fig. 16), tracheal atria fused, post-atrial sacs heavily sclerotized and with long distal extension (Fig. 17); male: setae c2 much longer (> 4x) than c1, tibia IV bearing attenuate terminal seta, solenidion subterminal (Fig. 18); larva: setae h1-2, ps2 attenuate, tibia and tarsus I separate (Fig. 19)........................................................................... Podotarsonemus indicus sp. nov. - Female: vestigial v2 formed as anterolateral invaginations of lateral edges of prodorsal shield (Fig. 25), tracheal atria separate, post-atrial sacs lightly sclerotized and without distal extension (Fig. 25); male: setae c2 slightly longer (c1, tibia IV without terminal seta, solenidion terminal (Fig. 26); larva: setae h1-2, ps2 short, tibia and tarsus I fused (Fig. 27).................................................................................. Podotarsonemus zuluensis sp. nov. 4. Female and male: femur I with 4 setae (rudimentary l ′, l′′, d and developed v′′), femur II bearing 3 setae (rudimentary l ′, d and developed v′′) (e.g., Fig. 8).............................................................................. 5 - Female and male: femur I with 2 or 3 setae (if present, d rudimentary, and developed l′′, v′′), femur II bearing 1 seta (developed v′′) (e.g., Fig. 20)..................................................................................... 6 5. Female: prodorsal, C and D plates with imbricate sculpturing; chelicerae with 1 tooth (Fig. 7); male: femorogenu IV with 1 seta, small genital capsule flanges present (Fig. 9)............................... Podotarsonemus boliviensis sp. nov. - Female: prodorsal, C and D plates with punctate sculpturing; chelicerae without teeth (Fig. 13); male: femorogenu IV without seta, genital capsule flanges absent (Fig. 14)................................. Podotarsonemus ecuadorensis sp. nov. 6. Female: setae sc1 present; vestigial setae v2 expressed as alveolar pits within prodorsal shield; palp with two setae; femur I with rudimentary seta d (Fig. 20)............................................ Podotarsonemus nipponicus sp. nov. - Female: setae sc1 absent; rudimentary setae v2 expressed as anterolateral invaginations of prodorsal shield; palp with one seta; femur I without rudimentary seta d (Fig. 23).............................. Podotarsonemus queenslandensis sp. nov.

Podotarsonemus Seeman, Lindquist & Husband gen. nov. Type species: Podotarsonemus boliviensis Seeman, Lindquist & Husband sp. nov. Diagnosis. With attributes of tribe Podotarsonemini, augmented as follows: Palpi much reduced, without clear segmentation; apical palpal seta flanked medially by small hyaline triangular flange. Cheliceral stylets of larvae and adults curved from base to apex, with 0–2 teeth subapically. Adults and larvae with scapular setae sc 2 at most slightly longer than interval between their insertions, not exceedingly long, attenuated. Adult female prodorsal stigmata with main tracheal trunks leading internally to atrial chamber(s) and then to distinctively enlarged pair of post-atrial structures. Legs II–III of larvae and adults often lacking all genual setae (v″ always absent), and leg III lacking femoral setae. Larvae with setae on coxisternal plates III well developed, often attenuated, and with setae ps well-developed, sometimes attenuated, along with setae h. Male leg IV without flanges, femorogenu IV with maximum of one seta Ge- vʹ. In contrast to Acarapis, leg IV is slender in addition to characters already described for the tribe. Description. Gnathosoma. Form and structures of gnathosoma similar on larva, female, and male, though generally slightly larger on female. Capsule broadly oval in dorsoventral view, not beak-like anteriorly; palpcoxal setae present on adults. Dorsal gnathosomatic setae variably long, slender, smooth on adults, similarly formed on larva. Palpi directed anteriorly, short, stubby, with short but prominent, stout, pointed genual seta projected anteriorly and sometimes a smaller, inconspicuous femoral seta posterolaterally, these flanked anteroventrally by small, bluntly pointed, apical hyaline flange-like margin of femorogenual segment which covers rudimentary tibiotarsus with its terminal ventro-apical zone of alveolar vestiges of 3–5 small circular structures, probably rudiments of setae, solenidia and claw elements. Cheliceral stylets prominent, thick, variably protruding from capsule, smooth or with 1–2 teeth subapically, curved laterally in attachment to large basal levers oriented transversely across anterior one-third of capsule. Pharynx striated, muscular, with thinly sclerotized walls, enlarged to about one-third the width of gnathosomatic capsule, with paired gland-like structures indiscernible posteriorly. Idiosoma. ADULT FEMALE. Dorsal shielding unornamented or reticulated, imbricated. Dorsal setae collectively smooth or with small barbs on setae c1, d, e, f. Prodorsal shield not extended hood-like over gnathosoma. Stigmata opening closely anterolaterad setae v1; main tracheal trunks with enlarged, slightly sclerotized atrial sacs (sometimes fused into single sac), and also with slightly to heavily sclerotized post-atrial sacs, these entire (much as in Hemitarsonemus). Pits v2 obliquely aligned with setae v1 and sc2, and situated about one-quarter to midway along interval from v1 to sc2, sometimes at anterolateral invaginations of prodorsal shield. Setae sc2 long, attenuate, more widely spaced from each other, and longer, than v1, but not exceedingly long, at most slightly longer than transverse interval between their insertions, and inserted near midlength of prodorsal shield. Bothridia vestigial or absent, their sensilla sc1 reduced, slender, or absent. Tergite EF with setae e and f transversely aligned, similar in length, not grouped closely together. Tergite H with setae h short to exceedingly long, attenuate. Ventral shielding with apodemes 1 forming Y-shaped juncture with prosternal apodeme. Apodemes 2 fully or weakly connecting with prosternal apodeme. Prosternal apodeme developed posteriad level of apodemes 2 but not reaching posterior margin of coxisternal plates II. Sejugal apodeme reduced to a lateral fragment on either side. Apodemes 3 not extending laterad trochanters III, nor mediad setae 3a. Apodemes 4 reduced medially and not connected to each other, but extending posterolaterad setae 3b to base of trochanters IV. Poststernal apodeme absent. Coxisternal plates I and II with setae 1a and 2a short to exceedingly long, attenuated, similar in size. Anterior margin of coxisternal plates III straight or convex, if convex then sometimes extending over posterior edge of coxisternal plates II. Coxisternal setae 3a and 3b short to exceedingly long, attenuated, similar in size. Bases of legs IV well spaced, separated by interval of three to four widths of trochanter IV; posterior margin of coxisternal plates IV weakly formed, not formed into tegular projection. Pseudanal setae ps exceedingly long, attenuate. ADULT MALE. Dorsal shielding unornamented. Dorsal setae collectively smooth, much as in female, but if attenuated, then less extremely so. Prodorsum with three pairs of setae, including short but well developed sc1; setae sc2 attenuate but not exceedingly long, no longer than transverse interval between their insertions; setae v2 lacking but vestiges of their alveoli indicated by a pair of pits. Plate CD with setae arranged in two or three transverse rows, pair c2 anteriorly, d posteriorly, and c1 either at same level between c2 or at level between (and often obliquely aligned with) c2 and d. Plate EF with setae e and f more or less aligned transversely. Genital capsule with adhesive copulatory flanges reduced or absent. Sejugal apodeme evident laterally but indistinct medially. Coxisternal plates III and IV partly or entirely separated from each other medially by soft cuticle, such that apodemes 3 and 4 not reaching each other medially and poststernal apodeme absent; apodemes 5 moderately to weakly formed on either side at bases of legs IV, sometimes connected to each other medially as a transverse apodeme, not uniting anteriorly with apodemes 3–4; leg III with femur and genu separate or usually fused. LARVA. Prodorsum with three pairs of simple or slightly barbed setae, alveolar pits of v2 posterolaterad or laterad of normally developed setae v1; setae sc1 similar in form to v1, and more widely spaced from each other than sc2; setae sc2 similar in length or considerably longer and more attenuate than v1, sc1, but shorter than transverse interval between their insertions. Tergites C, D, EF each entire, their setae collectively slightly barbed; EF not extended posteriorly over HPs; cupules ia, im, ih usually discernible on D, E, HPs, respectively; HPs formed as ventroterminal capsule, wider than long ventromedially; setae h1 and h2 simple, short to exceedingly long, attenuate, similar in length; two pairs of ps setae short to exceedingly long, sometimes dissimilar in length. Coxisternal plates I and II fully fused medially; pits 1b and 2b spaced well laterad setae 1a and 2a, respectively. Coxisternal plates III with setae 3a and 3b attenuate, short to exceedingly long. Legs. Ambulacrum of leg I of adult female and male well-developed but lacking claws; that of larva similar, but sometimes with rudiments of paired claws. Leg I with tibia and tarsus of adult male and larva separate or more often fused, as in adult female. Ambulacra of legs II and III of adults and larva with empodium and symmetrically paired claws. Tarsi II and III lacking any spinelike projection over base of ambulacrum. Femora I and II lacking flanges. Legs I–III moderately short, stubby, with no segments elongated; on leg II, combined length of genu and tibia exceeding length of tarsus, and equal to that of femur. Leg III of adult male not enlarged; femur and genu coalesced into one segment, as in female, or separate. Leg IV of adult female elongate-cylindrical, with femorogenu somewhat longer than tibiotarsus, femorogenu with 0–2 setae, tibiotarsus with two setae; trochanter moderately thick, subtriangular or subquadrate, shorter than femorogenu, with or without seta. Leg IV of adult male short, stubby, with or without a short seta on trochanter; femorogenu shorter than trochanter, lacking flanges, and with zero or one short seta, Ge -vʹ; tarsus, if present, bearing three setae and small claw, or more usually absent or rudimentary such that prominent tibial solenidion and sometimes one or two tibial setae retained as terminal structures, with tibial seta vʹ short or attenuate. Number of setae and solenidia on legs in adult female as presented in Table 1 and as follows: femur I with 2–4 setae (lʹ rudimentary if present, d small or rudimentary if present, lʹʹ rudimentary or developed, vʹʹ developed); femur II with 1–3 setae (vʹʹ developed if present, d small or rudimentary if present, lʹ rudimentary if present); femur III without setae; genua I–II with or without four developed setae, genu III with three setae (lʹ, l″, vʹ) or without setae; tibiotarsus I with 5(+ φ 1)+5(+ ω) phaneres, lacking solenidion φ 2 and seta k″; tarsal seta s spine-like, setae tcʹtcʹʹ eupathidial, setae ftʹ-ft″ absent or indicated by alveolar vestiges; tibiae II–III with four setae; tarsus II with 4(+ ω) to 5(+ ω) setae, seta uʹ spine-like, often bifid apically, seta uʹʹ present or absent; tarsus III with 4 or 5 setae, seta uʹ spine-like, often bifid apically, seta uʹʹ present or absent. Adult male setation similar to female, but with variation noted in Table 1 and as follows: tarsus I with developed eupathidial setae ftʹ-ftʹʹ, tarsi II–III sometimes with uʹʹ absent in female but present in male. Larval setation similar to female, but with variation noted in Table 2 and as follows: tarsi II–III with seta pʹ present in larva but absent in adults; tarsus III with seta uʹʹ present in larva, usually retained in adult male and female; tarsi II–II without seta pvʹ, added in adult male and female. Etymology. The prefix podo- (Gr. foot) refers to the distinctive, well-developed ambulacra of these mites that bear small claws on legs II–III but not leg I (excepting the larvae of two species). Few other tarsonemid mites have lost the claws on leg I but retained them on legs II–III, and all other acarapine mites have large, well-developed claws on legs I–III. The suffix -tarsonemus refers to the thread-legged aspect of leg IV of adult females of this genus, which resembles that of other members of the family Tarsonemidae. The name is also intended to indicate the morphological and ecological similarities of these mites with those of both families Podapolipidae and Tarsonemidae. Remarks. The designation of setae on tarsi II–III is problematic, showing differences between species, but also in ontogeny and sex. All active instars have three consistent setae, the tectal pair tcʹ-tcʹʹ and the spine-like unguinal seta uʹ, plus the tarsal solenidion on leg II. As in other Tarsonemidae, seta pvʹ on tarsi II–III is expressed in the adult. In addition to these, the larvae of all species have a tiny apical anterolateral seta, designated pʹ, on tarsi II–III; and most species have a posterolateral seta, designated u″, on tarsi II and III. Seta pʹ is not expressed in the adult instar, and seta u″ is sometimes lost. As seta pʹ is present in the larva, then lost, and seta pvʹ is present only in adults, they could be considered homologous. However, seta pvʹ is situated subapically and is as long as the tectal setae, in contrast to the short, apically positioned larval seta pʹ. Seta uʹʹ is present in the larvae in all but the Bolivian species on tarsus II and in all species on tarsus III. However, seta u″ on tarsus II is lost in the adult male and female of the South African species and, on tarsus III, on the adult female of the Bolivian species; another species from Ecuador, for which the larva is unknown, lacks seta u″ on tarsus III in both sexes (Table 2). The interpretation of the eupathidial pair added to the adult male as (ft) is based on (a) no example of the other eupathidia being suppressed or added in a sexually dimorphic way (Lindquist 1986, p. 153); (b) their position near the solenidion and the absence of well developed eupathidia in a typically proral position; (c) the addition of (ft) as eupathidia in at least two genera of pseudotarsonemoidines, i.e. Pseudotarsonemoides and Ununguitarsonemus (Khaustov 1998; Lindquist 1986, respectively). The fusion of tibia and tarsus I in the larva and adult male of most species is also unusual, as these are separate in other Tarsonemidae. One Australian species retains the typical, plesiomorphic condition of separation in the adult male and larva, and the Indian species has separate segments in the larva but not the adult male. This progression from fusion only in the adult female to fusion in all life stages is unique in the Tarsonemidae. Finally, the structure of the respiratory system of Podotarsonemus is mostly uniform throughout the genus, with one notable exception (the species from India). In most species the system is paired, with each stigma leading to a single trachea that runs into a very lightly sclerotized atrium. Distally, each trachea emerges from the atrium and leads into a more sclerotized post-atrial structure that usually appears to have one edge more sclerotized than the other. Distally, two tracheae emerge from the post-atrial structure, but these are not always apparent (e.g., Figs 15A–C). The Indian species differs in having an unpaired atrium and a single, sclerotized tube emerging distally from each of its heavily sclerotized post-atrial structures (Fig. 17).

Podotarsonemus Seeman, Lindquist & Husband gen. nov. Type species: Podotarsonemus boliviensis Seeman, Lindquist & Husband sp. nov. Diagnosis. With attributes of tribe Podotarsonemini, augmented as follows: Palpi much reduced, without clear segmentation; apical palpal seta flanked medially by small hyaline triangular flange. Cheliceral stylets of larvae and adults curved from base to apex, with 0���2 teeth subapically. Adults and larvae with scapular setae sc 2 at most slightly longer than interval between their insertions, not exceedingly long, attenuated. Adult female prodorsal stigmata with main tracheal trunks leading internally to atrial chamber(s) and then to distinctively enlarged pair of post-atrial structures. Legs II���III of larvae and adults often lacking all genual setae (v��� always absent), and leg III lacking femoral setae. Larvae with setae on coxisternal plates III well developed, often attenuated, and with setae ps well-developed, sometimes attenuated, along with setae h. Male leg IV without flanges, femorogenu IV with maximum of one seta Ge- v��. In contrast to Acarapis, leg IV is slender in addition to characters already described for the tribe. Description. Gnathosoma. Form and structures of gnathosoma similar on larva, female, and male, though generally slightly larger on female. Capsule broadly oval in dorsoventral view, not beak-like anteriorly; palpcoxal setae present on adults. Dorsal gnathosomatic setae variably long, slender, smooth on adults, similarly formed on larva. Palpi directed anteriorly, short, stubby, with short but prominent, stout, pointed genual seta projected anteriorly and sometimes a smaller, inconspicuous femoral seta posterolaterally, these flanked anteroventrally by small, bluntly pointed, apical hyaline flange-like margin of femorogenual segment which covers rudimentary tibiotarsus with its terminal ventro-apical zone of alveolar vestiges of 3���5 small circular structures, probably rudiments of setae, solenidia and claw elements. Cheliceral stylets prominent, thick, variably protruding from capsule, smooth or with 1���2 teeth subapically, curved laterally in attachment to large basal levers oriented transversely across anterior one-third of capsule. Pharynx striated, muscular, with thinly sclerotized walls, enlarged to about one-third the width of gnathosomatic capsule, with paired gland-like structures indiscernible posteriorly. Idiosoma. ADULT FEMALE. Dorsal shielding unornamented or reticulated, imbricated. Dorsal setae collectively smooth or with small barbs on setae c1, d, e, f. Prodorsal shield not extended hood-like over gnathosoma. Stigmata opening closely anterolaterad setae v1; main tracheal trunks with enlarged, slightly sclerotized atrial sacs (sometimes fused into single sac), and also with slightly to heavily sclerotized post-atrial sacs, these entire (much as in Hemitarsonemus). Pits v2 obliquely aligned with setae v1 and sc2, and situated about one-quarter to midway along interval from v1 to sc2, sometimes at anterolateral invaginations of prodorsal shield. Setae sc2 long, attenuate, more widely spaced from each other, and longer, than v1, but not exceedingly long, at most slightly longer than transverse interval between their insertions, and inserted near midlength of prodorsal shield. Bothridia vestigial or absent, their sensilla sc1 reduced, slender, or absent. Tergite EF with setae e and f transversely aligned, similar in length, not grouped closely together. Tergite H with setae h short to exceedingly long, attenuate. Ventral shielding with apodemes 1 forming Y-shaped juncture with prosternal apodeme. Apodemes 2 fully or weakly connecting with prosternal apodeme. Prosternal apodeme developed posteriad level of apodemes 2 but not reaching posterior margin of coxisternal plates II. Sejugal apodeme reduced to a lateral fragment on either side. Apodemes 3 not extending laterad trochanters III, nor mediad setae 3a. Apodemes 4 reduced medially and not connected to each other, but extending posterolaterad setae 3b to base of trochanters IV. Poststernal apodeme absent. Coxisternal plates I and II with setae 1a and 2a short to exceedingly long, attenuated, similar in size. Anterior margin of coxisternal plates III straight or convex, if convex then sometimes extending over posterior edge of coxisternal plates II. Coxisternal setae 3a and 3b short to exceedingly long, attenuated, similar in size. Bases of legs IV well spaced, separated by interval of three to four widths of trochanter IV; posterior margin of coxisternal plates IV weakly formed, not formed into tegular projection. Pseudanal setae ps exceedingly long, attenuate. ADULT MALE. Dorsal shielding unornamented. Dorsal setae collectively smooth, much as in female, but if attenuated, then less extremely so. Prodorsum with three pairs of setae, including short but well developed sc1; setae sc2 attenuate but not exceedingly long, no longer than transverse interval between their insertions; setae v2 lacking but vestiges of their alveoli indicated by a pair of pits. Plate CD with setae arranged in two or three transverse rows, pair c2 anteriorly, d posteriorly, and c1 either at same level between c2 or at level between (and often obliquely aligned with) c2 and d. Plate EF with setae e and f more or less aligned transversely. Genital capsule with adhesive copulatory flanges reduced or absent. Sejugal apodeme evident laterally but indistinct medially. Coxisternal plates III and IV partly or entirely separated from each other medially by soft cuticle, such that apodemes 3 and 4 not reaching each other medially and poststernal apodeme absent; apodemes 5 moderately to weakly formed on either side at bases of legs IV, sometimes connected to each other medially as a transverse apodeme, not uniting anteriorly with apodemes 3���4; leg III with femur and genu separate or usually fused. LARVA. Prodorsum with three pairs of simple or slightly barbed setae, alveolar pits of v2 posterolaterad or laterad of normally developed setae v1; setae sc1 similar in form to v1, and more widely spaced from each other than sc2; setae sc2 similar in length or considerably longer and more attenuate than v1, sc1, but shorter than transverse interval between their insertions. Tergites C, D, EF each entire, their setae collectively slightly barbed; EF not extended posteriorly over HPs; cupules ia, im, ih usually discernible on D, E, HPs, respectively; HPs formed as ventroterminal capsule, wider than long ventromedially; setae h1 and h2 simple, short to exceedingly long, attenuate, similar in length; two pairs of ps setae short to exceedingly long, sometimes dissimilar in length. Coxisternal plates I and II fully fused medially; pits 1b and 2b spaced well laterad setae 1a and 2a, respectively. Coxisternal plates III with setae 3a and 3b attenuate, short to exceedingly long. Legs. Ambulacrum of leg I of adult female and male well-developed but lacking claws; that of larva similar, but sometimes with rudiments of paired claws. Leg I with tibia and tarsus of adult male and larva separate or more often fused, as in adult female. Ambulacra of legs II and III of adults and larva with empodium and symmetrically paired claws. Tarsi II and III lacking any spinelike projection over base of ambulacrum. Femora I and II lacking flanges. Legs I���III moderately short, stubby, with no segments elongated; on leg II, combined length of genu and tibia exceeding length of tarsus, and equal to that of femur. Leg III of adult male not enlarged; femur and genu coalesced into one segment, as in female, or separate. Leg IV of adult female elongate-cylindrical, with femorogenu somewhat longer than tibiotarsus, femorogenu with 0���2 setae, tibiotarsus with two setae; trochanter moderately thick, subtriangular or subquadrate, shorter than femorogenu, with or without seta. Leg IV of adult male short, stubby, with or without a short seta on trochanter; femorogenu shorter than trochanter, lacking flanges, and with zero or one short seta, Ge -v��; tarsus, if present, bearing three setae and small claw, or more usually absent or rudimentary such that prominent tibial solenidion and sometimes one or two tibial setae retained as terminal structures, with tibial seta v�� short or attenuate. Number of setae and solenidia on legs in adult female as presented in Table 1 and as follows: femur I with 2���4 setae (l�� rudimentary if present, d small or rudimentary if present, l���� rudimentary or developed, v���� developed); femur II with 1���3 setae (v���� developed if present, d small or rudimentary if present, l�� rudimentary if present); femur III without setae; genua I���II with or without four developed setae, genu III with three setae (l��, l���, v��) or without setae; tibiotarsus I with 5(+ �� 1)+5(+ ��) phaneres, lacking solenidion �� 2 and seta k���; tarsal seta s spine-like, setae tc��tc���� eupathidial, setae ft��-ft��� absent or indicated by alveolar vestiges; tibiae II���III with four setae; tarsus II with 4(+ ��) to 5(+ ��) setae, seta u�� spine-like, often bifid apically, seta u���� present or absent; tarsus III with 4 or 5 setae, seta u�� spine-like, often bifid apically, seta u���� present or absent. Adult male setation similar to female, but with variation noted in Table 1 and as follows: tarsus I with developed eupathidial setae ft��-ft����, tarsi II���III sometimes with u���� absent in female but present in male. Larval setation similar to female, but with variation noted in Table 2 and as follows: tarsi II���III with seta p�� present in larva but absent in adults; tarsus III with seta u���� present in larva, usually retained in adult male and female; tarsi II���II without seta pv��, added in adult male and female. Etymology. The prefix podo- (Gr. foot) refers to the distinctive, well-developed ambulacra of these mites that bear small claws on legs II���III but not leg I (excepting the larvae of two species). Few other tarsonemid mites have lost the claws on leg I but retained them on legs II���III, and all other acarapine mites have large, well-developed claws on legs I���III. The suffix -tarsonemus refers to the thread-legged aspect of leg IV of adult females of this genus, which resembles that of other members of the family Tarsonemidae. The name is also intended to indicate the morphological and ecological similarities of these mites with those of both families Podapolipidae and Tarsonemidae. Remarks. The designation of setae on tarsi II���III is problematic, showing differences between species, but also in ontogeny and sex. All active instars have three consistent setae, the tectal pair tc��-tc���� and the spine-like unguinal seta u��, plus the tarsal solenidion on leg II. As in other Tarsonemidae, seta pv�� on tarsi II���III is expressed in the adult. In addition to these, the larvae of all species have a tiny apical anterolateral seta, designated p��, on tarsi II���III; and most species have a posterolateral seta, designated u���, on tarsi II and III. Seta p�� is not expressed in the adult instar, and seta u��� is sometimes lost. As seta p�� is present in the larva, then lost, and seta pv�� is present only in adults, they could be considered homologous. However, seta pv�� is situated subapically and is as long as the tectal setae, in contrast to the short, apically positioned larval seta p��. Seta u���� is present in the larvae in all but the Bolivian species on tarsus II and in all species on tarsus III. However, seta u��� on tarsus II is lost in the adult male and female of the South African species and, on tarsus III, on the adult female of the Bolivian species; another species from Ecuador, for which the larva is unknown, lacks seta u��� on tarsus III in both sexes (Table 2). The interpretation of the eupathidial pair added to the adult male as (ft) is based on (a) no example of the other eupathidia being suppressed or added in a sexually dimorphic way (Lindquist 1986, p. 153); (b) their position near the solenidion and the absence of well developed eupathidia in a typically proral position; (c) the addition of (ft) as eupathidia in at least two genera of pseudotarsonemoidines, i.e. Pseudotarsonemoides and Ununguitarsonemus (Khaustov 1998; Lindquist 1986, respectively). The fusion of tibia and tarsus I in the larva and adult male of most species is also unusual, as these are separate in other Tarsonemidae. One Australian species retains the typical, plesiomorphic condition of separation in the adult male and larva, and the Indian species has separate segments in the larva but not the adult male. This progression from fusion only in the adult female to fusion in all life stages is unique in the Tarsonemidae. Finally, the structure of the respiratory system of Podotarsonemus is mostly uniform throughout the genus, with one notable exception (the species from India). In most species the system is paired, with each stigma leading to a single trachea that runs into a very lightly sclerotized atrium. Distally, each trachea emerges from the atrium and leads into a more sclerotized post-atrial structure that usually appears to have one edge more sclerotized than the other. Distally, two tracheae emerge from the post-atrial structure, but these are not always apparent (e.g., Figs 15A���C). The Indian species differs in having an unpaired atrium and a single, sclerotized tube emerging distally from each of its heavily sclerotized post-atrial structures (Fig. 17)., Published as part of Seeman, Owen D., Lindquist, Evert E. & Husband, Robert W., 2018, A new tribe of tarsonemid mites (Trombidiformes: Heterostigmatina) parasitic on tetrigid grasshoppers (Orthoptera), pp. 1-54 in Zootaxa 4418 (1) on pages 7-10, DOI: 10.11646/zootaxa.4418.1.1, http://zenodo.org/record/1242435, {"references":["Lindquist, E. E. (1986) The world genera of Tarsonemidae (Acari: Heterostigmata): A morphological, phylogenetic, and systematic revision, with a reclassification of family group taxa in Heterostigmata. Memoirs of the Entomological Society of Canada, 136, 1 - 517. https: // doi. org / 10.4039 / entm 118136 fv","Khaustov, A. A. (1998) A review of mites of the genus Pseudotarsonemoides (Acari: Heterostigmata: Tarsonemidae) of Crimea, with description of two new species. Acarina, 6, 25 - 30."]}

Podotarsonemus australiensis Seeman, Lindquist & Husband sp. nov. Figures 1���6 Type material. Holotype. Adult female. AUSTRALIA, Spear Creek, Mt Molloy, Far North Queensland, 22 Jun 1975, G.B. Monteith, ex Loxilobus sp. (Tetrigidae) (Fig. 29A). In QM. Paratypes. 7 females, 1 male, 2 larvae, same data as holotype. All in QM except 1 female in each of CNCI and UMMZ. Non-type material. 1 female, PAPUA NEW GUINEA, Upper Gusap River, near Butemu, Madang Dist., Oct 1965, R. Pullen, 3500 ��� alt., ex Scelimena erimeta (G��nther) (Tetrigidae). In QM. Diagnosis. Female and male: Palps with 1 seta; chelicerae with 2 teeth; prodorsal plate entirely punctate; femur I with rudimentary setae l�� and d; femur II with tiny seta d, lacking l�� and v���; genua II���III with 3 setae; tarsi II���III with 5 setae, seta u���� present; trochanter IV with seta. Female: Setae sc1 absent; vestigial setae v2 expressed as anterolateral invaginations; coxisternal plates III���IV separated medially by longitudinal striae; setae h attenuate, 40���51 long; femorogenu IV with 2 setae; tracheal atria separate, post-atrial sac lightly sclerotized, not punctate. Male: Genital capsule without copulatory flanges; setae h2 developed, length 4���6; femorogenu IV with 1 seta; tibia IV with solenidion and attenuate seta; tarsus IV terminal, bearing 3 setae and claw. Male and larva: Leg I with tibia and tarsus I separate. Larva: Setae h1-2, ps2 attenuate, setae ps1 short; leg I with 2 small claws. Description. ADULT FEMALE (Figures 1���2, 5A, B, n = 8 measured) Gnathosoma. Length 63���69, width 58���80. Palps 10���12, with 1 seta (2 long). Cheliceral stylets 25���27, with 2 teeth. Pharynx length 30���33, width 17. Dorsal setae ch 30���35, ventral setae su 6���14, palpcoxal setae e 7���13. Idiosoma. Length 255���310, width 165���197. Tracheal atria separate, post-atrial sac lightly sclerotized, not punctate, terminal tracheae not visible. Prodorsal plate with punctate sculpturing, 102���104 long, 130���140 wide; with 2 pairs of setae: v1 22 ���25, thickened, tapering, with barbs, sc2 66���75 and vestigial v2 expressed as anterolateral invaginations. Distance between setae v1-v1 55 ���60, v2-v2 62 ���70, sc2-sc2 72��� 80, v1 -sc2 34. Plates C, D, EF and H with punctate sculpturing, plate H heavily so. Setae c1, d, e, f tapering, with small barbs, c 1 19���27, c2 54���64, d 21���27, e 20���24, f 24���28. Distance between setae c1-c1 97���110, c2-c2 148���165, c1-c 2 22���25, d-d 78���87, e-e 83���87, f-f 35���44. Setae h tapering, 50���51. Distance between setae h-h 25���31. Cupules ia present, tiny, im absent, ih present. Setae ps attenuate> 250, distance between setae ps-ps 12���14. Coxisternal setae short to attenuate, 1a 11���12, 2a > 140, 3a > 230, 3b > 150. Distance between setae 1a-1a 17���20, 2a-2a 42���46, 3a-3a 10��� 12, 3b-3b 14���24; alveolar vestiges of setae 1b, 2b barely discernible. Coxisternal plates III���IV separated by longitudinal striae; coxisternal plates III convex anteriorly, overlapping posterior margin of coxisterna II. Legs. Leg lengths: I 56 ���62, II 62 ���66, III 45 ���49. Leg IV Fe-Ge length 29���32, Ti-Ta length 14���16. Leg III, femur and genu fused. Femur I, l��, d rudimentary, l���� 1���2, v���� 10���14; genu I l�� 2���3, l���� 2���3, v�� 2���4, v���� 3���4; tibia and tarsus I fused, tibial setae d 30���35, l�� 4���6, l���� 4���5, v�� 8���10, v���� 4���7, solenidion �� 3; tarsal setae tc�� 14���15, tc���� 12���14, pv�� 7���10, pv���� 6���7, s spine-like, 8���9, solenidion �� 7. Femur II d 1���2; genu II l�� 3���4, l���� 4���5, v�� 3���5; tibia II d 20���22, l�� 7���8, v�� 15���16, v���� 15���18; tarsus II pv�� 12���16, tc�� 12���15, tc���� 17���21, u�� spine-like, 8���10, u���� 3���5, solenidion �� 6���7. Genu III l�� 3���4, l���� 5, v�� 3���4; Tibia III d 14���15, l�� 17, v�� 7���8, v���� 14���16; tarsus III pv�� 12���16, tc�� 16���18, tc���� 18, u�� spine-like, 8, u���� 2���3. Trochanter IV with ventral seta v�� 10���12; femorogenu IV with proximal femoral seta v�� 5���6, distal genual seta v�� 18���23; tibiotarsus with attenuate ventral tibial seta v�� > 100 and terminal tarsal seta tc��� > 350. ADULT MALE (Figs 3���4, 5C, n = 1) Gnathosoma. Length 48, width 54. Palps 8. Cheliceral stylets 19, with 2 teeth. Pharynx length 28, width 14. Dorsal setae 20, ventral setae 5, palpcoxal setae 5. Idiosoma. Length 200, width 160. Prodorsal plate punctate, 82 long, 110 wide; with 3 pairs of setae (v1 24, sc1 12, sc2 55) and vestigial v2 expressed as anterolateral invaginations. Distance between setae v1-v1 37, v2���v2 48, sc1-sc1 70, sc2-sc2 69, v1 -sc2 50. Plate CD punctate. Setae c1 tapering, with small barbs, 24, c2 fine, 44, d with small barbs, 27 (asymmetrical, extra d expressed on right hand side); setae c 1 in nearly transverse alignment with c2. Plate EF punctate, e 17, f 24, with small barbs. Distance between setae c1-c1 88, c2-c2 144, c1-c2 27, d-d 63, e- e 34, f-f 25. Cupules ia, im, ih indiscernible or absent. Genital capsule 47 long, 38 wide, bearing setae h1 6 long and alveolar remnants of h2; ventral flanges absent. Coxisternal setae short to attenuate 1a 14, 2a > 60, 3a> 80, 3b > 10 (broken). Distance between setae 1a-1a 20, 2a-2a 34, 3a-3a 8, 3b-3b 38; alveolar vestiges of setae 1b, 2b indiscernible or absent; setae 1a just posterior to ap1, setae 2a far posterior to ap2. Apodemes: ap2 almost reaching medially obsolete ap sj; ap5 well developed, fused, extending 45���50 anteriorly from base. Legs. Leg lengths: I 55, II 58, III 54, IV 45 ���51. Leg III, femur and genu separate. Femur I, l��, d rudimentary, l���� 2, v���� 10; genu I l�� 3, l���� 3, v�� 3, v���� 4; tibia and tarsus I separate, tibial setae d 28, l�� 6, l���� 5, v�� 5, v���� 11, solenidion �� 3; tarsal setae ft�� 8, ft���� 8, tc�� 14, tc���� 14, pv�� 10, pv���� 8, s spine-like, 8, solenidion �� 7. Femur II d 2; genu II l�� 4, l���� 6, v�� 3; tibia II d 14, l�� 15, v�� 7, v���� 20; tarsus II pv�� 10, tc�� 15, tc���� 20, u�� spine-like, 9, u���� 6, solenidion �� 10. Genu III l�� 3, l���� 5, v�� 5; tibia III d 19, l�� 18, v�� 8, v���� 13; tarsus III pv�� 13, tc�� 18, tc���� 20, u�� spine-like, 9, u���� 1. Trochanter IV with ventral seta 7 long; femorogenu IV with proximal seta 2; tibia IV with solenidion �� 17 and ventral seta attenuate,> 120; tarsus IV terminal, with u�� 4, tc���� 2, pv���� rudimentary, and claw 3. LARVA (Fig. 6, n = 2) Gnathosoma. Length 58���50, width 43���56. Palps 10���11. Cheliceral stylets 24���25, with 2 teeth. Pharynx length 26���30, width 14���15. Dorsal setae 7���10, ventral setae 4���5. Idiosoma. Length 255���350, width 142���225. Prodorsal plate smooth, 70���76 long, 111���125 wide; with 3 pairs of tapering, weakly barbed setae (v1 10, sc1 8���9, sc 2 20���21) and vestigial v2. Distance between setae v1-v1 35, v2-v2 56, sc1-sc1 77���79, sc2-sc2 68��� 70, v1 -sc2 24. Plates C, D, EF punctate. Setae tapering, barbed, c 1 15���17, c 2 13���15, d 16���18, e 16, f 20���21. Distance between setae c1-c1 76, c2-c2 127, c1-c 2 30���33, d-d 50���52, e-e 50���53, f-f 23���26. Plate H punctate, setae h1 attenuate,> 160, h2 ventral, attenuate, > 200. Distance between setae h1 24. Cupules ia, im present, ih indiscernible or absent. Setae ps1 fine, 10, ps2 attenuate 35���55. Coxisternal setae 1a 5���10, 2a 35���40, 3a attenuate,> 160, 3b 39. Distance between setae 1a-1a 17���21, 2a-2a 45, 3a-3a 32, 3b-3b 39; alveolar vestiges of setae 1b, 2b present. Legs. Femur I d, l��, l���� rudimentary, v���� 6���8; genu I l�� rudimentary, l���� 1, v�� rudimentary, v���� 1; tibia and tarsus I separate, tibial setae d 10���12, l�� 3���5, l���� 3, v�� 3���4, v���� 7���10, solenidion �� 3, tarsal setae tc�� 11���13, tc���� 10���12, pv�� 5, pv���� 3���5, s spine-like, 5, solenidion �� 5���6. Femur II d rudimentary; tibia II d 7���9, l�� 4���5, v�� 12, v���� 8���11; tarsus II tc�� 10, tc���� 11, p�� 1, u�� spine-like, 6, u���� 1��� 2, solenidion �� 4���5. Tibia III d 7���8, l�� 4���5, v�� 9���12, v���� 10���12; tarsus III pv�� 11, tc�� 10���14, tc���� 6, p�� 1���2, u�� spine-like, 6���7, u���� 1. Leg I with 2 small claws. Etymology. The species name australiensis refers to the name of the country where the type specimens were collected. Remarks. Podotarsonemus can be divided into two species groups. The first group (australiensis species group, including P. australiensis, P. indicus and P. zuluensis) all retains seta v�� on trochanter IV, has females with longitudinal striae between coxisterna III���IV and setae h not attenuated, and males that retain at least a rudimentary tarsus on leg IV. In contrast, females of the second group (boliviensis species group, including P. boliviensis, P. ecuadorensis, P. nipponicus, P. queenslandensis) have lost seta v�� on trochanter IV, have no longitudinal striae between coxisterna III���IV, have attenuate setae h, and males lack tarsus IV. The australiensis species group is possibly artificial because it is based on putative plesiomorphies. Of all its members, P. australiensis is perhaps the most plesiotypic species. In addition to the abovementioned features of the australiensis species group, P. australiensis is the only species that retains setae on genua II���III and has males with a divided tibia and tarsus I, tarsal structures on leg IV and a well-developed h seta. Despite these plesiomorphic attributes, P. australiensis also has the apomorphic loss of setae sc1 on the adult female, which is shared with P. queenslandensis and P. zuluensis. This feature is probably a homoplasious loss, as both these species belong in separate species groups. Another apomorphic loss is seta l�� on femur II (absent in all species except P. boliviensis and P. ecuadorensis) and the loss of seta v��� on femur II, which is autapomorphic in the Acarapinae. Exceptionally for the Tarsonemidae, the setae on genua II���III are absent in the larva but present in the adult. The delay in expression of these setae is an example of in-process evolutionary ontogenetic suppression, and while unusual in the Tarsonemidae, such ontogenetic delays are often reported for leg setae in other major taxa (e.g., see Beard et al. (2014) on the Tenuipalpidae). Curiously, considerable leg setal reductions also occur in the larvae of species of Acarapis, which retain just one tibial and two tarsal setae. However, adults of all three species retain a near-fully developed leg chaetome for the Acarapinae; hence the ontogenetic suppression of these setae is not connected with evolutionary trends towards simplification of leg setae. Instead, larvae of Acarapis show reductions in their gross leg morphology, suggesting their reductive trend is closely related to the apparent immobility of their larval stage. The material examined includes one non-type female from Papua New Guinea that we have included under P. australiensis. The specimen matches the Australian specimens reasonably well but is in poor condition, lacking setae sc2, c1, c2, d and e. However, many of its leg setae are longer, such as femur I d (2 versus rudimentary), and tibia I d (45 versus 30���35) and v��� (24 versus 4���7). Its host genus is also different: although both are Scelimeninae, Scelimena belongs to the Scelimenini while Loxilobus belongs to the Criotettigini. The differences in leg setal lengths and different host tribes suggest that this mite may be a different species, but more collections are required to test this hypothesis., Published as part of Seeman, Owen D., Lindquist, Evert E. & Husband, Robert W., 2018, A new tribe of tarsonemid mites (Trombidiformes: Heterostigmatina) parasitic on tetrigid grasshoppers (Orthoptera), pp. 1-54 in Zootaxa 4418 (1) on pages 10-18, DOI: 10.11646/zootaxa.4418.1.1, http://zenodo.org/record/1242435, {"references":["Beard, J. J., Seeman, O. D. & Bauchan, G. R. (2014) Tenuipalpidae (Acari: Trombidiformes) from Casuarinaceae (Fagales). Zootaxa, 3778 (1), 1 - 157. https: // doi. org / 10.11646 / zootaxa. 3778.1.1"]}

Subfamily Acarapinae Schaarschmidt, 1959 Type genus: Acarapis Hirst, 1921 Description. As follows Lindquist (1986, p. 244), with following modifications as indicated in italics as variation noted to accommodate the genus Podotarsonemus and tribe Podotarsonemini. Gnathosoma. Palpcoxal setae present in adults (Podotarsonemini), or lacking (Coreitarsonemini, Acarapini). Cheliceral stylets robust, nearly straight or curved along their length (Podotarsonemini), attached to conspicuous basal levers. Idiosomatic dorsum. Prodorsal stigmata of adult female situated anterolaterad of vertical setae, usually on protuberances (Acarapini, Coreitarsonemini, some Podotarsonemini). Prodorsal bothridia lacking, their sensilla a slender seta (most Podotarsonemini), vestigial or lacking (Acarapini, Coreitarsonemini, few Podotarsonemini) on adult female, and corresponding setae sc1 a slender seta (Podotarsonemini), minute, vestigial, or lacking (Acarapini, Coreitarsonemini) on larva and adult male. Prodorsum of adult male also lacking setae v2, such that only two (v1, sc2 ��� Acarapini, most Coreitarsonemini) or three (v1, sc1, sc2 ���Podotarsonemini, few Coreitarsonemini) pairs of setae usually present. Plate EF of adult male with one or two pairs of setae, sometimes retaining e in addition to f and cupules im. Genital capsule HPs of adult male with pliable adhesive copulatory flanges well-developed, reduced or absent. Idiosomatic venter. Coxisternal plates I and II of adult each with one pair of setae (1a and 2a, respectively) but alveolar remnants of setae 1b and 2b present or lacking; plates I and II of larva with setae 1a and 2a present (Podotarsonemini) or vestigial or lacking (Acarapini, Coreitarsonemini), and alveolar remnants of 1b and 2b present (Podotarsonemini) or lacking (Acarapini, Coreitarsonemini). Metapodosomal venter of adult with no setae added to basic larval complement, such that coxisternal plates III with two pairs of setae, and plates IV lacking setae; plates III of larva with the basic two pairs of setae, 3a and 3b, present or lacking. Tegula of adult female either short and broadly truncate, or not developed. Aggenital plate of adult female lacking setae. Idiosoma of larva moderately (Podotarsonemini) to greatly (Acarapini, Coreitarsonemini) distensible due to extensive plication of membranous cuticle between various plates. Legs. Leg I of adult female with single, normal claw on pretarsus (Acarapini, Coreitarsonemini) or claw absent on well-developed ambulacrum (Podotarsonemini); adult male with claw single (Acarapini), paired (Coreitarsonemini) or absent on well-developed ambulacrum (Podotarsonemini); larva with claw paired (Acarapini, Coreitarsonemini, few Podotarsonemini) or absent on well-developed ambulacrum (most Podotarsonemini). Leg III of adult female with trochanter no longer than femorogenu, and retaining a somewhat subtriangular form with articulated mobility similar to that of larva and adult male. Leg III of adult male with femur and genu fused or separate. Leg IV of adult female with zero to two (Podotarsonemini) or two or three setae on femorogenu (a second genual seta, l����, sometimes present) (Acarapini, Coreitarsonemini), and with two or rarely three setae on tibiotarsus (a second long tibial seta, v���, rarely present). Leg IV of adult male with or without a seta on trochanter; with tibia and tarsus separate (few Coreitarsonemini, few Podotarsonemini), fused (most Coreitarsonemini) or tarsus absent (Acarapini, most Podotarsonemini); tarsal setae present, vestigial, or lacking; tarsal claw vestigial or usually lacking. Number of setae and solenidia on femur, genu, tibia, and tarsus variable, but consistent within Acarapini and Coreitarsonemini, more variable in Podotarsonemini; see descriptions of tribes in Lindquist (1986). Tibial sensory cluster of leg I of adult incomplete, lacking �� 1 (Acarapini, Coreitarsonemini) and sometimes �� 2 (Coreitarsonemini) but retaining seta k����, or retaining �� 1 and lacking k���� (Podotarsonemini). Tarsal setation of leg I lacking pl�� and sometimes pl����; on adult female setae ft��-ft���� lacking, on adult male setae ft��-ft���� well developed (Podotarsonemini) or lacking (Acarapini, Coreitarsonemini); unguinals u��-u����, when present, well separated from each other. Femur I setation complete (Acarapini, Coreitarsonemini, some Podotarsonemini) or incomplete (some Podotarsonemini) but seta l���� always present. Tarsus II lacking dorsoproximal spinelike seta pl����. Tarsi II and III with seta u��� present or absent; with unguinal u�� spinelike or slender. Tarsus III with seta pv�� expressed on adult of both sexes (exceptionally absent on male) but not on larva. Leg III with one femoral seta (Acarapini, Coretarsonemini) or femoral setae absent (Podotarsonemini). Larvae and adults parasitic on insects., Published as part of Seeman, Owen D., Lindquist, Evert E. & Husband, Robert W., 2018, A new tribe of tarsonemid mites (Trombidiformes: Heterostigmatina) parasitic on tetrigid grasshoppers (Orthoptera), pp. 1-54 in Zootaxa 4418 (1) on page 5, DOI: 10.11646/zootaxa.4418.1.1, http://zenodo.org/record/1242435, {"references":["Schaarschmidt, L. (1959) Systematik und okologie der Tarsonemiden. Beitrage zur Systematik und Okologie mitteleuropaischer Acarina I, 2 (5), 713 - 823.","Hirst, S. (1921) On some new or little-known Acari, mostly parasitic in habit. Proceedings of the Zoological Society of London, 1921, 357 - 328. https: // doi. org / 10.1111 / j. 1096 - 3642.1921. tb 03268. x","Lindquist, E. E. (1986) The world genera of Tarsonemidae (Acari: Heterostigmata): A morphological, phylogenetic, and systematic revision, with a reclassification of family group taxa in Heterostigmata. Memoirs of the Entomological Society of Canada, 136, 1 - 517. https: // doi. org / 10.4039 / entm 118136 fv"]}

Podotarsonemus zuluensis Seeman, Lindquist & Husband sp. nov. Figures 25–27 Type material. Holotype. Adult female. SOUTH AFRICA, Natal, Ntumeni, D. Otte, 7 Mar. 1965, ex probable Pelusca sp. (Tetrigidae), specimen # 1. RWH 10VIII2010 -12. In UMMZ. Paratypes. 7 females, 1 male, 5 larvae, as follows. 4 females, 2 larvae, same data as holotype. 4 females, 1 male, 4 larvae, same data as holotype except host specimen #2. In UMMZ except 1 female, 1 larva in each of CNCI and QM. Diagnosis. Female and male: Palps with 2 setae (1 minute); chelicerae without teeth; prodorsal plate entirely punctate; femora I–II without setae d, lʹ; tibia I seta l″ short (3–5); genua II–III bare; tarsus II with 4 setae, seta uʹʹ absent; tarsus III with 5 setae, seta uʹʹ present; trochanter IV with seta. Female: Setae sc1 present; vestigial setae v2 expressed as anterolateral invaginations; coxisternal plates III–IV separated by longitudinal striae; setae h short, 5– 7 long; tracheal atria separate, post-atrial sac lightly sclerotized, not punctate. Male: Genital capsule without copulatory flanges; setae h1 rudimentary, like h2; tibia IV with solenidion, without setae; tarsus IV rudimentary, subterminal. Male and larva: Leg I with tibia and tarsus fused. Larva: setae h1-2 and ps1-2 short, 5–12 long leg I without claws. Description. ADULT FEMALE (Fig. 25, n = 7 measured) Gnathosoma. Length 36–45, width 40–41. Palps 6–7, with 2 setae (2–3 and 1 long). Cheliceral stylets 18–22, without teeth. Pharynx length 25–27, width 11–14. Dorsal setae ch 19–21, ventral setae su 13–19, palpcoxal setae e 6–8. Idiosoma. Length 205–260, width 145–165. Tracheal atria separate, post-atrial sac lightly sclerotized, not punctate, with two small prongs indicating separate terminal tracheae. Prodorsal plate with punctate sculpturing, 76–90 long, 93–104 wide; with 3 pairs of setae: v 1 14–16, thickened, blunt, with barbs, sc 1 11–14, sc2 51–60, and vestigial v2 expressed as anterolateral invaginations. Distance between setae v1-v 1 24–27, v2-v2 32 –34, sc1-sc1 37–39, sc2-sc2 70– 81, v1 -sc 2 17–22. Plates C, D, EF and H with punctate sculpturing. Setae c1, d, e, f blunt, with small barbs, c 1 13–15, c2 37–42, d 13–15, e 7–10, f 10–11. Distance between setae c1-c1 56–61, c2-c2 107–115, c1-c 2 22–26, d-d 26–33, e-e 40–50, f-f 16–23. Setae h short, 5–7. Distance between setae h-h 22–25. Cupules ia, im, ih present. Setae ps attenuate> 60, distance between setae ps-ps 11–13. Coxisternal setae short to attenuate, 1a 13–19, 2a 13–25, 3a > 80, 3b > 45. Distance between setae 1a-1a 15–18, 2a-2a 31–39, 3a-3a 13–17, 3b-3b 25–34; alveolar vestiges of setae 1b present, 2b not evident. Coxisternal plates III–IV separated by longitudinal striae; coxisternal plates III convex anteriorly, sometimes just overlapping posterior margin of coxisterna II. Legs. Leg lengths: I 45 –46, II 41 –45, III 41 –45. Leg IV Fe-Ge length 17–20, Ti-Ta length 15–17. Leg III, femur and genu fused. Femur I lʹʹ rudimentary, vʹʹ 18–24; genu I lʹ 3, lʹʹ 2–3, vʹ 4–5, vʹʹ 4–5; tibia and tarsus I fused, tibial setae d 24– 33, lʹ 7–8, lʹʹ 3–5, vʹ 5–9, vʹʹ 8–11, solenidion φ 3, tarsal setae tcʹ 12–13, tcʹʹ 11–12, pvʹ 8–9, pvʹʹ 5–6, s spine-like, 6, solenidion ω 7–8. Femur II vʹʹ 11–14; tibia II d 15–31, lʹ 6–8, vʹ 15–19, vʹʹ 17–19; tarsus II tcʹ 16–17, tcʹʹ 18–23, pvʹ 14–19, uʹ spine-like, 5–6, solenidion ω 3–4. Tibia III d 23–25, lʹ 7–11, vʹ 18–23, vʹʹ 17–22; tarsus III tcʹ 17–20, tcʹʹ 22–27, pvʹ 16–18, uʹ spine-like, 5–6, uʹʹ 3–4. Trochanter IV with ventral seta 10–13; femorogenu IV bare; tibiotarsus IV with ventral tibial seta 14–18 and terminal tarsal seta> 240. ADULT MALE (Fig. 26, n = 1 measured) Gnathosoma. Damaged. Cheliceral stylets 18, without teeth. Pharynx length 27, width 10. Dorsal setae 13, ventral setae 6, palpcoxal setae 8. Idiosoma. Length 139, width 108. Prodorsal plate punctate, 57 long, 81 wide; with 3 pairs of barbed setae (v1 8, sc1 6, sc2 21) and vestigial v2. Distance between setae v1-v1 25, v2-v2 31, sc1-sc1 57, sc2-sc2 26, v1 -sc2 32. Plate CD punctate. Setae with small barbs, setae c1, 8, c2 9, d 8; setae c 1 in transverse alignment with setae c2. Plate EF punctate, e 5, f 6. Distance between setae c1-c1 53, c2-c2 96, c1-c2 21, d-d 8, e-e 21, f-f 8. Cupules ia, im present. Genital capsule 26 long, 28 wide, bearing alveolar remnants of h1, h2; ventral flanges absent. Coxisternal setae small, 1a 3, 2a 8, 3a 13, 3b 10. Distance between setae 1a-1a 14, 2a-2a 36, 3a-3a 19, 3b-3b 20; setae 1a just posterior to ap1, setae 2b far posterior to ap2. Apodemes: ap2 almost reaching medially obsolete ap sj; ap5 small, extend 12 from base, not fused medially. Legs. Leg lengths: I 36, II 35, III 32, IV 21. Leg III, femur and genu separate. Femur I, lʹʹ rudimentary, vʹʹ 12; genu I lʹ 3, lʹʹ 3, vʹ 2, vʹʹ 2; tibia and tarsus I fused, tibial setae d 21, lʹ 3, lʹʹ 2, vʹ 2, vʹʹ 5, solenidion φ 2, tarsal setae ftʹ 9, ftʹʹ 8, tcʹ 10, tcʹʹ 9, pvʹ 6, pvʹʹ 5, s spine-like, 5, solenidion ω 6. Femur II vʹʹ 13; tibia II d 15, lʹ 3, vʹ 12, vʹʹ 13; tarsus II tcʹ 7, tcʹʹ 14, pvʹ 7, uʹ spine-like, 5, solenidion ω 6. Tibia III d 14, lʹ 4, vʹ 10, vʹʹ 8; tarsus III tcʹ 15, tcʹʹ 14, pvʹ 12, uʹ spine-like, 5, uʹʹ 3. Trochanter IV with ventral seta 3 long; femorogenu IV bare; tibia IV solenidion φ 7; tarsus IV rudimentary, subterminal, with vestigial seta. LARVA (Fig. 27, n = 4 measured) Gnathosoma. Length 36–44, width 39–46. Palps 5. Cheliceral stylets 19–22, without teeth. Pharynx length 20–26, width 10–12. Dorsal setae 10–11, ventral setae 6–8. Idiosoma. Length 190–195, width 115–120. Prodorsal plate smooth, 55 long, split in all specimens; with 3 pairs of weakly barbed setae (v1 5–7, sc1 5–6, sc 2 6–10) and vestigial v2. Distance between setae v1-v 1 13–25, v2- v2 24, sc1-sc1 59, sc2-sc2 39, v1 -sc 2 24–25. Plates C, D, EF punctate. Setae tapering, barbed, c 1 8–10, c2 8, d 9– 10, e 8–9, f 8. Distance between setae c1-c1 51–54, c2-c2 96–100, c1-c 2 24–26, d-d 22–27, e-e 32–34, f-f 14–16. Plate H punctate, setae h1 short, 7–8, h2 ventral, 5–12 (unpaired h 3 in one specimen). Distance between setae h 1 17–21. Cupules ia, im, ih present. Setae ps1 fine, 8–10, ps2 fine, 10. Coxisternal setae 1a 4–5, 2a 4–7, 3a attenuate, 10–17, 3b 5–10. Distance between setae 1a-1a 13–26, 2a-2a 27–32, 3a-3a 34, 3b-3b 26. Legs. Femur I, lʹʹ rudimentary, vʹʹ 7–10; genu I lʹ 1, lʹʹ 1–2, vʹ 2–3, vʹʹ 2–3; tibia and tarsus I fused, tibial setae d 10–11, lʹ 2–3, lʹʹ 2, vʹ 2–3, vʹʹ 5–6, solenidion φ 2–4, tarsal setae tcʹ 7–8, tcʹʹ 8, pvʹ 5, pvʹʹ 3–4, s spine-like, 5, solenidion ω 3–4. Femur II vʹʹ 3–4; tibia II d 7–8, lʹ 1–2, vʹ 3–5, vʹʹ 9; tarsus II tcʹ 4–7, tcʹʹ 9–10, pʹ 1, uʹ spine-like, 5, uʹʹ 1, solenidion ω 2. Tibia III d 7–8, lʹ 2–3, vʹ 6–8, vʹʹ 9–10; tarsus III tcʹ 7–8, tcʹʹ 12–15, pʹ 1, uʹ spine-like, 5, uʹʹ 1. Leg I without claws. Etymology. The species name zuluensis refers to the presence of this species in Zululand, which is now a part of KwaZulu-Natal, South Africa. Remarks. Podotarsonemus zuluensis is most similar to P. indicus but differs by several character states as noted in the Differential diagnosis for P. indicus, and also detailed in the key below.

Podotarsonemus nipponicus Seeman, Lindquist & Husband sp. nov. Figures 20���22 Type material. Holotype. Adult female. JAPAN, Maesato, Ishigaki-jimi Is., Okinawa Pref., T. Takahashi, 22 Aug. 1975, ex hind wings of Euparatettix tricarinatus (Bol��var) (Tetrigidae), Kurosa No. 4880-1. In NMST. Paratypes. 16 females, 7 males, 32 larvae, as follows. 3 females, 3 larvae, same data as holotype (slides 2���7). 2 females, 2 larvae, same data as holotype except 17 Jun. 1975, Kurosa No. 4877 (slides 1���4). 3 females, 1 male, 4 larvae, same data as holotype except 17 Jun. 1975, Kurosa No. 4879 (slides 1, 3, 4, 5, 6). 1 female, 3 males, 4 larvae, same data as holotype except 17 Jun. 1975, Kurosa No. 4881 (slides 1, 2, 3, 4���8). 3 females, 1 male, 3 larvae, same data as holotype except 17 Jun. 1975, Kurosa No. 4882 (slides 1���6, 8). 2 larvae, same data as holotype except Kurosa No. 4883 (slides 1���2). 4 females, 2 males, 14 larvae, same data as holotype except Kurosa No. 4884 (slides 1���18, 20���24). In NMST except 1 female, 1 male, 1 larva in each of CNCI, QM, UMMZ. Diagnosis. Female and male: Palps with 2 setae; chelicerae with 1 tooth (female) or without teeth (male); prodorsal plate with punctate sculpturing anteriorly, imbricate sculpturing behind sc2; femora I���II without seta l��; genua II���III bare; tarsi II���III with 5 setae, seta u���� present; trochanter IV without seta. Female: Setae sc1 present; vestigial setae v2 expressed as alveoli within prodorsal shield; coxisternal plates III���IV separated by soft cuticle; setae h attenuate,> 100 long; femorogenu IV with 2 setae; tracheal atria separate, post-atrial sac lightly sclerotized, not punctate. Male: Genital capsule without copulatory flanges; setae h1 rudimentary, like h2; femorogenu IV with 1 seta; tibia IV with solenidion and rudimentary seta; tarsus IV absent. Male and larva: Leg I with tibia and tarsus fused. Larva: Setae h1-2, ps1-2 attenuate; leg I without claws. Description. ADULT FEMALE (Fig. 20, n = 15 measured) Gnathosoma. Length 41���42, width 43���45. Palps 10, with 2 setae (3���5 and 1���4 long). Cheliceral stylets 1 7���20, each with 1 tooth. Pharynx length 23���29, width 22���24. Dorsal setae ch 20���24, ventral setae su 14���23, palpcoxal setae e 8���13. Idiosoma. Length 166���180 (physogastric 275���500), width 135���139. Tracheal atria separate, post-atrial sac lightly sclerotized, not punctate, with two small distal prongs indicating separate terminal tracheae. Prodorsal plate with punctate sculpturing anteriorly, imbricate posterior to sc1, 72���78 long, 110���111 wide; with 3 pairs of setae: v 1 16���20, thickened, blunt, with small barbs, sc 1 6���12, sc2 58���61, and vestigial v2. Distance between setae v1-v1 34 ��� 38, v2-v2 42 ���50, sc1-sc1 90���95, sc2-sc2 54��� 62, v1 -sc 2 22���26. Plates C, D with punctate sculpturing anteriorly, imbricate posteriorly, EF and H with punctate sculpturing. Setae c1, d, e, f thickened, with small barbs, c 1 15���17, c2 40���47, d 15���19, e 15���17, f 17���20. Distance between setae c1-c1 70���77, c2-c2 127���137, c1-c 2 23���28, d-d 42��� 46, e-e 41���46, f-f 20���26. Setae h attenuate,> 100. Distance between setae h-h 22���25. Cupules ia, im present, ih indiscernible. Setae ps attenuate 220, distance between setae ps-ps 9���11. Coxisternal setae short to attenuate, 1a 9��� 11, 2a 19���25, 3a > 200, 3b 30. Distance between setae 1a-1a 15���21, 2a-2a 21���25, 3a-3a 5���13, 3b-3b 24���35; alveolar vestiges of setae 1b and 2b present. Legs. Leg lengths: I 49 ���51, II 48 ���50, III 50. Leg IV Fe-Ge length 19���21, Ti-Ta length 17���19. Leg III, femur and genu fused. Femur I d 1���2, l���� 3���12, v���� 22���27; genu I l�� 9���15, l���� 7���11, v�� 10���13, v���� 9���12; tibia and tarsus I fused, tibial setae d 23, l�� 11���12, l���� 4���6, v�� 9���11, v���� 12���13, solenidion �� 3���4, tarsal setae tc�� 13���14, tc���� 11���12, pv�� 14���17, pv���� 10���12, s spine-like, 7���8, solenidion �� 7���8. Femur II v���� 12���19; tibia II d 21���23, l�� 11���17, v�� 16���22, v���� 18���22; tarsus II tc�� 15���20, tc���� 20���23, pv�� 16���20, u�� spine-like, 7���8, u���� 2���4, solenidion �� 6. Tibia III d 18���21, l�� 17���23, v�� 16���20, v���� 16���21; tarsus III pv�� 22���25, tc�� 21���26, tc���� 22���24, u�� spine-like, 5���6, u���� 6���8. Femorogenu IV with proximal femoral seta 13���18, distal genual seta 26���28; tibiotarsus with attenuate ventral tibial seta> 80 and terminal tarsal seta> 180. ADULT MALE (Fig. 21, n = 5 measured) Gnathosoma. Length 30���31, width 32���36. Palps 6���7. Cheliceral stylets 12���13, without teeth. Pharynx length 20���21, width 11���12. Dorsal setae 9���16, ventral setae 11���13, palpcoxal setae 6���8. Idiosoma. Length 127���138, width 99���108. Prodorsal plate punctate, 59���62 long, 83���94 wide; with 3 pairs of setae (v 1 10���11, sc 1 7���10, sc 2 28���40) and vestigial v2. Distance between setae v1-v 1 21���25, v2-v 2 27���30, sc1-sc1 56���59, sc2-sc 2 30���33, v1-sc2 33���39. Plate CD punctate. Setae c1 tapering, with small barbs, 11���13, c2 fine, 16��� 21, d with small barbs, 10���12; setae c1 obliquely aligned between setae c2 and d. Plate EF punctate, e 5���7, f 6���8, with small barbs. Distance between setae c1-c1 51���59, c2-c2 85���92, c1-c 2 23���26, d-d 15���18, e-e 19���21, f-f 6���8. Cupules ia, im present. Genital capsule 27���30 long, 24���27 wide, bearing alveolar remnants of h1, h2; ventral flanges absent. Coxisternal setae attenuate 1a 9���11, 2a 13���17, 3a 80���100, 3b 16���20. Distance between setae 1a-1a 10���13, 2a-2a 23���25, 3a-3a 14���16, 3b-3b 14���17; setae 1a at level of or just posterior to ap1, setae 2b at level of ap2. Apodemes: ap2 almost reaching medially obsolete ap sj; ap5 small, extend 11���13 from base, not fused medially. Legs. Leg lengths: I 38 ���39, II 40, III 38, IV 2 6���29. Leg III, femur and genu fused. Femur I, d 1���2, l���� 3���4, v���� 10���15; genu I l�� 3���4, l���� 3���4, v�� 3���4, v���� 3���4; tibia and tarsus I fused, tibial setae d 14��� 19, l�� 3���4, l���� 2, v�� 3���5, v���� 6���8, solenidion �� 2, tarsal setae ft�� 7���9, ft���� 7���9, tc�� 11���13, tc���� 9���10, pv�� 10���11, pv���� 7���8, s spine-like, 6, solenidion �� 5���6. Femur II v���� 6���7; tibia II d 14���17, l�� 7���8, v�� 10���11, v���� 11���14; tarsus II tc�� 10���12, tc���� 11, pv�� 7���9, u�� spine-like, 6, u���� 2, solenidion �� 8. Tibia III d 15���20, l�� 7���10, v�� 11���15, v���� 13���15; tarsus III tc�� 11���16, tc���� 11���15, pv�� 7���10, u�� spine-like, 6, u���� 5���6. Trochanter IV bare; femorogenu IV with genual seta 3���6; tibia IV solenidion �� 7, v�� 1; tarsus IV absent. LARVA (Fig. 22, n = 30 measured) Gnathosoma. Length 37���41, width 33���39. Palps 6���7. Cheliceral stylets 11���14, without teeth. Pharynx length 15���17, width 11���12. Dorsal setae 11���14, ventral setae 8���10. Idiosoma. Length 167���250, width 114���155. Prodorsal plate smooth, 47���52 long, 89���99 wide; with 3 pairs of weakly barbed setae (v 1 8���10, sc1 6, sc 2 14���17) and vestigial v2. Distance between setae v1-v 1 25���29, v2-v2 37 ��� 44, sc1-sc1 59���65, sc2-sc2 41��� 44, v1 -sc 2 16���21. Plates C, D, EF smooth. Setae weakly barbed, c 1 11���13, c 2 9���11, d 11���13, e 8���10, f 11���12. Distance between setae c1-c1 53���59, c2-c2 90���105, c1-c 2 20���24, d-d 28���33, e-e 28���30, f-f 11���15. Plate H smooth, setae h1 attenuate,> 70, h2 ventral, > 100. Distance between setae h 16���21. Cupules ia, im, ih present. Setae ps1-2 attenuate, ps1 ca. 40, ps2 > 40. Coxisternal setae 1a 4���8, 2a 8���10, 3a attenuate,> 110, 3b 10���14. Distance between setae 1a-1a 15���16, 2a-2a 30���31, 3a-3a 34���43, 3b-3b 37���48. Legs. Femur I, l����, d rudimentary, v���� 6���10; genu I l�� 1���2, l���� 1���2, v�� 1���2, v���� 1���2; tibia and tarsus I fused, tibial setae d 10���16, l�� 2, l���� 1���2, v�� 2���3, v���� 6���7, solenidion �� 2, tarsal setae tc�� 9���10, tc���� 8, pv�� 7���8, p���� 2���3, s spine-like, 5��� 6, solenidion �� 3���5. Femur II v���� 3; tibia II d 7���10, l�� 3���5, v�� 7, v���� 8���9; tarsus II tc�� 10���13, tc���� 9���11, p�� 5���6, u�� spine-like, 5���6, u���� 7��� 8, solenidion �� 3���5. Tibia III d 7, l�� 3, v�� 7���8, v���� 7���9; tarsus III tc�� 11���12, tc���� 10���11, p�� 4���5, u�� spine-like, 5���6, u���� 7���11. Leg I without claws. Etymology. The species name nipponicus refers to the country where the type specimens were collected. Differential diagnosis. Podotarsonemus nipponicus belongs to the boliviensis species group (P. boliviensis, P. ecuadorensis, P. nipponicus, and P. queenslandensis) whose females lack a seta on trochanter IV, lack longitudinal striae between coxisterna III���IV, and have setae h attenuated. Of these species, P. nipponicus is most similar to P. queenslandensis in having no more than three setae on femur I and just one seta on femur II. Podotarsonemus nipponicus is distinguished from P. queenslandensis, which is known only from the female, by having setae sc1 (absent in P. queenslandensis), retaining a rudimentary seta d on femur I (absent in P. queenslandensis) and having the prodorsal, C and D plates with some imbricate sculpturing (punctate in P. queenslandensis). Remarks. One specimen (4881-4) is a pharate male, demonstrating that both sexes have active, sexually monomorphic larvae. This is the plesiomorphic condition in the Tarsonemidae and other Acariformes, but within the Podapolipidae, only the genera Chrysomelobia and Tarsopolipus retain an active, monomorphic male (Husband 1997; Seeman & Nahrung 2005). Most other genera of Podapolipidae have free-living larval and adult females, but adult males that emerge directly from their eggs as sexually mature paedomorphic forms (i.e. larviform males). The only exception is Scarabapolipus which have active larvae in both sexes, but they are dimorphic (Husband & Kurosa 1994)., Published as part of Seeman, Owen D., Lindquist, Evert E. & Husband, Robert W., 2018, A new tribe of tarsonemid mites (Trombidiformes: Heterostigmatina) parasitic on tetrigid grasshoppers (Orthoptera), pp. 1-54 in Zootaxa 4418 (1) on pages 35-39, DOI: 10.11646/zootaxa.4418.1.1, http://zenodo.org/record/1242435, {"references":["Husband, R. W. (1997) Two new species of Tarsopolipus (Acari: Podapolipidae) from scarab beetles from Central and Western Africa. International Journal of Acarology, 23, 275 - 282. https: // doi. org / 10.1080 / 01647959708683577","Seeman, O. D. & Nahrung, H. F. (2005) New Parobia (Acari: Podapolipidae) parasitic on pestiferous paropsine beetles (Coleoptera: Chrysomelidae) in eucalypt plantations. Systematic & Applied Acarology, 10, 110 - 135.","Husband, R. W. & Kurosa, K. (1994) Four new species of Scarabapolipus (Acari: Podapolipidae) ectoparasites of Scarabaeidae (Coleoptera) from the Western Pacific Region. Annals of the Entomological Society of America, 87, 424 - 435. https: // doi. org / 10.1093 / aesa / 87.4.424"]}

Podotarsonemus ecuadorensis Seeman, Lindquist & Husband sp. nov. Figures 13–14, 15C Type material. Holotype. Adult female. ECUADOR, Puyo Napo-Pastaza Prov., 2500’ alt., R.W. Hodges, 18 Apr. 1958, ex Batrachidea mucronata, RWH 18102010 (slide 7). In A. J. Cook Entomology Museum of Michigan State University. Paratypes. 1 female, 4 males, same data as holotype. In UMMZ except 1 male in each of CNCI and QM. Non-type material. 2 pharate females, 2 larvae, same data as holotype. Larvae in very poor condition, not described. In UMMZ. Diagnosis. Female and male: Palps with 1 seta; chelicerae without teeth; prodorsal plate punctate; femora I–II with rudimentary setae lʹ, d; genua II–III bare; tarsus II with 5 setae, with seta uʹʹ, tarsus III with 4 setae, lacking seta u″; trochanter IV bare. Female: Setae sc1 present; vestigial setae v2 expressed as alveoli within prodorsal shield; coxisternal plates III–IV separated by soft cuticle; setae h attenuate,> 70 long; femorogenu IV with 2 setae; tracheal atria separate, post-atrial sac lightly sclerotized, not punctate. Male: Genital capsule without copulatory flanges; setae h1 rudimentary, like h2; leg I with tibia and tarsus separate; femorogenu IV bare; tibia IV with solenidion and small vʹ; tarsus IV absent. Description. ADULT FEMALE (Figures 13, 15C, n = 2 measured; specimens distorted and fragmented to some extent, so only diagnostic features are illustrated) Gnathosoma. Length 56–57, width 60–61. Palps 13–14, with 1 apical seta 4. Cheliceral stylets 22–25, without teeth. Pharynx length 31–35, width 14–16. Dorsal setae ch 30–32, ventral setae su 19–23, palpcoxal setae e 6–7. Idiosoma. Length and width not measureable. Tracheal atria separate, post-atrial sac lightly sclerotized, not punctate, terminal tracheae separate. Prodorsal plate with punctate sculpturing, 96–100 long, 135–137 wide; with 3 pairs of setae: v 1 29–30, thickened, with small barbs, sc 1 19–25, sc2 90 and vestigial v2. Distance between setae v1-v1 54 –56, v2-v2 66 –68, sc1-sc1 118, sc2-sc2 78– 85, v1 -sc 2 23–31. Plates C, D, EF with punctate sculpturing. Setae c1, d, e, f thickened, blunt, with small barbs, c1 27, c2 59–65, d 25, e 23–24, f 23–24. Distance between setae c1-c1 not measureable, c2-c2 195, c1-c2 33–37, d-d 79–93, e-e 67–68, f-f 36–41. Setae h attenuate,> 70. Distance between setae h-h 30. Cupules ia, im, ih not visible or absent. Setae ps attenuate> 100, distance between setae psps 10. Coxisternal setae short to attenuate, 1a 5–21, 2a 10–32, 3a > 100, 3b > 30; alveolar vestiges of setae 1b and 2b present. Distances between setae impossible to measure accurately. Legs. Leg lengths: I 61 –63, II 67 –71, III 67 –70. Leg IV Fe-Ge length 32–33, Ti-Ta length 26–28. Leg III, femur and genu fused. Femur I, setae lʹ, d rudimentary, lʹʹ 16–17, vʹʹ 24–27; genu I lʹ 11–20, lʹʹ 20–21, vʹ 13–14, vʹʹ 12–17; tibia and tarsus I fused, tibial setae d 45–47, lʹ 18–20, l″ 15, vʹ 15–19, vʹʹ 16, solenidion φ 6, tarsal setae tcʹ 17–20, tcʹʹ 17–20, pvʹ 19–20, pvʹʹ 14–17, s spine-like, 8, solenidion ω 8–10. Femur II lʹ, d rudimentary, vʹʹ 16–19; tibia II d 30–35, lʹ 24–34, vʹ 30–31, vʹʹ 26–27; tarsus II tcʹ 22–25, tcʹʹ 28– 29, pvʹ 20–23, uʹ spine-like, 7–8, uʹʹ 6–8, solenidion ω 7. Tibia III d 37, lʹ 21–26, vʹ 28–34, vʹʹ 28; tarsus III tcʹ 27–33, tcʹʹ 35–41, pvʹ 30, uʹ spine-like, 6–8. Femorogenu IV with short proximal femoral seta vʹ 10–16, and fine distal genual seta vʹ 26–30; tibiotarsus with attenuate ventral tibial seta 90 and terminal tarsal seta> 200. ADULT MALE (Fig. 14, n = 4 measured) Gnathosoma. Similar to female. Length 35–40, width 36–40. Palps 8–9. Cheliceral stylets 15–18, without teeth. Pharynx length 25–30, width 12–13. Dorsal setae 13–15, ventral setae 10–14, palpcoxal setae 7. Idiosoma. Length 160–180, width 147–150. Prodorsal plate smooth, 61–82 long, 97–112 wide; with 3 pairs of setae, v 1 10–11, sc1 6–9, sc2 34–37 and vestigial v2. Distance between setae v1-v 1 25–28, v2-v2 36 –41, sc1-sc1 56–69, sc2-sc2 49– 60, v1 -sc2 40. Plate CD smooth. Setae sparsely barbed, setae c1 11, c 2 25–35, d 11–12; setae c1 obliquely aligned between setae c2 and d. Plate EF smooth, e 4–5, f 4–5. Distance between setae c1-c1 65–84, c2- c2 111–130, c1-c 2 23–27, d-d 24, e–e 18, f-f 9. Cupules ia, im present. Genital capsule 33–34 long, 30–31 wide, bearing alveolar remnants of h1, h2; ventral flanges absent. Coxisternal setae 1a short, 2a, 3a, 3b short to attenuate, 1a 4–6, 2a 20–40, 3a 100, 3b > 70. Distance between setae 1a-1a 10–13, 2a-2a 21–32, 3a-3a 20, 3b-3b 23; setae 1a at level of ap1, setae 2b at level of or just posterior to ap2. Apodemes: ap2 not reaching medially obsolete ap sj; ap5 extend anteriorly 34–39 from base, not fused medially. Legs. Leg lengths: I 47, II 46, III 45, IV 30–31. Leg III, femur and genu fused. Femur I, lʹ, lʹʹ, d rudimentary, vʹʹ 10–13; genu I lʹ 2–4, lʹʹ 2–3, vʹ 2, vʹʹ 5–7; tibia and tarsus I fused, tibial setae d 20–24, lʹ 7–10, lʹʹ 2–3, vʹ 6–7, vʹʹ 10–11, solenidion φ 4–5, tarsal setae ftʹ 13–14, ftʹʹ 12–13, tcʹ 13, tcʹʹ 12, pvʹ 9–10, pvʹʹ 6–7, s spine-like, 5, solenidion ω 7. Femur II lʹ, d rudimentary, vʹʹ 6–7; tibia II d 19, lʹ 10, vʹ 14, vʹʹ 23; tarsus II tcʹ 17, tcʹʹ 20–22, pvʹ 12, uʹʹ 7–8, uʹ spine-like, 9, solenidion ω 9. Tibia III d 19–26, lʹ 11–13, vʹ 19–25, vʹʹ 23–27; tarsus III tcʹ 20–23, tcʹʹ 16–20, pvʹ 10–15, uʹ spine-like, 6–7. Femorogenu IV bare; tibia IV vʹ 2–3, solenidion φ 10–12; tarsus IV absent. Etymology. The species name ecuadorensis refers to the country where the type specimens were collected. Remarks. Podotarsonemus ecuadorensis is most similar to P. boliviensis and is differentiated in the Differential diagnosis for that species. However, the separation of these species is complicated by several female specimens from Ecuador and Honduras, as discussed below under the specimens referred to as Podotarsonemus sp.

Podotarsonemus indicus Seeman, Lindquist & Husband sp. nov. Figures 16–19 Type material. Holotype. Adult female. INDIA, Madras, Dist. Coimbatore, Coimbatore, P.S. Nathan, 5 Jul. 1946, ex Scelimena kempi (Hancock), RWH VII2010 (slide 1). In UMMZ. Paratypes. 15 females, 11 males, 2 larvae, as follows. 11 females, 11 males, 1 larva same data as holotype. 3 females, 1 larva, same data as holotype except 26 Nov. 1945. 1 female, same data as holotype except 3 Nov. 1947. In UMMZ, except 1 female, 1 male, 1 larva in CNCI and 1 female, 1 male in QM. Diagnosis. Female and male: Palps with 2 setae; chelicerae without teeth; prodorsal plate with punctate sculpturing; femora I–II without setae lʹ, d; genua II–III bare; tarsi II–III with 5 setae, uʹʹ present; trochanter IV with 1 seta. Female: Setae sc1 present; vestigial setae v2 expressed as alveoli within prodorsal shield; coxisternal plates III–IV separated by longitudinal striae; setae h not attenuate, h1 rudimentary, like h2; leg I with tibia and tarsus fused; femorogenu IV bare; tibia IV with solenidion and attenuate terminal seta; tarsus IV rudimentary, subterminal. Larva: Setae h1-2, ps2 attenuate, setae ps1 short; leg I with separate tibia and tarsus, with 2 rudimentary claws. Description. ADULT FEMALE (Figures 16–17, n = 9 measured) Gnathosoma. Length 57–67, width 68–71. Palps 9–13, with 2 setae (8 and 3 long). Cheliceral stylets 28–31, without teeth. Pharynx length 41–50, width 20–23. Dorsal setae ch 35–42, ventral setae su 27–32, palpcoxal setae e 7–8. Idiosoma. Length 242–300 (physogastric 600), width 226–240. Tracheal atria fused, post-atrial sac heavily sclerotized, punctate, with single long terminal extension of trachea. Prodorsal plate with punctate sculpturing, 103–110 long, 148–174 wide; with 3 pairs of setae: v1 21 –26, thickened, with small barbs, sc 1 17–21, sc2 67–78, and vestigial v2. Distance between setae v1-v1 63 –75, v2-v2 68 –82, sc1-sc1 124–140, sc2-sc2 83–106, v1 -sc2 33– 38. Plates C, D, EF with punctate sculpturing. Setae c1, d, e, f thickened, with small barbs, c 1 18–20, c2 51–65, d 17–20, e 15–21, f 15–20. Distance between setae c1-c1 122–128, c2-c2 201–209, c1-c2 35–38, d-d 64–76, e-e 73– 80, f-f 28–34. Plate H variably covered by EF, setae h not attenuate, 11–26. Distance between setae h-h 31–37. Cupules ia, im, ih present. Setae ps attenuate> 300, distance between setae ps-ps 9–12. Coxisternal setae attenuate, 1a 80 (often broken), 2a > 120 (often broken), 3a > 100, 3b > 100. Distance between setae 1a-1a 26–31, 2a-2a 59– 63, 3a-3a 15–18, 3b-3b 35–43; alveolar vestiges of setae 1b and 2b present. Coxisternal plates III–IV separated by longitudinal striae; coxisternal plates III convex anteriorly, sometimes just reaching posterior margin of coxisterna II. Legs. Leg lengths: I 71 –73, II 75 –79, III 70 –73. Leg IV Fe-Ge length 33–34, Ti-Ta length 20–22. Leg III, femur and genu fused, line of fusion apparent and one specimen with asymmetric separation of femur and genu. Femur I lʹʹ 5–6, vʹʹ 15–28; genu I lʹ 9–11, lʹʹ 9–10, vʹ 3–5, vʹʹ 4–5; tibia and tarsus I fused, tibial setae d 31–34, lʹ 9–12, lʹʹ 9–11, vʹ 6–7, vʹʹ 11–13, solenidion φ 3–4, tarsal setae tcʹ 12–14, tcʹʹ 12–13, pvʹ 10–13, pvʹʹ 8–10, s spinelike, 6–7, solenidion ω 7–8. Femur II vʹʹ 15–24; tibia II d 24–30, lʹ 6–11, vʹ 13–21, vʹʹ 15–25; tarsus II tcʹ 16–17, tcʹʹ 18, pvʹ 13–15, uʹ spinelike, 7, uʹʹ 6–8, solenidion ω 6–7. Tibia III d 24–32, lʹ 14–20, vʹ 18–22, vʹʹ 17–23; tarsus III tcʹ 17–22, tcʹʹ 26–32, pvʹ 15–17, uʹ spine-like, 8, uʹʹ 5– 6. Trochanter IV with seta 11–13; tibiotarsus with ventral tibial seta 45–60, terminal tarsal seta> 400. ADULT MALE (Fig. 18, n = 5 measured) Gnathosoma. Similar to female. Length 61, width 66. Palps 11–12. Cheliceral stylets 2 5–26, without teeth. Pharynx length 34–35, width 18. Dorsal setae 39–41, ventral setae 7, palpcoxal setae 3. Idiosoma. Length 170–238, width 168–179. Prodorsal plate punctate, 74 long, 104–105 wide; with 3 pairs of setae (v1 14 –18, sc 1 17–18, sc2 75–78) and vestigial v2. Distance between setae v1-v1 34 –38, v2-v2 44 –50, sc1-sc1 87–90, sc2-sc2 64– 65, v1 -sc 2 29–32. Plate CD punctate. Setae c1 tapering, with few barbs, 7, c2 attenuate, 70–84, d with few barbs, 19–20; setae c 1 in nearly transverse alignment with c2. Plate EF punctate, e 18–19, f 16, with few barbs. Distance between setae c1-c1 64, c2-c2 134, c1-c 2 21–25, d-d 33, e-e 47–51, f-f 16. Cupules ia, im present. Genital capsule 43 long, 51 wide, bearing alveolar remnants of h1, h2; ventral flanges absent. Coxisternal setae attenuate 1a 30–36, 2a 28–30, 3a > 190, 3b > 30. Distance between setae 1a-1a 33–35, 2a-2a 56–59, 3a-3a 19–23, 3b-3b 29–31; setae 1a at level of or just posterior to ap1, setae 2b usually just posterior to ap2 (one specimen with asymmetrical position of setae 2b far posterior to ap2). Apodemes: ap2 not reaching medially obsolete ap sj; ap5 obsolete. Legs. Leg lengths: I 70 –72, II 83 –89, III 83 –85, IV 52 –54. Leg III, femur and genu separate. Femur I, lʹʹ 5–8, vʹʹ 30; genu I lʹ 8–12, lʹʹ 7–8, vʹ 4, vʹʹ 3–4; tibia and tarsus I fused, tibial setae d 44–50, lʹ 7–10, lʹʹ 10, vʹ 7, vʹʹ 15, solenidion φ 5, tarsal setae ftʹ 16, ftʹʹ 15, tcʹ 15–17, tcʹʹ 15–17, pvʹ 12, pvʹʹ 14, s spine-like, 7, solenidion ω 7–8. Femur II vʹʹ 25; tibia II d 38–42, lʹ 27–29, vʹ 10–18, vʹʹ 31–32; tarsus II pvʹ 6–9, tcʹ 18–20, tcʹʹ 18–20, uʹʹ 5–6, uʹ spine-like, 6, solenidion ω 8. Tibia III d 32–35, lʹ 27–30, vʹ 26–32, vʹʹ 23–25; tarsus III pvʹ 17–20, tcʹ 27–28, tcʹʹ 25–29, uʹ spine-like, 7, uʹʹ 5– 6. Trochanter IV with seta, 8; femorogenu IV bare; tibia IV φ 11, terminal seta attenuate, 150; tarsus IV rudimentary, subterminal. LARVA (Fig. 19, n = 1 measured) Gnathosoma. Length 46, width not measurable. Palp length 5, with 2 setae (lengths 6 and minute). Cheliceral stylets 26, without teeth. Pharynx length 29, width 18. Dorsal setae 18, ventral setae 9. Idiosoma. Length 225, width 160. Prodorsal plate smooth, 60 long, 95 wide; with 3 pairs of tapering, weakly barbed setae (v1 10, sc1 8, sc2 17) and vestigial v2. Distance between setae v1-v1 28, v2-v2 49, sc1-sc1 not measurable as sc1 missing on left hand side, sc2-sc2 58, v1 -sc2 41. Plates C, D, EF smooth. Setae tapering, weakly barbed, c1 17, c2 14, d 14, e 12, f 15. Distance between setae c1-c1 76, c2-c2 130, c1-c2 30, d-d 28, e-e 41, f-f 13. Plate H smooth, setae h1 long, 65, h2 ventral,> 120. Distance between setae h1 24. Cupules ia, im, ih present. Setae ps1 fine, 8, ps2 attenuate 60. Coxisternal setae attenuate, 1a 36, 2a 41, 3a > 40, 3b 30. Distance between setae 1a-1a 36, 2a-2a 41, 3a-3a 37, 3b-3b 35. Longitudinal striae between coxisterna III–IV. Legs. Femur I, lʹʹ rudimentary, vʹʹ 10; genu I lʹ, lʹʹ rudimentary, vʹ 3, vʹʹ 2; tibia and tarsus I separate, tibial setae d 15, lʹ 4, lʹʹ 4, vʹ 3, vʹʹ 6, solenidion φ 3, tarsal setae tcʹ 9, tcʹʹ 9, pvʹ 3, pvʹʹ 6, s spine-like, 5, solenidion ω 4. Femur II vʹʹ 7; tibia II d 9, lʹ 5, vʹ 11, vʹʹ 11; tarsus II tcʹ 7, tcʹʹ 12, pʹ 4, uʹ spine-like, 5, uʹʹ 2, solenidion ω 3. Tibia III d 9, lʹ 6, vʹ 10, vʹʹ 10; tarsus III tcʹ 6, tcʹʹ 13, pʹ 4, uʹ spine-like, 5, uʹʹ 3. Leg I with 2 rudimentary claws. Etymology. The species name indicus refers to the country where the type specimens were collected. Differential diagnosis. Podotarsonemus indicus belongs to the australiensis species group (P. australiensis, P. indicus, P. zuluensis) whose females retain a seta on trochanter IV, have longitudinal striae between coxisterna III– IV and have setae h not attenuated. Of these species, P. indicus most closely resembles P. zuluensis but is distinguished by having: females with tracheal atria fused (free in other Podotarsonemus), females with alveolar remnants of setae v2 within the prodorsal shield (anterolateral invaginations of prodorsal shield in P. zuluensis), females and males retaining seta u″ on tarsus II (absent in P. zuluensis), males with setae c2 much longer (> ×4) than c1 (P. zuluensis) and tibia IV having an attenuate terminal seta and subterminal solenidion (lacking terminal seta and solenidion terminal in P. zuluensis), and larvae with setae h1-2 and ps2 attenuate (short in P. zuluensis).

Podotarsonemus ecuadorensis Seeman, Lindquist & Husband sp. nov. Figures 13���14, 15C Type material. Holotype. Adult female. ECUADOR, Puyo Napo-Pastaza Prov., 2500��� alt., R.W. Hodges, 18 Apr. 1958, ex Batrachidea mucronata, RWH 18102010 (slide 7). In A. J. Cook Entomology Museum of Michigan State University. Paratypes. 1 female, 4 males, same data as holotype. In UMMZ except 1 male in each of CNCI and QM. Non-type material. 2 pharate females, 2 larvae, same data as holotype. Larvae in very poor condition, not described. In UMMZ. Diagnosis. Female and male: Palps with 1 seta; chelicerae without teeth; prodorsal plate punctate; femora I���II with rudimentary setae l��, d; genua II���III bare; tarsus II with 5 setae, with seta u����, tarsus III with 4 setae, lacking seta u���; trochanter IV bare. Female: Setae sc1 present; vestigial setae v2 expressed as alveoli within prodorsal shield; coxisternal plates III���IV separated by soft cuticle; setae h attenuate,> 70 long; femorogenu IV with 2 setae; tracheal atria separate, post-atrial sac lightly sclerotized, not punctate. Male: Genital capsule without copulatory flanges; setae h1 rudimentary, like h2; leg I with tibia and tarsus separate; femorogenu IV bare; tibia IV with solenidion and small v��; tarsus IV absent. Description. ADULT FEMALE (Figures 13, 15C, n = 2 measured; specimens distorted and fragmented to some extent, so only diagnostic features are illustrated) Gnathosoma. Length 56���57, width 60���61. Palps 13���14, with 1 apical seta 4. Cheliceral stylets 22���25, without teeth. Pharynx length 31���35, width 14���16. Dorsal setae ch 30���32, ventral setae su 19���23, palpcoxal setae e 6���7. Idiosoma. Length and width not measureable. Tracheal atria separate, post-atrial sac lightly sclerotized, not punctate, terminal tracheae separate. Prodorsal plate with punctate sculpturing, 96���100 long, 135���137 wide; with 3 pairs of setae: v 1 29���30, thickened, with small barbs, sc 1 19���25, sc2 90 and vestigial v2. Distance between setae v1-v1 54 ���56, v2-v2 66 ���68, sc1-sc1 118, sc2-sc2 78��� 85, v1 -sc 2 23���31. Plates C, D, EF with punctate sculpturing. Setae c1, d, e, f thickened, blunt, with small barbs, c1 27, c2 59���65, d 25, e 23���24, f 23���24. Distance between setae c1-c1 not measureable, c2-c2 195, c1-c2 33���37, d-d 79���93, e-e 67���68, f-f 36���41. Setae h attenuate,> 70. Distance between setae h-h 30. Cupules ia, im, ih not visible or absent. Setae ps attenuate> 100, distance between setae psps 10. Coxisternal setae short to attenuate, 1a 5���21, 2a 10���32, 3a > 100, 3b > 30; alveolar vestiges of setae 1b and 2b present. Distances between setae impossible to measure accurately. Legs. Leg lengths: I 61 ���63, II 67 ���71, III 67 ���70. Leg IV Fe-Ge length 32���33, Ti-Ta length 26���28. Leg III, femur and genu fused. Femur I, setae l��, d rudimentary, l���� 16���17, v���� 24���27; genu I l�� 11���20, l���� 20���21, v�� 13���14, v���� 12���17; tibia and tarsus I fused, tibial setae d 45���47, l�� 18���20, l��� 15, v�� 15���19, v���� 16, solenidion �� 6, tarsal setae tc�� 17���20, tc���� 17���20, pv�� 19���20, pv���� 14���17, s spine-like, 8, solenidion �� 8���10. Femur II l��, d rudimentary, v���� 16���19; tibia II d 30���35, l�� 24���34, v�� 30���31, v���� 26���27; tarsus II tc�� 22���25, tc���� 28��� 29, pv�� 20���23, u�� spine-like, 7���8, u���� 6���8, solenidion �� 7. Tibia III d 37, l�� 21���26, v�� 28���34, v���� 28; tarsus III tc�� 27���33, tc���� 35���41, pv�� 30, u�� spine-like, 6���8. Femorogenu IV with short proximal femoral seta v�� 10���16, and fine distal genual seta v�� 26���30; tibiotarsus with attenuate ventral tibial seta 90 and terminal tarsal seta> 200. ADULT MALE (Fig. 14, n = 4 measured) Gnathosoma. Similar to female. Length 35���40, width 36���40. Palps 8���9. Cheliceral stylets 15���18, without teeth. Pharynx length 25���30, width 12���13. Dorsal setae 13���15, ventral setae 10���14, palpcoxal setae 7. Idiosoma. Length 160���180, width 147���150. Prodorsal plate smooth, 61���82 long, 97���112 wide; with 3 pairs of setae, v 1 10���11, sc1 6���9, sc2 34���37 and vestigial v2. Distance between setae v1-v 1 25���28, v2-v2 36 ���41, sc1-sc1 56���69, sc2-sc2 49��� 60, v1 -sc2 40. Plate CD smooth. Setae sparsely barbed, setae c1 11, c 2 25���35, d 11���12; setae c1 obliquely aligned between setae c2 and d. Plate EF smooth, e 4���5, f 4���5. Distance between setae c1-c1 65���84, c2- c2 111���130, c1-c 2 23���27, d-d 24, e���e 18, f-f 9. Cupules ia, im present. Genital capsule 33���34 long, 30���31 wide, bearing alveolar remnants of h1, h2; ventral flanges absent. Coxisternal setae 1a short, 2a, 3a, 3b short to attenuate, 1a 4���6, 2a 20���40, 3a 100, 3b > 70. Distance between setae 1a-1a 10���13, 2a-2a 21���32, 3a-3a 20, 3b-3b 23; setae 1a at level of ap1, setae 2b at level of or just posterior to ap2. Apodemes: ap2 not reaching medially obsolete ap sj; ap5 extend anteriorly 34���39 from base, not fused medially. Legs. Leg lengths: I 47, II 46, III 45, IV 30���31. Leg III, femur and genu fused. Femur I, l��, l����, d rudimentary, v���� 10���13; genu I l�� 2���4, l���� 2���3, v�� 2, v���� 5���7; tibia and tarsus I fused, tibial setae d 20���24, l�� 7���10, l���� 2���3, v�� 6���7, v���� 10���11, solenidion �� 4���5, tarsal setae ft�� 13���14, ft���� 12���13, tc�� 13, tc���� 12, pv�� 9���10, pv���� 6���7, s spine-like, 5, solenidion �� 7. Femur II l��, d rudimentary, v���� 6���7; tibia II d 19, l�� 10, v�� 14, v���� 23; tarsus II tc�� 17, tc���� 20���22, pv�� 12, u���� 7���8, u�� spine-like, 9, solenidion �� 9. Tibia III d 19���26, l�� 11���13, v�� 19���25, v���� 23���27; tarsus III tc�� 20���23, tc���� 16���20, pv�� 10���15, u�� spine-like, 6���7. Femorogenu IV bare; tibia IV v�� 2���3, solenidion �� 10���12; tarsus IV absent. Etymology. The species name ecuadorensis refers to the country where the type specimens were collected. Remarks. Podotarsonemus ecuadorensis is most similar to P. boliviensis and is differentiated in the Differential diagnosis for that species. However, the separation of these species is complicated by several female specimens from Ecuador and Honduras, as discussed below under the specimens referred to as Podotarsonemus sp., Published as part of Seeman, Owen D., Lindquist, Evert E. & Husband, Robert W., 2018, A new tribe of tarsonemid mites (Trombidiformes: Heterostigmatina) parasitic on tetrigid grasshoppers (Orthoptera), pp. 1-54 in Zootaxa 4418 (1) on pages 26-28, DOI: 10.11646/zootaxa.4418.1.1, http://zenodo.org/record/1242435

Podotarsonemus boliviensis Seeman, Lindquist & Husband sp. nov. Figures 7���12, 15A, B Type material. Holotype. Adult female. BOLIVIA, Dept. Beni, Puertofonbeni, Luis E. Pe��a, 12 Nov. 1956, ex Batrachidea mucronata (Serville) (Tetrigidae). G# 23, RWH 15I 2010 (slide 2). In UMMZ. Paratypes. 20 females, 5 males, 11 larvae, as follows: 14 females, 4 males, 8 larvae, same data as holotype. 1 female, 1 male, same data as holotype except additional collection data ���on way to Reyes from Rio Beni ���, 6���10 Nov. 1956. 5 females, 3 larvae, Dept. La Laz, Altamarani on Beni River, Luis E. Pe��a, 5���11 Nov. 1956, ex B. mucronata. In UMMZ except 1 female, 1 male, 1 larva in each of CNCI and QM. Non-type material. 1 adult female, same data as holotype, RWH 15I 2010 (slide 8)���aberrant specimen. 1 pharate female, 8 eggs, same data as holotype. In UMMZ. Diagnosis. Female and male: Palps with 1 seta; chelicera with 1 tooth (female) or without teeth (male); prodorsal plate with imbricate sculpturing; femora I���II with rudimentary seta l��, d; genua II���III bare; tarsi II���III with 4 setae, lacking seta u��� (except male tarsus III with 5 setae, u��� present); trochanter IV bare. Female: Setae sc1 present; vestigial setae v2 expressed as alveoli within prodorsal shield; coxisternal plates III���IV separated by soft cuticle; setae h attenuate,> 70 long; femorogenu IV with 2 setae; tracheal atria separate, post-atrial sac lightly sclerotized, not punctate. Male: Genital capsule with small copulatory flanges; setae h1 rudimentary, like h2; femorogenu IV with 1 seta; tibia IV with solenidion, without setae; tarsus IV absent. Male and larva: Leg I with tibia and tarsus fused. Larva: Setae h1-2, ps1-2 attenuate; leg I without claws. Description. ADULT FEMALE (Figures 7���8, 15A, B, n = 11 measured) Measurements are in the form of: Range [Measurement of aberrant specimen if outside of range]. Gnathosoma. Length 51���63, width 59���66 [69]. Palps 9���14, with 1 apical seta 7���9 flanked medially by pointed triangular hyaline process. Cheliceral stylets 1 9���23, each with 1 tooth. Pharynx length 26���42, width 19���25. Dorsal setae ch 25���34, ventral setae su 8���23 [30], palpcoxal setae e 7���13. Idiosoma. Length 228���245, width 159���198. Tracheal atria separate, post-atrial sac lightly sclerotized, not punctate, terminal tracheae separate. Prodorsal plate with imbricate sculpturing, irregular anteriorly [similar, but smooth between sc2-sc2], 91���106 long, 140���163 wide; with 3 pairs of setae: v 1 20���26 [29] with few barbs, sc 1 5��� 22 variable, mostly 5���6, but 14 and 22 in two specimens, sc2 47���75 [100] and vestigial v2. Distance between setae v1-v1 46 ���56 [45], v2-v2 56 ���64 [68], sc1-sc1 120���131 [110], sc2-sc2 67��� 82, v1 -sc 2 26���32. Plates C, D, EF with imbricate sculpturing [similar, but smooth between c1-c1]. Setae c1, d, e, f with few barbs [many barbs], c 1 15���23 [37], c2 44���54 [> 100], d 15���22 [33], e 15���22 [25], f 15���22. Distance between setae c1-c1 109���123 [95], c2-c2 154���170, c1-c 2 23���30, d-d 82���96 [68], e-e 71���80, f-f 36���50 [31]. Plate H folded beneath EF, setae h attenuate,> 130. Distance between setae h-h 26���37. Cupules ia, im, ih present. Setae ps attenuate> 130, distance between setae ps-ps 10���13 [14]. Coxisternal setae attenuate, 1a 90 (often broken), 2a 96���135 (often broken), 3a > 250, 3b > 210. Distance between setae 1a-1a 21���28 [31], 2a-2a 33���47 [57], 3a-3a 18���31 [11], 3b-3b 33���50 [55]; alveolar vestiges of setae 1b and 2b present. Legs. Leg lengths: I 55 ���60 [75], II 51 ���56 [74], III 53 ���55 [78]. Leg IV Fe-Ge length 25���29, Ti-Ta length 18���21. Leg III, femur and genu fused. Femur I, l��, l����, d rudimentary [l���� 2], v���� 12���21 [27]; genu I l�� 4���9 [14], l���� 6���10, v�� 4��� 5 [9], v���� 4���6 [9]; tibia and tarsus I fused, tibial setae d 28���41, l�� 4���9 [15], l��� 3 [5], v�� 5���9 [15], v���� 8���13 [14], solenidion �� 3���4 [6], tarsal setae tc�� 15 [17], tc���� 13���15 [17], pv�� 9���15, pv���� 6���10 [17], s spine-like, 5���9, solenidion �� 8���11. Femur II l��, d rudimentary, v���� 7���19 [24]; tibia II d 30���39, l�� 8���21 [22], v�� 7���24 [38], v���� 19���33 [35]; tarsus II pv�� 12���16 [22], tc�� 19���22 [25], tc���� 27���45, u�� spine-like, 6���10, solenidion �� 8���12. Tibia III d 23���38 [40], l�� 11���15 [30], v�� 17���33, v���� 25���31 [34]; tarsus III pv�� 11���24, tc�� 21���29 [31], tc���� 26���46, u�� spine-like, 8���9. Femorogenu IV with 2 attenuate ventral setae (Fe- v��, Ge- v��); proximal femoral seta 18���25 [36], distal genual seta 36���45 [28]; tibiotarsus with attenuate ventral tibial seta and terminal tarsal seta> 100. ADULT MALE (Figs 9���10, n = 2 measured) Gnathosoma. Similar to female. Length 33���36, width 35���36. Palps 10���12. Cheliceral stylets 15���17, without teeth. Pharynx length 20���22, width 10���11. Dorsal setae 10���12, ventral setae 7���10, palpcoxal setae 2���3. Idiosoma. Length 165���175, width 140. Prodorsal plate punctate, 65���73 long, 105���110 wide; with 3 pairs of setae (v 1 16���17, sc1 4���5, sc2 39���42) and vestigial v2. Distance between setae v1-v 1 29���31, v2-v 2 30���34, sc1-sc1 72���75, sc2-sc2 41��� 44, v1 -sc2 41���43. Plates CD, EF punctate. Setae smooth, setae c1 6���7, c 2 25���26, d 35���40 (left side d 8 on one specimen); setae c1 obliquely aligned between setae c2 and d. Plate EF, e 5, f 6. Distance between setae c1-c1 75, c2-c2 100���115, c1-c 2 28���32, d-d 26���27, e-e 36���37, f-f 29���30. Cupules ia, im present. Genital capsule 30���35 long, 32 wide, bearing alveolar remnants of h1, h2; ventral flanges present, small. Coxisternal setae attenuate, 1a 28, 2a 70, 3a > 90, 3b > 80. Distance between setae 1a-1a 13���15, 2a-2a 38���40, 3a-3a 10���13, 3b-3b 37���40; setae 1a at level of or just posterior to ap1, setae 2b at level of or just posterior to ap2. Apodemes: ap2 not reaching medially obsolete ap sj; ap5 small, extending 17���21 from base, not fused medially. Legs. Leg lengths: I 47, II 46, III 45, IV 30���31. Leg III, femur and genu fused. Femur I, l��, l����, d rudimentary, l���� on small tubercle, v���� 8���10; genu I l�� 2���3, l���� 4���5, v�� 3, v���� 2���3; tibia and tarsus I fused, tibial setae d 23���25, l�� 4, l���� 2���3, v�� 6���7, v���� 10���15, solenidion �� 3���4, tarsal setae ft�� 12, ft���� 12, tc�� 11���12, tc���� 9��� 11, pv�� 9���11, pv���� 9���11, s spine-like, 8, solenidion �� 9���10. Femur II l��, d rudimentary, v���� 7���11; tibia II d 20���22, l�� 6���9, v�� 12���16, v���� 15���22; tarsus II pv�� 5���7, tc�� 10���16, tc���� 25���28, u�� spine-like, 8���9, solenidion �� 10. Tibia III d 22, l�� 9, v�� 20, v���� 24; tarsus III pv�� 7, tc�� 22���25, tc���� 24���30, u�� spine-like, 8���9, u���� 7. Trochanter IV bare; femorogenu IV v�� 4���5, tibia IV solenidion �� 7; tarsus IV absent. LARVA (Figs 11���12, n = 4) Gnathosoma. Length 39���50, width 37���47. Palps 8���10. Cheliceral stylets 15���19, without teeth. Pharynx length 21���34, width 12���16. Dorsal setae 16���18, ventral setae 10���19. Idiosoma. Length 249���283, width 140���194. Prodorsal plate smooth, 66���70 long, 98���114 wide; with 3 pairs of weakly barbed setae (v 1 14���17, sc1 7���9, sc 2 20���27) and vestigial v2. Distance between setae v1-v 1 25���38, v2-v2 37 ���48, sc1-sc1 67���84, sc2-sc2 40��� 48, v1 -sc2 50���55. Plates C, D, EF smooth. Setae weakly barbed, c 1 10���13, c 2 11���14, d 10���12, e 7���12, f 8���12. Distance between setae c1-c1 65���86, c2-c2 131���138, c1-c 2 30���35, d-d 48���53, e-e 42���51, f-f 20���26. Plate H smooth, setae h1 attenuate,> 100, h2 ventral, > 50. Distance between setae h 1 27���39. Cupules ia, im, ih present. Setae ps1-2 attenuate, ps1 > 40, ps2 > 100. Coxisternal setae attenuate, 1a 18���34, 2a 20��� 32, 3a > 190, 3b > 110. Distance between setae 1a-1a 21���24, 2a-2a 39���52, 3a-3a 44���54, 3b-3b 60���64. Legs. Femur I, l��, l����, d rudimentary, v���� 11���15; genu I l�� 2���5, l���� 4���5, v�� 3���4, v���� 3���4; tibia and tarsus I fused, tibial setae d 15���26, l�� 3���4, l���� 2���4, v�� 5���6, v���� 6���8, solenidion �� 3���4, tarsal setae tc�� 9���12, tc���� 9���12, pv�� 5���11, pv���� 5���8, s spine-like, 6���7, solenidion �� 6���7. Femur II l��, d rudimentary, v���� 7���10; tibia II d 13���20, l�� 8���13, v�� 10���13, v���� 12���16; tarsus II tc�� 15, tc���� 24���30, p�� 6���7, u�� spine-like, 6���7, solenidion �� 4���5. Tibia III d 12���15, l�� 5���8, v�� 12���14, v���� 20���23; tarsus III tc�� 15���19, tc���� 30���57, p�� 5���6, u�� spine-like, 6���7, u���� rudimentary. Leg I without claws. Etymology. The species name boliviensis refers to the country where the type specimens were collected. Differential diagnosis. Adults of Podotarsonemus boliviensis are very similar to those of Podotarsonemus ecuadorensis but can be distinguished by the following character states. Female P. boliviensis has extensive imbricate sculpturing on the dorsal plates, has a cheliceral tooth and lacks seta u���� on tarsus II. In contrast, female P. ecuadorensis has smooth dorsal plates, lacks cheliceral teeth and has seta u���� on tarsus II. Male P. boliviensis and P. ecuadorensis are easily distinguished by several characters: P. boliviensis is the only Podotarsonemus with copulatory flange remnants on the genital capsule; P. boliviensis has v�� on femorogenu IV and lacks setae v ��� and v���� on tibia IV (absent and present, respectively, in P. ecuadorensis); P. boliviensis has a shorter solenidion on tibia IV (7 vs 10���12); P. boliviensis lacks seta u���� on tarsus II (present in P. ecuadorensis) and retains seta u���� on tarsus III (absent in P. ecuadorensis); and setae e-e and f-f are placed closer: e -e 36���37 vs 18, f-f 29���30 vs 9. Remarks. Among the Bolivian specimens, one female (considered in the description as the aberrant specimen) has a similar body size but has numerous measurements that are generally greater than the ranges recorded for the type series. For example, its body setae are longer dorsally: v1 (29 vs 20���21), sc2 (100 vs 50���75), c1 (37 vs 15���18), c2 (> 100 vs 44���54), d (33 vs 15���20), e (25 vs 15���8), but shorter ventrally, 1a (21 vs attenuate> 35), 2b (35 vs attenuate> 60). Its leg setae are also longer: Leg I���Fe l���� (2 vs rudimentary), Fe v���� (27 vs 12���21), Ge l�� (14, 5���9), Ge v�� (9 vs 4���5), Ti l�� (15 vs 4���7), Ti v�� (15 vs 5���9), Ti �� (6 vs 3���4); Leg II���Ti l�� (22 vs 8���11), Ti v�� (38 vs 18���24); Leg III���Ti l�� (30 vs 11���15). However its distances between some body setae are shorter: d -d (68 vs 82���91), f -f (31 vs 39���50). This specimen also has imbricate sculpturing only behind sc2-sc2, c1-c1 and d -d., Published as part of Seeman, Owen D., Lindquist, Evert E. & Husband, Robert W., 2018, A new tribe of tarsonemid mites (Trombidiformes: Heterostigmatina) parasitic on tetrigid grasshoppers (Orthoptera), pp. 1-54 in Zootaxa 4418 (1) on pages 18-26, DOI: 10.11646/zootaxa.4418.1.1, http://zenodo.org/record/1242435

Key to species of Podotarsonemus 1. Female and male: with seta on trochanter IV (e.g., Figs 2, 4); female: coxisternal plates III���IV separate, longitudinal striae between coxisterna (Fig. 2); setae h small to moderate length, not attenuate (length australiensis species group................................................................... 2 - Female and male: without seta on trochanter IV; female: coxisternal plates III���IV not separate, lacking longitudinal striae between coxisterna (Fig. 8); setae h attenuate (length> 70); male: tarsus IV absent ��� boliviensis species group........... 4 2. Female: setae sc1 absent (Fig. 1); genua II���III with setae (l ���, l������, v ���), femur I with four setae (l ���, l������, d, v������), femorogenu IV with two setae; male: tibia and tarsus I separate; femorogenu IV with seta, tarsus IV developed, with claw and setae (Fig. 3)............................................................................ Podotarsonemus australiensis sp. nov. - Female: setae sc1 present (Figs 16, 25); genua II���III without setae, femur I with two setae (l���, v ���), femorogenu IV bare; male: tibia and tarsus I fused; femorogenu IV without seta, tarsus IV rudimentary, without claw and setae (Figs. 18, 26)......... 3 3. Female: vestigial v2 formed as alveoli separate from lateral edges of prodorsal shield (Fig. 16), tracheal atria fused, post-atrial sacs heavily sclerotized and with long distal extension (Fig. 17); male: setae c2 much longer (> 4x) than c1, tibia IV bearing attenuate terminal seta, solenidion subterminal (Fig. 18); larva: setae h1-2, ps2 attenuate, tibia and tarsus I separate (Fig. 19)........................................................................... Podotarsonemus indicus sp. nov. - Female: vestigial v2 formed as anterolateral invaginations of lateral edges of prodorsal shield (Fig. 25), tracheal atria separate, post-atrial sacs lightly sclerotized and without distal extension (Fig. 25); male: setae c2 slightly longer (c1, tibia IV without terminal seta, solenidion terminal (Fig. 26); larva: setae h1-2, ps2 short, tibia and tarsus I fused (Fig. 27).................................................................................. Podotarsonemus zuluensis sp. nov. 4. Female and male: femur I with 4 setae (rudimentary l ���, l������, d and developed v������), femur II bearing 3 setae (rudimentary l ���, d and developed v������) (e.g., Fig. 8).............................................................................. 5 - Female and male: femur I with 2 or 3 setae (if present, d rudimentary, and developed l������, v������), femur II bearing 1 seta (developed v������) (e.g., Fig. 20)..................................................................................... 6 5. Female: prodorsal, C and D plates with imbricate sculpturing; chelicerae with 1 tooth (Fig. 7); male: femorogenu IV with 1 seta, small genital capsule flanges present (Fig. 9)............................... Podotarsonemus boliviensis sp. nov. - Female: prodorsal, C and D plates with punctate sculpturing; chelicerae without teeth (Fig. 13); male: femorogenu IV without seta, genital capsule flanges absent (Fig. 14)................................. Podotarsonemus ecuadorensis sp. nov. 6. Female: setae sc1 present; vestigial setae v2 expressed as alveolar pits within prodorsal shield; palp with two setae; femur I with rudimentary seta d (Fig. 20)............................................ Podotarsonemus nipponicus sp. nov. - Female: setae sc1 absent; rudimentary setae v2 expressed as anterolateral invaginations of prodorsal shield; palp with one seta; femur I without rudimentary seta d (Fig. 23).............................. Podotarsonemus queenslandensis sp. nov., Published as part of Seeman, Owen D., Lindquist, Evert E. & Husband, Robert W., 2018, A new tribe of tarsonemid mites (Trombidiformes: Heterostigmatina) parasitic on tetrigid grasshoppers (Orthoptera), pp. 1-54 in Zootaxa 4418 (1) on page 47, DOI: 10.11646/zootaxa.4418.1.1, http://zenodo.org/record/1242435

Podotarsonemus queenslandensis Seeman, Lindquist & Husband sp. nov. Figures 23���24 Type material. Holotype. Adult female. AUSTRALIA, Split Rock, 14 km SW of Laura, Far North Queensland, 23���26 Jun 1975, GB Monteith, ex Hedotettix amplus (Sj��stedt) (Tetrigidae) (Fig. 29B). In QM. Paratypes. 9 females, same data as holotype. In QM except 1 female in each of CNCI and UMMZ. Diagnosis. Female: Palps with 1 seta; chelicerae with 1 weak tooth; prodorsal plate with imbricate sculpturing behind setae sc2; setae sc1 absent; vestigial setae v2 expressed as anterolateral invaginations; femora I���II without seta l��, d; genua II���III bare; tarsi II���III with 5 setae, seta u���� present; trochanter IV without seta; coxisternal plates III���IV separated by soft cuticle; setae h attenuate,> 150 long; femorogenu IV with 2 setae; tracheal atria separate, post-atrial sac lightly sclerotized, not punctate. Description. ADULT FEMALE (Figures 23���24, n = 10 measured) Gnathosoma. Length 46���49, width 42���47. Palps 8���11, with 1 seta (5 long). Cheliceral stylets 16���19, each with 1 weakly formed tooth. Pharynx length 31���34, width 15���16. Dorsal setae ch 15���22, ventral setae su 15���23, palpcoxal setae e 11���13. Idiosoma. Length 200 (physogastric 525), width 130���140. Tracheal atria separate, post-atrial sac lightly sclerotized, not punctate, with two small distal prongs indicating separate tracheae. Prodorsal plate 78���83 long, 102���114 wide, with punctate sculpturing anteriorly, imbricate sculpturing posterior to sc2; with 2 pairs of setae: v 1 16���18, thickened, with small barbs, sc2 57���64, attenuate, and vestigial v2 expressed as anterolateral invaginations. Distance between setae v1-v 1 31���35, v2-v2 49, sc2-sc2 55��� 60, v1 -sc 2 18���23. Plates C, D with punctate sculpturing anteriorly, imbricated posteriorly, EF, H with punctate sculpturing, H heavily so. Setae c1, d, e, f thickened, with small barbs, c 1 15���18, c 2 30���36, d 13���15, e 10���13, f 11���13. Distance between setae c1-c1 66���91, c2-c2 120���150, c1-c 2 25���31, d-d 50���59, e-e 45���49, f-f 25���30. Setae h attenuate,> 140. Distance between setae h-h 21���28. Cupules ia, im present, ih indiscernible. Setae ps attenuate 260, distance between setae ps-ps 8���10. Coxisternal setae attenuate, 1a 8���10, 2a 15���22, 3a > 130, 3b > 90. Distance between setae 1a-1a 19���20, 2a-2a 40���41, 3a-3a 9���18, 3b-3b 26; alveolar vestiges of setae 1b, 2b present. Legs. Leg lengths: I 45 ���47, II 46 ���48, III 46 ���47. Leg IV Fe-Ge length 18���20, Ti-Ta length 17���18. Leg III, femur and genu fused. Femur I l���� 3���5, v���� 13���18; genu I l�� 7���12, l���� 6���8, v�� 4���5, v���� 5���7; tibia and tarsus I fused, tibial setae d 17���20, l�� 6���7, l���� 5���6, v�� 6���8, v���� 7���13, solenidion �� 3���4, tarsal setae tc�� 10���11, tc���� 10���12, pv�� 9���10, pv���� 7���8, s spine-like, 6��� 7, solenidion �� 7. Femur II v���� 11���15; tibia II d 15���18, l�� 9���11, v�� 13���16, v���� 13���17; tarsus II tc�� 12���14, tc���� 16���21, pv�� 12���14, u�� spine-like, 7, u���� 1, solenidion �� 4���6. Tibia III d 19���26, l�� 13���15, v�� 11���15, v���� 14���19; tarsus III tc�� 17���19, tc���� 16���17, pv�� 16���17, u�� spine-like, 6���7, u���� 1. Femorogenu IV with 2 setae, proximal femoral seta 20���23, distal genual seta 40���47; tibiotarsus with attenuate ventral tibial seta> 80 and terminal tarsal seta> 210. ADULT MALE���Unknown LARVA���Unknown Etymology. The species name queenslandensis refers to the state (Queensland, Australia) where the type specimens were collected. Remarks. The female of Podotarsonemus queenslandensis is most similar to that of P. nipponicus and can be distinguished in the Differential diagnosis for that species., Published as part of Seeman, Owen D., Lindquist, Evert E. & Husband, Robert W., 2018, A new tribe of tarsonemid mites (Trombidiformes: Heterostigmatina) parasitic on tetrigid grasshoppers (Orthoptera), pp. 1-54 in Zootaxa 4418 (1) on page 40, DOI: 10.11646/zootaxa.4418.1.1, http://zenodo.org/record/1242435

Podotarsonemus boliviensis Seeman, Lindquist & Husband sp. nov. Figures 7–12, 15A, B Type material. Holotype. Adult female. BOLIVIA, Dept. Beni, Puertofonbeni, Luis E. Peña, 12 Nov. 1956, ex Batrachidea mucronata (Serville) (Tetrigidae). G# 23, RWH 15I 2010 (slide 2). In UMMZ. Paratypes. 20 females, 5 males, 11 larvae, as follows: 14 females, 4 males, 8 larvae, same data as holotype. 1 female, 1 male, same data as holotype except additional collection data “on way to Reyes from Rio Beni ″, 6–10 Nov. 1956. 5 females, 3 larvae, Dept. La Laz, Altamarani on Beni River, Luis E. Peña, 5–11 Nov. 1956, ex B. mucronata. In UMMZ except 1 female, 1 male, 1 larva in each of CNCI and QM. Non-type material. 1 adult female, same data as holotype, RWH 15I 2010 (slide 8)—aberrant specimen. 1 pharate female, 8 eggs, same data as holotype. In UMMZ. Diagnosis. Female and male: Palps with 1 seta; chelicera with 1 tooth (female) or without teeth (male); prodorsal plate with imbricate sculpturing; femora I–II with rudimentary seta lʹ, d; genua II–III bare; tarsi II–III with 4 setae, lacking seta u″ (except male tarsus III with 5 setae, u″ present); trochanter IV bare. Female: Setae sc1 present; vestigial setae v2 expressed as alveoli within prodorsal shield; coxisternal plates III–IV separated by soft cuticle; setae h attenuate,> 70 long; femorogenu IV with 2 setae; tracheal atria separate, post-atrial sac lightly sclerotized, not punctate. Male: Genital capsule with small copulatory flanges; setae h1 rudimentary, like h2; femorogenu IV with 1 seta; tibia IV with solenidion, without setae; tarsus IV absent. Male and larva: Leg I with tibia and tarsus fused. Larva: Setae h1-2, ps1-2 attenuate; leg I without claws. Description. ADULT FEMALE (Figures 7–8, 15A, B, n = 11 measured) Measurements are in the form of: Range [Measurement of aberrant specimen if outside of range]. Gnathosoma. Length 51–63, width 59–66 [69]. Palps 9–14, with 1 apical seta 7–9 flanked medially by pointed triangular hyaline process. Cheliceral stylets 1 9–23, each with 1 tooth. Pharynx length 26–42, width 19–25. Dorsal setae ch 25–34, ventral setae su 8–23 [30], palpcoxal setae e 7–13. Idiosoma. Length 228–245, width 159–198. Tracheal atria separate, post-atrial sac lightly sclerotized, not punctate, terminal tracheae separate. Prodorsal plate with imbricate sculpturing, irregular anteriorly [similar, but smooth between sc2-sc2], 91–106 long, 140–163 wide; with 3 pairs of setae: v 1 20–26 [29] with few barbs, sc 1 5– 22 variable, mostly 5–6, but 14 and 22 in two specimens, sc2 47–75 [100] and vestigial v2. Distance between setae v1-v1 46 –56 [45], v2-v2 56 –64 [68], sc1-sc1 120–131 [110], sc2-sc2 67– 82, v1 -sc 2 26–32. Plates C, D, EF with imbricate sculpturing [similar, but smooth between c1-c1]. Setae c1, d, e, f with few barbs [many barbs], c 1 15–23 [37], c2 44–54 [> 100], d 15–22 [33], e 15–22 [25], f 15–22. Distance between setae c1-c1 109–123 [95], c2-c2 154–170, c1-c 2 23–30, d-d 82–96 [68], e-e 71–80, f-f 36–50 [31]. Plate H folded beneath EF, setae h attenuate,> 130. Distance between setae h-h 26–37. Cupules ia, im, ih present. Setae ps attenuate> 130, distance between setae ps-ps 10–13 [14]. Coxisternal setae attenuate, 1a 90 (often broken), 2a 96–135 (often broken), 3a > 250, 3b > 210. Distance between setae 1a-1a 21–28 [31], 2a-2a 33–47 [57], 3a-3a 18–31 [11], 3b-3b 33–50 [55]; alveolar vestiges of setae 1b and 2b present. Legs. Leg lengths: I 55 –60 [75], II 51 –56 [74], III 53 –55 [78]. Leg IV Fe-Ge length 25–29, Ti-Ta length 18–21. Leg III, femur and genu fused. Femur I, lʹ, lʹʹ, d rudimentary [lʹʹ 2], vʹʹ 12–21 [27]; genu I lʹ 4–9 [14], lʹʹ 6–10, vʹ 4– 5 [9], vʹʹ 4–6 [9]; tibia and tarsus I fused, tibial setae d 28–41, lʹ 4–9 [15], l″ 3 [5], vʹ 5–9 [15], vʹʹ 8–13 [14], solenidion φ 3–4 [6], tarsal setae tcʹ 15 [17], tcʹʹ 13–15 [17], pvʹ 9–15, pvʹʹ 6–10 [17], s spine-like, 5–9, solenidion ω 8–11. Femur II lʹ, d rudimentary, vʹʹ 7–19 [24]; tibia II d 30–39, lʹ 8–21 [22], vʹ 7–24 [38], vʹʹ 19–33 [35]; tarsus II pvʹ 12–16 [22], tcʹ 19–22 [25], tcʹʹ 27–45, uʹ spine-like, 6–10, solenidion ω 8–12. Tibia III d 23–38 [40], lʹ 11–15 [30], vʹ 17–33, vʹʹ 25–31 [34]; tarsus III pvʹ 11–24, tcʹ 21–29 [31], tcʹʹ 26–46, uʹ spine-like, 8–9. Femorogenu IV with 2 attenuate ventral setae (Fe- vʹ, Ge- vʹ); proximal femoral seta 18–25 [36], distal genual seta 36–45 [28]; tibiotarsus with attenuate ventral tibial seta and terminal tarsal seta> 100. ADULT MALE (Figs 9–10, n = 2 measured) Gnathosoma. Similar to female. Length 33–36, width 35–36. Palps 10–12. Cheliceral stylets 15–17, without teeth. Pharynx length 20–22, width 10–11. Dorsal setae 10–12, ventral setae 7–10, palpcoxal setae 2–3. Idiosoma. Length 165–175, width 140. Prodorsal plate punctate, 65–73 long, 105–110 wide; with 3 pairs of setae (v 1 16–17, sc1 4–5, sc2 39–42) and vestigial v2. Distance between setae v1-v 1 29–31, v2-v 2 30–34, sc1-sc1 72–75, sc2-sc2 41– 44, v1 -sc2 41–43. Plates CD, EF punctate. Setae smooth, setae c1 6–7, c 2 25–26, d 35–40 (left side d 8 on one specimen); setae c1 obliquely aligned between setae c2 and d. Plate EF, e 5, f 6. Distance between setae c1-c1 75, c2-c2 100–115, c1-c 2 28–32, d-d 26–27, e-e 36–37, f-f 29–30. Cupules ia, im present. Genital capsule 30–35 long, 32 wide, bearing alveolar remnants of h1, h2; ventral flanges present, small. Coxisternal setae attenuate, 1a 28, 2a 70, 3a > 90, 3b > 80. Distance between setae 1a-1a 13–15, 2a-2a 38–40, 3a-3a 10–13, 3b-3b 37–40; setae 1a at level of or just posterior to ap1, setae 2b at level of or just posterior to ap2. Apodemes: ap2 not reaching medially obsolete ap sj; ap5 small, extending 17–21 from base, not fused medially. Legs. Leg lengths: I 47, II 46, III 45, IV 30–31. Leg III, femur and genu fused. Femur I, lʹ, lʹʹ, d rudimentary, lʹʹ on small tubercle, vʹʹ 8–10; genu I lʹ 2–3, lʹʹ 4–5, vʹ 3, vʹʹ 2–3; tibia and tarsus I fused, tibial setae d 23–25, lʹ 4, lʹʹ 2–3, vʹ 6–7, vʹʹ 10–15, solenidion φ 3–4, tarsal setae ftʹ 12, ftʹʹ 12, tcʹ 11–12, tcʹʹ 9– 11, pvʹ 9–11, pvʹʹ 9–11, s spine-like, 8, solenidion ω 9–10. Femur II lʹ, d rudimentary, vʹʹ 7–11; tibia II d 20–22, lʹ 6–9, vʹ 12–16, vʹʹ 15–22; tarsus II pvʹ 5–7, tcʹ 10–16, tcʹʹ 25–28, uʹ spine-like, 8–9, solenidion ω 10. Tibia III d 22, lʹ 9, vʹ 20, vʹʹ 24; tarsus III pvʹ 7, tcʹ 22–25, tcʹʹ 24–30, uʹ spine-like, 8–9, uʹʹ 7. Trochanter IV bare; femorogenu IV vʹ 4–5, tibia IV solenidion φ 7; tarsus IV absent. LARVA (Figs 11–12, n = 4) Gnathosoma. Length 39–50, width 37–47. Palps 8–10. Cheliceral stylets 15–19, without teeth. Pharynx length 21–34, width 12–16. Dorsal setae 16–18, ventral setae 10–19. Idiosoma. Length 249–283, width 140–194. Prodorsal plate smooth, 66–70 long, 98–114 wide; with 3 pairs of weakly barbed setae (v 1 14–17, sc1 7–9, sc 2 20–27) and vestigial v2. Distance between setae v1-v 1 25–38, v2-v2 37 –48, sc1-sc1 67–84, sc2-sc2 40– 48, v1 -sc2 50–55. Plates C, D, EF smooth. Setae weakly barbed, c 1 10–13, c 2 11–14, d 10–12, e 7–12, f 8–12. Distance between setae c1-c1 65–86, c2-c2 131–138, c1-c 2 30–35, d-d 48–53, e-e 42–51, f-f 20–26. Plate H smooth, setae h1 attenuate,> 100, h2 ventral, > 50. Distance between setae h 1 27–39. Cupules ia, im, ih present. Setae ps1-2 attenuate, ps1 > 40, ps2 > 100. Coxisternal setae attenuate, 1a 18–34, 2a 20– 32, 3a > 190, 3b > 110. Distance between setae 1a-1a 21–24, 2a-2a 39–52, 3a-3a 44–54, 3b-3b 60–64. Legs. Femur I, lʹ, lʹʹ, d rudimentary, vʹʹ 11–15; genu I lʹ 2–5, lʹʹ 4–5, vʹ 3–4, vʹʹ 3–4; tibia and tarsus I fused, tibial setae d 15–26, lʹ 3–4, lʹʹ 2–4, vʹ 5–6, vʹʹ 6–8, solenidion φ 3–4, tarsal setae tcʹ 9–12, tcʹʹ 9–12, pvʹ 5–11, pvʹʹ 5–8, s spine-like, 6–7, solenidion ω 6–7. Femur II lʹ, d rudimentary, vʹʹ 7–10; tibia II d 13–20, lʹ 8–13, vʹ 10–13, vʹʹ 12–16; tarsus II tcʹ 15, tcʹʹ 24–30, pʹ 6–7, uʹ spine-like, 6–7, solenidion ω 4–5. Tibia III d 12–15, lʹ 5–8, vʹ 12–14, vʹʹ 20–23; tarsus III tcʹ 15–19, tcʹʹ 30–57, pʹ 5–6, uʹ spine-like, 6–7, uʹʹ rudimentary. Leg I without claws. Etymology. The species name boliviensis refers to the country where the type specimens were collected. Differential diagnosis. Adults of Podotarsonemus boliviensis are very similar to those of Podotarsonemus ecuadorensis but can be distinguished by the following character states. Female P. boliviensis has extensive imbricate sculpturing on the dorsal plates, has a cheliceral tooth and lacks seta uʹʹ on tarsus II. In contrast, female P. ecuadorensis has smooth dorsal plates, lacks cheliceral teeth and has seta uʹʹ on tarsus II. Male P. boliviensis and P. ecuadorensis are easily distinguished by several characters: P. boliviensis is the only Podotarsonemus with copulatory flange remnants on the genital capsule; P. boliviensis has vʹ on femorogenu IV and lacks setae v ′ and vʹʹ on tibia IV (absent and present, respectively, in P. ecuadorensis); P. boliviensis has a shorter solenidion on tibia IV (7 vs 10–12); P. boliviensis lacks seta uʹʹ on tarsus II (present in P. ecuadorensis) and retains seta uʹʹ on tarsus III (absent in P. ecuadorensis); and setae e-e and f-f are placed closer: e -e 36–37 vs 18, f-f 29–30 vs 9. Remarks. Among the Bolivian specimens, one female (considered in the description as the aberrant specimen) has a similar body size but has numerous measurements that are generally greater than the ranges recorded for the type series. For example, its body setae are longer dorsally: v1 (29 vs 20–21), sc2 (100 vs 50–75), c1 (37 vs 15–18), c2 (> 100 vs 44–54), d (33 vs 15–20), e (25 vs 15–8), but shorter ventrally, 1a (21 vs attenuate> 35), 2b (35 vs attenuate> 60). Its leg setae are also longer: Leg I—Fe lʹʹ (2 vs rudimentary), Fe vʹʹ (27 vs 12–21), Ge lʹ (14, 5–9), Ge vʹ (9 vs 4–5), Ti lʹ (15 vs 4–7), Ti vʹ (15 vs 5–9), Ti φ (6 vs 3–4); Leg II—Ti lʹ (22 vs 8–11), Ti vʹ (38 vs 18–24); Leg III—Ti lʹ (30 vs 11–15). However its distances between some body setae are shorter: d -d (68 vs 82–91), f -f (31 vs 39–50). This specimen also has imbricate sculpturing only behind sc2-sc2, c1-c1 and d -d.

Podotarsonemus queenslandensis Seeman, Lindquist & Husband sp. nov. Figures 23–24 Type material. Holotype. Adult female. AUSTRALIA, Split Rock, 14 km SW of Laura, Far North Queensland, 23–26 Jun 1975, GB Monteith, ex Hedotettix amplus (Sjöstedt) (Tetrigidae) (Fig. 29B). In QM. Paratypes. 9 females, same data as holotype. In QM except 1 female in each of CNCI and UMMZ. Diagnosis. Female: Palps with 1 seta; chelicerae with 1 weak tooth; prodorsal plate with imbricate sculpturing behind setae sc2; setae sc1 absent; vestigial setae v2 expressed as anterolateral invaginations; femora I–II without seta lʹ, d; genua II–III bare; tarsi II–III with 5 setae, seta uʹʹ present; trochanter IV without seta; coxisternal plates III–IV separated by soft cuticle; setae h attenuate,> 150 long; femorogenu IV with 2 setae; tracheal atria separate, post-atrial sac lightly sclerotized, not punctate. Description. ADULT FEMALE (Figures 23–24, n = 10 measured) Gnathosoma. Length 46–49, width 42–47. Palps 8–11, with 1 seta (5 long). Cheliceral stylets 16–19, each with 1 weakly formed tooth. Pharynx length 31–34, width 15–16. Dorsal setae ch 15–22, ventral setae su 15–23, palpcoxal setae e 11–13. Idiosoma. Length 200 (physogastric 525), width 130–140. Tracheal atria separate, post-atrial sac lightly sclerotized, not punctate, with two small distal prongs indicating separate tracheae. Prodorsal plate 78–83 long, 102–114 wide, with punctate sculpturing anteriorly, imbricate sculpturing posterior to sc2; with 2 pairs of setae: v 1 16–18, thickened, with small barbs, sc2 57–64, attenuate, and vestigial v2 expressed as anterolateral invaginations. Distance between setae v1-v 1 31–35, v2-v2 49, sc2-sc2 55– 60, v1 -sc 2 18–23. Plates C, D with punctate sculpturing anteriorly, imbricated posteriorly, EF, H with punctate sculpturing, H heavily so. Setae c1, d, e, f thickened, with small barbs, c 1 15–18, c 2 30–36, d 13–15, e 10–13, f 11–13. Distance between setae c1-c1 66–91, c2-c2 120–150, c1-c 2 25–31, d-d 50–59, e-e 45–49, f-f 25–30. Setae h attenuate,> 140. Distance between setae h-h 21–28. Cupules ia, im present, ih indiscernible. Setae ps attenuate 260, distance between setae ps-ps 8–10. Coxisternal setae attenuate, 1a 8–10, 2a 15–22, 3a > 130, 3b > 90. Distance between setae 1a-1a 19–20, 2a-2a 40–41, 3a-3a 9–18, 3b-3b 26; alveolar vestiges of setae 1b, 2b present. Legs. Leg lengths: I 45 –47, II 46 –48, III 46 –47. Leg IV Fe-Ge length 18–20, Ti-Ta length 17–18. Leg III, femur and genu fused. Femur I lʹʹ 3–5, vʹʹ 13–18; genu I lʹ 7–12, lʹʹ 6–8, vʹ 4–5, vʹʹ 5–7; tibia and tarsus I fused, tibial setae d 17–20, lʹ 6–7, lʹʹ 5–6, vʹ 6–8, vʹʹ 7–13, solenidion φ 3–4, tarsal setae tcʹ 10–11, tcʹʹ 10–12, pvʹ 9–10, pvʹʹ 7–8, s spine-like, 6– 7, solenidion ω 7. Femur II vʹʹ 11–15; tibia II d 15–18, lʹ 9–11, vʹ 13–16, vʹʹ 13–17; tarsus II tcʹ 12–14, tcʹʹ 16–21, pvʹ 12–14, uʹ spine-like, 7, uʹʹ 1, solenidion ω 4–6. Tibia III d 19–26, lʹ 13–15, vʹ 11–15, vʹʹ 14–19; tarsus III tcʹ 17–19, tcʹʹ 16–17, pvʹ 16–17, uʹ spine-like, 6–7, uʹʹ 1. Femorogenu IV with 2 setae, proximal femoral seta 20–23, distal genual seta 40–47; tibiotarsus with attenuate ventral tibial seta> 80 and terminal tarsal seta> 210. ADULT MALE—Unknown LARVA—Unknown Etymology. The species name queenslandensis refers to the state (Queensland, Australia) where the type specimens were collected. Remarks. The female of Podotarsonemus queenslandensis is most similar to that of P. nipponicus and can be distinguished in the Differential diagnosis for that species.

Lasioseius fuscina n. sp. (Figures 8, 9, 11C, 12D, 13 E-F, 14D) Zoobank: F088E970-F8B8-4E76-8382-87DA2EF12B7F Diagnosis — Adult female: Sternal shield lineated laterally and anteromedially, its anterior margin with blunt medial projection, its posterolateral corners not emarginated; poroids iv3 and setae st4 on slender metasternal platelets; epigynal shield reticulated over nearly entire surface, its lateral margins concave, narrowed anterior to setae st5; ventrianal shield with posterolateral margins on either side of anus nearly parallel, and with four pairs of opisthogastric setae, JV1 on shield; metapodal plates nearly connected or coalesced on each side. Dorsal setae r6 on edge of dorsal shield; Z5, S5 attenuated like adjacent setae. Adult male: Sternitigenital shield incompletely reticulated, nearly smooth medially. Dimorphic leg setal structures minimal. Adult attributes: Gnathotectum anterior margin with 3 elongated, denticulated tines. Fixed chela with file of 8-9 teeth in female, ca 4 in male. Sternal setae st1 inserted on anterior edge of sternal shield. Tarsi II-IV with ventral apical process bluntly pointed, smooth apically; tarsi II-III with setae (l-1) shorter, more spinelike than (v- 1); tarsus IV with distal seta pv-1 longer, more attenuate than av-1, (l-1); pretarsus with anterior paradactylus slightly longer than posterior one. Description Idiosomatic dorsum — Adult female. Dorsal shield 424 – 450 long, 201 – 245 wide at level of r3, 267 – 296 wide at level of r6 (n=7), lightly reticulate-lineate on lateral surfaces but nearly smooth medially between bases of setae j4 to J4, and sparsely punctate behind level of setae Z4; shield with 38 pairs of setae generally similar in size and acutely pointed form, with few apical barbs, more so on Z4, Z5, S4, S5 (Figs 8A, B); J1-J3 and Z1-Z4 slightly longer than longitudinal intervals between their bases, S2-S4 clearly so; J5 minute, barbed (4 – 6): j1 (28 – 31), j2 (26 – 32), j3 (41 – 44), j4 (26 – 32), j5 (32 – 35), j6 (45 – 47), z1 (19 – 24), z2 (30 – 35), z3 (37 – 42), z4 (41 – 45), z5 (32 – 37), z6 (41 – 46); s1 (32 – 38), s2 (34 – 43), s3 (40 – 45), s4 (43 – 48), s5 (45 – 50), s6 (47 – 52), r2 (35 – 42), r3 erect (21 – 30), r4 (32 – 48), r5 (32 – 50), r6 (32 – 48), J1 (32 – 42), J2 (40 – 50), J3 (43 – 52), J4 (43 – 52), Z1 (48 – 51), Z2 (51 – 54), Z3 (54 – 58), Z4 (58 – 62), Z5 (63 – 70), S1 (43 – 50), S2 (44 – 53), S3 (46 – 58), S4 (46 – 58), S5 (43 – 52). Lateral soft cuticle with seven pairs of smooth setae, including R1 (40 – 48), R2-R6 (35 – 42) and ZV5 posteriorly (30 – 33) (Fig. 8D). Peritrematal shield fused to dorsal shield at level of s1. Adult male. Dorsal shield 338 – 340 long, 185 – 200 wide at level of setae r3, 200 – 215 wide at level of r6 (n = 2); shield ornamentation, complement of pore-like structures and setation as in female except slightly more expanded laterally so as to become united with peritrematal shields at level of setae s2 and bear setae R1. Dorsal shield with 39 pairs of setae, their attenuated form and relative lengths as in female. Setae j1 (21 – 23), j2 (16 – 26), j3 (35 – 36), j4 (21 – 25), j5 (24 – 25), j6 (32 – 33), z1 (14 – 16), z2 (22), z3 (32), z4 (34 – 35), z5 (27 – 28), z6 (32 – 36), s1 (29 – 30), s2 (30 – 31), s3 (34 – 38), s4 (36), s5 (42), s6 (35 – 36), r2, r4, r5 (33 – 37), r3 erect, shorter (23 – 27), r6 (36 – 38), J1-J3 (27 – 32), J4 (25 – 27), J5 (3 – 4), Z1-Z2 (40 – 43), Z3 (44 – 45), Z4, Z5 longest (48 – 54), S1, S2 (35 – 41), S3 (41 – 45), S4 (40 – 43), S5 (32 – 35). Lateral soft cuticle with six pairs of setae, R2-R6 (27 – 37) followed by ZV5. Idiosomatic venter — Adult female. Tritosternum base 26 – 28 long and 10 – 14 wide at its base, with paired laciniae fused for about 33 – 37 of entire length (96 – 100). Presternal region with circumscribed pair of jagged-lineate, granulated areas adhered to anterior margin of sternal shield, sometimes flanked anteriorly by a pair of narrow slightly sclerotized strips on either side of tritosternum base. Sternal shield 87 – 93 long from slight, blunt medial projection of anterior margin to gently concave posterior margin, 85 – 95 wide, weakly reticulate-lineate over much of surface or nearly smooth medially, with arcuate transverse line between poroids iv1 (Fig. 8D); sternal shield setae of similar lengths, st1 (15 – 20), st2, st3 (11 – 16). Sternal poroids iv3 and setae st4 (6 – 14) inserted together on small metasternal platelets (15 – 20). Endopodal strips narrow between coxae III and IV. Epigynal shield lightly reticulate over entire surface, its narrowly rounded hyaline margin nearly reaching or slightly overlapping posterior edge of sternal shield, its lateral margins concave, narrowed anterior to setae st5 (11 – 15), and its posterolateral corners slightly emarginated behind st5; shield 140 – 147 long, 78 – 87 at widest level at posterior rounded corners, 66 – 71 at level of st5, and 49 – 54 at narrowest width between legs IV. Opisthosomatic venter with two pairs of metapodal platelets touching or coalesced on each side, primary pair ovoid to subtriangular, 20 – 25 long, 10 – 12 wide, secondary pair smaller, greatest transverse dimension 8 – 12. Ventrianal shield inversely subtriangular with irregularly rounded anterior corners, with posterolateral margins on either side of anus nearly parallel; surface entirely lightly sculptured, with primarily transverse lineation changing to reticulation behind setae JV2-ZV2, except anal region sparsely punctate on either side of postanal seta; shield wider (160 – 183) than long including cribrum (136 – 147), width at anal level (95 – 104); gland pore gv3 on shield edge posterior to anal opening; shield with four pairs of ventral setae: JV1 (12 – 17), JV2 (22 – 27), JV3 (24 – 31), ZV2 (22 – 27); with paranal setae (11 – 14) shorter than postanal seta (28 – 35). Soft cuticle with four pairs of simple opisthogastric setae, ZV1 (11 – 15) anterior to ventrianal shield, ZV3 (17 – 22), JV5 attenuated, barbed (48 – 55) flanking the shield, and ZV5 (30 – 33) somewhat aligned behind R6; three pairs of poroids (including ivp) on soft opisthogastric cuticle, ivo2 sometimes captured on anterolateral edge of ventrianal shield. Peritrematal-exopodal shield with lines extending from stigma into rounded posterior margin (Fig. 8D); peritreme extending to level between setae s1 and z1 (Fig. 8A). Spermathecal apparatus with cup-like calyx (12 – 15 long including embolus). Adult male. Tritosternum much as in female, but with base biarticulate, wider (6 – 11) than long (5 – 10), and laciniae (37 – 40) fused (8) for only about 1/5 of total length. Presternal region weakly ornamented, sometimes with pair of narrow strips on either side of tritosternum base. Sternitigenital shield length 158 – 162 from genital opening, widths 65 – 72 at level between coxae II, 50 between coxae IV; shield with a concave arcuate line between poroids iv1, lightly lineate-reticulate along lateral margins and posteriorly between coxae IV, unornamented medially between st2 and iv3, with endopodal apices freely acuminate beside coxae IV; setae st1-st5 of dissimilar lengths as in female, st1 (14 – 16), st2 (11 – 12), st3 (8 – 11), st4 shorter (4 – 8), st5 (7 – 8). Ventrianal shield reticulate as in female, its convex anterolateral margins entire behind apices of exopodal strips behind coxae IV, and posterolateral margins not constricted in anal region, its greatest width (173 – 189) larger than its length (123 – 141, including cribrum ca. 3 – 8 long, 73 – 80 wide); shield with seven pairs of opisthogastric setae, JV1, ZV1 (10 – 12), JV2, JV3 (25 – 29), ZV2 (20 – 26), ZV3 (17 – 20), and longest JV5 (40 – 43); postanal seta smooth (26 – 28), about three times as long as paranal setae (8 – 12) (Fig. 9A). Soft opisthogastric cuticle with two pairs of lyrifissures including ivp. Gnathosoma — Adult female. Gnathotectum anterior margin with three denticulate prongs of similar moderately elongate length or medial prong longer than laterals (Fig. 8C). Cheliceral shaft, excluding basal section, 139 – 155 long, with slender digits; dorsal seta simple (10 – 11); fixed digit with tiny subapical tooth and row of 7-10 fine teeth, pilus dentilis minute, at level of 3 rd apical tooth; movable digit (48 – 51) tridentate (Fig. 8E). Subcapitulum with deutosternal rows differing little in width, most anterior row with 9-10 teeth, nd 2 to 5 th rows with 8-15 teeth, basalmost two rows slightly wider, with ca 15-20 teeth (Fig. 8F); subcapitular setae hp1 (40 – 50) about as long as capitular setae pc (40 – 46), longer than hp3 (27 – 29), and hp2 (20 – 22). Palp length (113 – 122), apical margin of palptrochanter with two or usually three spines anterolaterally; palptrochanter with internal seta v1 sickle-shaped, longer (35 – 44) and thicker than v2 (24 – 30) (Fig. 8G). Adult male. Gnathotectum anterior margin with elongated prongs as in female (Fig. 9B). Cheliceral shaft, excluding short basal section 91 – 98 long; fixed digit with offset subapical tooth followed by pilus dentilis and sparse row of 4-5 small teeth (Fig. 9E); movable digit (27 – 28) unidentate; spermatodactyl 30 – 40 long, 3 – 6 wide at base, extending anteriorly ca 23 – 30 beyond apex of digit, with narrowed neck and small head apically (Fig. 9D). Corniculi slightly more widely spaced but not more arched than in female, 24 – 26 long, their bases flanked medially by pair of short (5 – 7) spiculate processes (Figs 9C, 11C); internal malae and labrum as in female. Subcapitulum with rows of deutosternal denticles much as in female, anterior five rows with ca 8-15 teeth, 6 th and 7 th rows with ca 15-20 teeth; hypostomatic setae hp1 (28 – 30) slightly attenuated, about as long as pc (24 – 31), longer than hp3 (20 – 21), and hp2 shortest (10 – 12) (Fig. 9D). Palpi (length 96 – 101) similar in structure, form of setae as in female; apical margin of palptrochanter with three spines anterolaterally; palptrochanter with inner seta (26 – 30) longer than outer seta (8 – 11) (Fig. 9D). Legs — Adult female. Legs I (357 – 385) subequally as long as legs IV (360 – 380), and shorter than dorsal shield; other leg lengths (excluding pretarsi): II 286 – 319, III 288 – 303. Leg I pretarsus with biarticulated shaft (16 – 21 to base of claws), claws 10 – 12 long; tarsus (107 – 112) about 1.9-2.1 longer than tibia; other length ratios, femur (70 – 77): genu (50 – 52): tibia (52 – 60), about 1.3: 0.9: 1.0. Legs II, III with tarsus/tibia length ratio about 1.8-2.0, leg IV with that ratio about 1.9-2.0. Chaetotactic formulae of leg segments as given for species-group. Tarsi II-IV with ventroapical process (7 – 9) bluntly pointed, smooth apically (Fig. 14D), and with setae (d- 1) (15 – 20) nearly as long as pretarsus to base of claws (17 – 20). Tarsi II-III with paradactyli reaching tips of tarsal claws, but tarsus IV with anterior paradactylus extending about 3-5 beyond claw (Fig. 14D). Tarsi II (Fig. 13F), III with setae (l-1) shorter (16 – 19), more spinelike than (v-1) (28 – 34); dorsal seta md (34 – 37) no longer than pd-2, pl-3, mv (34 – 35); setae al-2, ad-2 (36 – 41), pl-2 (43 – 44), al-3, (pd-3) (28 – 32); all tarsal setae smooth. Tarsus IV with distal setae v- (1) symmetrical in size (26 – 31), slenderer than somewhat spinelike (l- 1) (15 – 22). Adult male. Relative leg thicknesses and lengths, excluding pretarsi, similar to those of female, I 305 – 310, II 255 – 260, III 242 – 250, IV 302 – 305. Legs with segment length ratios as in female. Tarsus I pretarsus 10 – 15 to base of claws, claws 5 – 9 long. Tarsi II-IV with dorsoapical setal processes (d-1), ventroapical process and pretarsal structures formed as in female, pretarsus lengths 12 – 17 excluding claws (10) (Fig. 13E). Apical setal processes (d-1) (11 – 13) shorter than pretarsi to bases of claws (14 – 17). Dimorphic leg setal structures not apparent. Type material — All specimens collected at the La Selva Biological Station, Heredia Province, Costa Rica (10°26´N, 84°01´W, elevation 50-150 m): HOLOTYPE: adult female, 12 June 1997, ex Heliconia sp. leaves, successional plots, coll. ALAS. PARATYPES: 1 adult female, 1 adult male, with same data as holotype; 1 adult female, 12 June 1997, ex coexistent adults of Cephaloleia belti Baly and C. ornatrix Donckier from Heliconia leaves, coll. ALAS; 3 adult females, 15 April 1994, ex adult Chrysomelidae, no host plant data, coll. M. Paniagua; 2 adult females, 18 February 1994, ex adult Chrysomelidae, no host plant data, coll. ALAS; 3 adult females, 1 adult male, date not indicated, ex adult Cephaloleia belti on Calathea leaf, coll. not indicated. Host identifications — As noted for the records of Lasioseius serripes, the identities of the associated beetles given above are in accord with current molecular studies, which indicate that records of Cephololeia belti and C. ornatrix on Heliconia pogognatha are in accord with field diet records. While C. belti is a host generalist, C. ornatrix is a host specialist of Heliconia pogonantha (Staines & Garcia-Robledo 2014, Garcia-Robledo et al. 2017). Etymology — The specific name is based on the Latin noun,‘fuscina’, indicating a threepronged fork or trident, and refers to the elongated three-pronged gnathotectum characteristic of this species. Remarks — Limited collection data indicate a possible host association preference for Cephaloleia belti, a plant host generalist. If so, the mite/host size would correlate, as adults of L. ‘fuscina’ are the smallest of the described mites of this species-group, and among species of Cephaloleia, beetles of C. belti are relatively small-sized as well, although those of C. ornatrix are mid-sized. However, collection data uncertainties must be kept in mind, as two of the collection samples for this species also included many adults of Lasioseius serripes, one of them as well with a few adults of L. cassidini which also co-occurred with fuscina in two other samples (Table 1). Tentatively, subject to further studies, we conclude that, at La Selva, Lasioseius fuscina may be associated regularly with the host generalist Cephaloleia belti and to some extent with the host specialist C. ornatrix on Heliconia pogonantha.

Lasioseius cassidini n. sp. (Figures 1, 2, 11A, 12C, 13 A-B, 14A) Zoobank: EB77E489-1EB6-4120-9375-44E765ABEFF3 Diagnosis ��� Adult female: Sternal shield weakly lineate along lateral margins, smooth medially, its posterolateral corners not emarginated; poroids iv3 alone, on soft cuticle, setae st4 on tiny metasternal platelets; epigynal shield partially reticulated, its lateral margins concave, narrowed anterior to setae st5; ventrianal shield with posterolateral margins on either side of anus divergent anteriorly, and with four pairs of opisthogastric setae, JV1 on shield; metapodal plates well separated on each side. Dorsal setae r6 on edge of dorsal shield; Z5, S5 attenuated like adjacent setae. Adult male: Dorsal shield setae S5 dimorphically less attenuated. Sternitigenital shield incompletely reticulated, nearly smooth medially. Tibiae III-IV with setae (v) dimorphically spinelike, blunt; genu IV av similarly so. Adult attributes: Gnathotectum anterior margin truncate, irregularly denticulate. Fixed chela with file of ca 20 teeth in female, ca 18 in male. Sternal setae st1 inserted in weakly sclerotized presternal region. Tarsi II-IV with ventral apical process bluntly pointed, smooth apically; tarsi II-III with seta av-1 slightly longer than pv-1; pretarsi III-IV with anterior paradactylus asymmetrically elongate, twice as long as claws; tarsus IV with distal seta av-1 clearly larger, thicker than adjacent setae. Description Idiosomatic dorsum ��� Adult female. Dorsal shield 703 ��� 757 long, 375 ��� 410 wide at level of r3, 428 ��� 477 wide at r6 (n=9), lightly reticulated over entire surface, without puncta behind level of setae Z4; shield with 38 pairs of setae generally similar in size and acutely pointed form, J1-J4 about as long or slightly longer than longitudinal intervals between their bases; J5 minute, barbed (8 ��� 12): j1, j2 (45 ��� 60), j3 (67 ��� 76), j4 (50 ��� 57), j5 (56 ��� 62), j6 (70 ��� 77), z1 (37 ��� 42), z2 (56 ��� 65), z3 (60 ��� 68), z4 (72 ��� 77), z5 (52 ��� 62), z6 (62 ��� 74); s1 (58 ��� 75), s2 (65 ��� 75), s3 (70 ��� 75), s4 (70 ��� 80), s5 (82 ��� 92), s6 (85 ��� 96), r2 (62 ��� 74), r3 erect (60 ��� 67), r4 (75 ��� 80), r5 (75 ��� 82), r6 (70 ��� 79), J1 (65 ��� 75), J2 (68 ��� 77), J3 (67 ��� 82), J4 (70 ��� 75), Z1, Z2 (92 ��� 103), Z3 (100 ��� 112), Z4 (107 ��� 122), Z5 (93 ��� 110), S1 (77 ��� 95), S2, S3 (88 ��� 103), S4 (97 ��� 110), S5 (82 ��� 105). Lateral soft cuticle with seven pairs of smooth setae, including R1-R3 (48 ��� 62), R4-R6 (42 ��� 53) and ZV5 (47 ��� 55) posteriorly (Figs. 1A, B, C). Adult male. Dorsal shield 493 ��� 610 long, 315 ��� 342 wide at level of setae r3, 345 ��� 367 wide at r6 (n = 8); shield ornamentation, complement of pore-like structures and setation as in female; posterolateral setae attenuated as in female. Setae j1 (40 ��� 50), j2 (35 ��� 47), j3 (50 ��� 60), j4 (38 ��� 45), j5 (43 ��� 47), j6 (53 ��� 62), z1 (28 ��� 38), z2, z3 (43 ��� 55), z4 (52 ��� 62), z5 (40 ��� 45), z6 (48 ��� 55), s1 (47 ��� 57), s2-s4 (50 ��� 65), s5, s6 (60 ��� 75), r2 (50 ��� 60), r3 erect (38 ��� 55), r4, r5 (53 ��� 67), r6 (45 ��� 57), J1, J2 (43 ��� 55), J3 (50 ��� 62), J4 (40 ��� 47), J5 (5 ��� 9), Z1 (63 ��� 80), Z2 (65 ��� 92), Z3 (66 ��� 98), Z5 (61 ��� 77), S1 (54 ��� 72), S2 (55 ��� 77), S3, S4 (60 ��� 78), S5 (50 ��� 70), Z4 longest (80 ��� 102) of opisthonotal setae. Lateral soft cuticle with five or six pairs of R- setae (33 ��� 48) (Fig. 2A), R2 sometimes on small platelet (Fig. 2E). Idiosomatic venter ��� Adult female. Tritosternum base sometimes biarticulate, 38 ��� 43 long and 20 ��� 22 wide at its base, with paired laciniae fused for about 22 ��� 25 of entire length (122 ��� 135). Presternal region with circumscribed pair of weakly granulated, lineated areas, flanked anteriorly by a pair of weak or distinct narrow strips on either side of tritosternal base, and with setae st1 (33 ��� 37) inserted on this region, anterior to sternal shield. Sternal shield 127 ��� 135 long from anterior margin at transverse level with poroids iv1 to slightly concave posterior margin, 125 ��� 136 wide, weakly lineated along lateral margins, smooth medially; sternal setae st2, st3 of similar length (30 ��� 37). Sternal poroids iv3 alone on soft cuticle, setae st4 (28 ��� 35) on tiny metasternal platelets (13 ��� 15 x 8 ��� 11). Endopodal strips narrow between coxae III and IV. Epigynal shield sparsely reticulated, its broadly rounded hyaline anterior margin not overlapping posterior edge of sternal shield, its lateral margins slightly concave, narrowed anterior to setae st5 (38 ��� 47), and its posterolateral corners emarginated behind st5; shield 208 ��� 225 long, 118 ��� 137 at widest level at posterior rounded corners, 80 ��� 87 at narrowest width between legs IV. Opisthosomatic venter with two pairs of well separated metapodal platelets, primary pair elongate-oval, 33 ��� 40 long, 15 ��� 19 wide, secondary pair small, 10 ��� 16 by 5 ��� 7. Ventrianal shield inversely subtriangular with angulose or rounded anterior corners, with posterolateral margins on either side of anus divergent anteriorly; surface entirely reticulate, and punctate on either side of postanal seta; shield wider (293 ��� 337) than long (212 ��� 232), width at anal level (130 ��� 155); poroids ivo2 on shield anterior edge behind setae ZV1; gland pore gv3 on or off shield posterior to anal opening; shield with four pairs of ventral setae: JV1 (37 ��� 42), JV2 (43 ��� 50), JV3 (44 ��� 55), ZV2 (35 ��� 48); with paranal setae (21 ��� 27) shorter than postanal seta (43 ��� 65). Soft cuticle with five pairs of simple opisthogastric setae, ZV1 (35 ��� 44) anterior to ventrianal shield, ZV3 (33 ��� 43), ZV4 (37 ��� 44), JV5 (55 ��� 67) flanking the shield, and ZV5 aligned behind R6; three pairs of poroids on soft cuticle. Peritrematal-exopodal shield with lines extending from stigma into rounded posterior margin (Fig. 1C); peritreme extending to level between setae s1 and z1 (Fig. 1A). Spermathecal apparatus with cup-like calyx 28 ��� 29 long to embolus, 30 ��� 31 including it (Fig. 1F). Adult male. Tritosternum much as in female, but with base biarticulate, longer (20 ��� 32) than wide (15 ��� 19), and with laciniae fused for only about 10 ��� 15 of entire length (88 ��� 107). Presternal region faintly sclerotized, with a pair of distinct narrow strips on either side of tritosternal base, and with setae st1 inserted on this region on either side of genital opening. Sternitigenital shield length 225 ��� 267, narrowest widths 88 ��� 113 at level between coxae II, 77 ��� 95 between coxae III, 58 ��� 87 between coxae IV; shield slightly lineated along lateral margins and posteriorly between coxae IV, unornamented medially, with endopodal apices freely acuminate beside coxae IV; setae st1 (28 ��� 32) and st5 (28 ��� 37) longer, st2-st4 (17 ��� 26); poroids iv3 sometimes hardly discernible flanking faint pair of pits (Fig. 2A). Ventrianal shield well reticulated and with punctate fields in anal region as in female, its convex anterior margin sometimes excised near apices of exopodal strips behind coxae IV, and posterolateral margins slightly constricted in anal region; shield much wider (278 ��� 337) than long (170 ��� 228, including cribrum ca. 8 ��� 11 long, 58 ��� 65 wide); shield with six or usually seven pairs of opisthogastric setae, JV5 asymmetrically on or off shield (Figs. 1A, B); JV1 and ZV1 (23 ��� 32), JV2 (28 ��� 37), ZV2 (28 ��� 32), JV3 (28 ��� 42), ZV3 (23 ��� 30), JV5 (39 ��� 50); postanal seta smooth (39 ��� 47), about twice as long as paranal setae (15 ��� 22). Soft opisthogastric cuticle with two pairs of setae, ZV4 (26 ��� 30), and ZV5 (30 ��� 42) aligned behind R6, one pair of lyrifissures, ivp. Gnathosoma ��� Adult female. Gnathotectum anterior margin broadly truncate, with two lateral smooth prongs and serrate medial edge (Fig. 1D). Cheliceral shaft, excluding basal section, 160 ��� 173 long, with slender digits; dorsal seta blunt (13 ��� 14); fixed digit with tiny offset subapical tooth and row of 18-22 very fine teeth (Fig. 1G), pilus dentilus minute, at level between 4 th- 5 th apical teeth; movable digit (71 ��� 75) tridentate (Fig. 1H). Subcapitulum rows of deutosternal denticles progressively slightly wider posteriorly, each with numerous (ca 30-35) fine denticles, 6th and sometimes 7th widest and with ca 40-45 and 35-40 teeth, respectively (Fig. 1E); subcapitular setae hp1 (40 ��� 45) slightly shorter than hp3 (48 ��� 52) and capitular seta pc (43 ��� 52), hp3 much longer than hp2 (23 ��� 27). Palp length (195 ��� 212); apical margin of palptrochanter with a single spine anterolaterally; palptrochanter setae dissimilar in length, v1 48 ��� 52, v2 38 ��� 42. Adult male. Gnathotectum as in female (Fig. 2C). Cheliceral shaft, excluding short basal section 105 ��� 130 long; fixed digit with offset subapical tooth followed by minute pilus dentilis inserted on small angular protrusion and row of 11-16 small teeth (Fig. 2H); movable digit (40 ��� 45) unidentate; spermatodactyl 45 ��� 54 long, 8 ��� 9 wide at medial region, extending anteriorly 28 ��� 37 beyond apex of digit, with narrowed neck and small bifid head apically (Figs. 2F, G). Corniculi more widely spaced but not more arched than in female, 33 ��� 37 long, and flanked medially by pair of short (7-10) spiculate processes (Figs 2D, 11A); internal malae and labrum as in female. Subcapitulum with seven rows of deutosternal denticles much as in female, sixth and seventh rows slightly widest; hypostomatic setae hp1 (28 ��� 32) similarly as long as pc (25 ��� 37), hp3 usually longest (30 ��� 50), hp2 (12 ��� 20). Palpi (length 145 ��� 187) similar in structure, form of setae as in female; palptrochanter with inner seta longer (30 ��� 44) than outer seta (17 ��� 32) but not markedly whip-like. Legs ��� Adult female. Legs I (707 ��� 755) about as long as legs IV (715 ��� 750), and about as long as dorsal shield; other leg lengths (excluding pretarsi): II (565 ��� 600), III (587 ��� 615). Leg I pretarsus with biarticulate shaft (38 ��� 42 to base of claws), tarsus (187 ��� 205) about 1.6 longer than tibia; other length ratios, femur (135 ��� 147): genu (112 ��� 118): tibia (117 ��� 125) about 1.15: 0.95: 1.0. Legs II-IV with tarsus/tibia length ratios about 2.0-2.3. Chaetotactic formulae of leg segments as given for species-group. Tarsi II-IV with ventroapical process (12 ��� 17) bluntly pointed, smooth apically (Figs 13A, H), and with setae (d- 1) (23 ��� 30) slightly shorter than pretarsus to base of claws (28 ��� 37). Tarsi II-IV with paradactyli asymmetrical, anterior element increasingly extending beyond tip of claw from II to IV, its length 30 ��� 35, twice as long as claw (15) on leg IV (Fig. 14A). Tarsi II and III with setae (l- 1) and (v-1) symmetrical in form, pl-1 (32 ��� 35) and av-1 (36 ��� 47) slightly longer than al-1 (27 ��� 30) and pv-1 (26 ��� 37), respectively. Tarsus IV with distal seta av-1 (50 ��� 60) blunt, clearly larger, thicker than pv-1 (37 ��� 45); adjacent setae (l-1) symmetrically short (20 ��� 27), slender (Fig. 14A). Tarsi II-IV with dorsal seta md nearly reaching base of claws, and nearly as long as smooth attenuated setae (d-2) (67 ��� 80 on II, 85 ��� 100 on III, 95 ��� 110 on IV). Adult male. Legs I clearly longer than dorsal shield, not thicker than those of female; relative leg lengths, excluding pretarsi, similar to those of female, I 550 ��� 660, II 445 ��� 524, III 455 ��� 552, IV 575 ��� 675. Legs with segment length ratios as in female. Tarsus I with long pretarsus (25 ��� 37 to base of claws), claws 10 ��� 12 long. Tarsi II-IV with dorsoapical setal processes (d-1), ventroapical process and pretarsal structures formed as in female, pretarsus lengths 27 ��� 35 excluding claws (10) (Fig. 13B). Legs III-IV with tibial setae (v) and leg IV genual seta av dimorphically thicker, spinelike, less attenuate; length and form of other setae much as in female; seta pl-3 of basitarsus II smooth, attenuate (36 ��� 38) (Fig. 13B), but pl-3 of basitarsi III, IV (21 ��� 22) similar to pl- 2 in thicker, slightly shorter form than other three basitarsal setae. Type material ��� Except the final one, noted below, all specimens collected at the La Selva Biological Station, Heredia Province, Costa Rica (10��26��N, 84��01��W, elevation 50-150 m): HOLOTYPE: adult female, 8 July 1997, ex adult Cephaloleia bella Baly on Heliconia pogonantha leaves, coll. M. Paniagua. PARATYPES: 1 adult female, 1 adult male, with same data as holotype; 2 adult females, 1 adult male, 12 June 1997, ex coexistent adults of Cephaloleia belti Baly and C. ornatrix Donckier from Heliconia leaves, coll. ALAS; 3 adult females, 2 adult males, 15 April 1994, ex adult Chrysomelidae, no host plant data, M. Paniagua coll.; 1 adult female, 1 adult male, June 1997, ex adult Cephaloleia bella Baly, no host plant data, coll. D. Johnson; 1 adult female (gravid with larva), 1 adult male, June 1997, ex adult Cephaloleia belti Baly, no host plant data, coll. D. Johnson; 2 adult males, February 1994, associated with adult Chelobasis sp. on leaves of Heliconia pogonantha, coll. E.E. Lindquist & ALAS; 1 adult female, 2 adult males, 18 February 1994, ex adult Chrysomelidae, no host plant data, coll. ALAS; 3 adult females, 2 June 1997, with adult hispine beetles on Heliconiaceae leaves, coll. E.E. Lindquist; 2 adult females, June 1997, ex adult Cephaloleia vicina on Heliconia pogonantha, coll. not indicated; 1 adult male, February 1994, associated with adult Cephaloleia vicina on leaf of Heliconia pogonantha, coll. E.E. Lindquist & ALAS. Costa Rica, Osa Peninsula, Corcovado National Park (8��27.39��N, 83��25.45��W,elevation 10-500 m): 1 adult male, May 1997, ex adult Chelobasis bicolor Gray, no host plant data, coll. D. Johnson. Host identifications ��� Some of the identities of the associated beetles given above are in accord with current molecular studies, i.e., those as noted below for Lasioseius serripes. However, the following identities are discordant. Cephaloleia vicina does not occur at La Selva, and specimens identified as such from there are, in all cases checked by DNA barcodes, C. bella. Further, C. bella normally does not occur on Heliconia pogonantha; thousands of field diet records indicate it feeds only on H. imbricata (Garcia-Robledo et al., 2017). Etymology ��� The specific name is based on the chrysomelid subfamilial name Cassidinae, which includes the tribes Arescini and Imatidiini (or Cephaloleini) to which beetle associates of the genera Chelobasis and Cephaloleia belong, respectively. We have chosen this name for the species with the widest array of host associates and as the species-group name, anticipating that rolled-leaf beetles of other genera in these tribes will be found to live with these mites. Remarks ��� The male from Corcovado measured ca 12-19% smaller in dimensions than seven males measured from La Selva; however, in other descriptive aspects it fully resembles those from La Selva. Two females are asymmetrical in having humeral seta r3 inserted off the dorsal shield on one side, yet the aligned r2 and r4-r6 are on the shield. Two females show asymmetry in size and arrangement of the vertical cluster of setae j1, j2, z1; one male lacks one j2. Collection records of this mite species associated with beetles of three different species of Cephaloleia and one of Chelobasis leave uncertainty as to host preferences, particularly as beetles of these species are often coexistent (Garcia-Robledo 2010, Staines 2011, Schmitt & Frank 2013, Moraza & Lindquist 2015). Adults of Lasioseius cassidini are clearly the largest of the described species of this group. However, the data available are inadequate to test whether a mite/beetle size correlation exists, such as was hypothesized for larger mites being associated with larger beetles, for species of Hispiniphis and Makarovaia, which are more intimate subelytral associates of these same beetles (Moraza & Lindquist 2015, 2016). Although there are records of a few specimens of L. cassidini with relatively large beetles of Chelobasis, including one from the Osa Peninsula, they are based only on adult male mites, which may not present inherent preferences phoretically. Tentatively, subject to further studies, we conclude that, at La Selva, Lasioseius cassidini may be associated regularly with small beetles of the host specialist Cephaloleia bella and to some extent with the host generalist C. belti on Heliconia imbricata., Published as part of Moraza, Maria L. & Lindquist, Evert E., 2018, A new species-group with new species of the genus Lasioseius (Acari: Mesostigmata: Blattisociidae) associated with Neotropical hispine beetles in furled leaves ofHeliconia, pp. 62-98 in Acarologia 58 (1) on pages 69-74, DOI: 10.24349/acarologia/20184227, http://zenodo.org/record/4487537, {"references":["Garcia-Robledo C., Horvitz C. C., Kress J. W., Carvajal-Acosta N., Erwin T. L., Staines C. L. (2017) - Experimental assemblage of novel plant-herbivore interactions: ecological host shifts after 40 million years of isolation - Biotropica, 49 (6): 803 - 810. doi: 10.1111 / btp. 12464","Garcia-Robledo C., Horvitz C. C., Staines C. L. 2010 - Larval morphology and development, host plants, adult longevity, sexual dimorphism and notes on natural history in Cephaloleia rolled-leaf beetles (Coleoptera: Chrysomelidae: Hispinae) - Zootaxa, 2610: 50 - 68.","Staines C. L. 2011 - Hispines (Chrysomelidae, Cassidinae) of La Selva Biological Station, Costa Rica","Schmitt M., Frank M. 2013 - Notes on the ecology of rolled-leaf hispines (Chrysomelidae, Cassidinae) at La Gamba (Costa Rica) - ZooKeys, 332: 55 - 69. doi: 10.3897 / zookeys. 332.5215","Moraza M. L., Lindquist E. E. 2015 - Systematics and biology of mites associated with neotropical hispine beetles in unfurled leaves of Heliconia, with descriptions of two new genera of the family Melicharidae (Acari: Mesostigmata: Gamasina: Ascoides) - Zootaxa, 3931 (3): 301 - 351. doi: 10.11646 / zootaxa. 3931.3.1"]}

Lasioseius serripes n. sp. (Figures 3-7, 11B, 12A, B, G, 13 C-D, 14B) Zoobank: F1A4EE0C-0031-410D-B993-20CDC3D08D71 Diagnosis — Adult female: Sternal shield reticulated over entire surface, its posterolateral corners emarginated lateral to setae st3; poroids iv3 and setae st4 on soft cuticle; epigynal shield fully reticulated, its lateral margins parallel anterior to setae st5; ventrianal shield with posterolateral margins on either side of anus divergent, and with three pairs of opisthogastric setae, JV1 on soft cuticle; metapodal plates connected or coalesced on each side. Dorsal setae r6 usually on soft cuticle; Z5, S5 attenuated like adjacent setae. Adult male: Sternitigenital shield reticulated over entire surface. Leg II with dorsal setae dimorphically short, blunt on genu, tibia, tarsus. Tibiae III-IV with seta pv dimorphically hooked, blunt. Adult attributes: Gnathotectum anterior margin truncate, irregularly denticulate. Fixed chela with file of ca 20 teeth in female, ca 18 in male. Sternal setae st1 inserted on anterior edge of sternal shield. Tarsi II-IV with ventroapical process truncate-spiculate apically; tarsi II-III with seta av-1 slightly longer and thicker at base than pv-1 and both slightly longer than l- (1); tarsus IV with apical setae (v-1) symmetrically thicker than, and about twice as long as d- (1); pretarsal paradactyli small, symmetrical. Description Idiosomatic dorsum — Adult female: Dorsal shield 651 – 695 long, 337 – 385 wide at level of r3, 368 – 397 wide at level of r5 (n=9), lightly reticulated over entire surface, more faintly so behind setae Z4, where also faintly punctate. Shield with 37 pairs of setae, all acutely pointed (Figs. 3A, B), with few barbs, except J5 short (15 – 17), clearly barbed; J1-J4 about as long as longitudinal intervals between their bases, Z1-Z4 and S2-S4 longer than intervals between their bases; r3 erect, similarly as long as r2, r4: j1 (41 – 50), j2 (45 – 55), j3 (60 – 70), j4 (43 – 50), j5 (48 – 55), j6 (58 – 63), z1 (20 – 29), z2 (50 – 55), z3 (52 – 60), z4 (64 – 70), z5 (47 – 55), z6 (60 – 70), s1 (48 – 55), s2 (50 – 60), s3 (53 – 65), s4 (64 – 70), s5 (63 – 75), s6 (57 – 72), r2 (50 – 60), r3 (54 – 62), r4 (55 – 62), r5 (56 – 67), J1 (45 – 55), J2-J4 (48 – 62), Z1-Z3 (71 – 97), Z4 (87 – 102), Z5 (70 – 88), S1 (57 – 70), S2 (66 – 77), S3 (64 – 75), S4 (66 – 78), S5 (64 – 75). Lateral soft cuticle with eight pairs of smooth setae, including r6 (52 – 62), R1 (48 – 55), R2-R6 (44 – 52) and ZV5 (43 – 47) posteriorly. Adult male. Dorsal shield 557 – 587 long, 332 – 357 wide at level of setae r3, 325 – 353 wide at level of r6 (n = 5); shield ornamentation, complement of pore-like structures and setation as in female except slightly more expanded laterally so as to become united with peritrematal shields at level of setae s2 and bear marginal setae r6 and sometimes R1 (Fig. 5A). Dorsal shield with 37-38 pairs of setae, their form and relative lengths as in female, except bases of Z4, Z5, S4, S5 swollen sub-basally. Setae j1 (28 – 33), j2 (40 – 43), j3 (53 – 60), j4 (38 – 40), j5 (38 – 47), j6 (48 – 54), z1 (17 – 20), z2, z3 (45 – 50), z4 (55 – 60), z5 (38 – 44), z6 (50), s1 (46 – 48), s2 (52 – 56), s3 (50 – 61), s4 (58 – 65), s5 (58 – 65), s6 (55 – 58), r2-r4 (48 – 53), r5 (57 – 60), r6 (50 – 52), J1-J4 (40 – 47), J5 (9 – 12), Z1-Z3 (66 – 75), Z5 (61 – 72), S1 (50 – 52), S2 (55 – 57), S3-S5 (57 – 66), Z4 the longest opisthonotal setae (75 – 86). Lateral soft cuticle with usually six pairs of marginal setae (40 – 47). Form of peritrematal shields as in female. Idiosomatic venter — Adult female. Tritosternum base 28 – 33 long, 16 – 20 wide at its base, with paired laciniae fused along basal 15 – 22 of entire length (95 – 115). Presternal region with circumscribed pair of weakly lineated granulated areas adhered to anterior margin of sternal shield. Sternal shield 100 – 112 long, with irregularly concave posterior margin, 118 – 125 wide, well reticulated over entire surface, midsurface with narrower reticula; sternal setae of similar length, st1 (38 – 42), st2 (33 – 39), st3 (35 – 42). Sternal poroids iv3 and setae st4 (26 – 32) inserted on soft cuticle. Endopodal strips narrow between coxae III and IV. Epigynal shield well reticulated over entire surface, with narrower reticula anterior to insertion of st5 (36 – 42), its broadly rounded hyaline anterior margin not reaching posterior edge of sternal shield, its lateral margins nearly parallel anterior to setae st5, and its posterolateral corners slightly to clearly emarginated behind st5; shield 188 – 210 long, 95 – 107 at widest level at posterior angulate margin, 62 – 70 at narrowest width between legs IV. Opisthosomatic venter with two pairs of metapodal platelets connected or coalesced on each side, primary pair ovate, 25 – 30 long, 13 – 17 wide, secondary pair with greatest dimension transverse, 13 – 22; one adjacent set of small, narrow platelets near anterior margin of ventrianal shield, lateral to JV1. Ventrianal shield inversely subtriangular with rounded anterior corners, with posterolateral margins on either side of anus divergent anteriorly; surface strongly reticulate except faintly in anal region where also faintly punctate on either side of postanal seta; shield wider (253 – 295) than long (179 – 205), including cribrum (8 – 12); shield with three pairs of ventral setae and sometimes pores gv3: JV2 (33 – 40), JV3 (35 – 45), ZV2 (34 – 40); with paranal setae (20 – 27) shorter than postanal seta (49 – 62). Soft cuticle with six pairs of simple opisthogastric setae, JV1 and ZV1 (26 – 33) anterior to ventrianal shield, ZV3 (32 – 40), ZV4 (40 – 47), JV5 (55 – 65) flanking the shield, and ZV5 (43 – 47) aligned behind R6; three pairs of poroids (including ivp) and sometimes gland pore gv3 on soft opisthogastric cuticle. Peritrematal-exopodal shield with lines extending from stigma to irregularly rounded posterior margin (Fig. 4A); peritreme extending to level slightly anterior to insertion of setae s1 (Fig. 3A). Spermathecal apparatus with cup-like calyx ca. 30 – 35 long from atrium at base, 23 – 26 wide at distal margin (Fig. 3D). Adult male. Tritosternum much as in female, but with base biarticulate, longer (21 – 24) than wide (13 – 17), and laciniae (78 – 88) fused along basal 10 – 12 of entire length (Fig. 5C). Presternal region weakly granulated, with or without pair of weakly sclerotized transverse strips anteriorly. Sternitigenital shield length 245 – 283 from anterior margin of genital opening, widths 104 – 110 at level between coxae II, 92 – 100 between coxae III, and 58 – 71 between coxae IV; shield nearly fully reticulated, but bare near posterior margin where endopodal apices freely truncated beside coxae IV; shield medial region more weakly ornamented but punctate between setae iv1 and st3, and with poroids iv3 appearing as distinctively rounded pits; setae st1 on anterior margin of sclerotized region of shield; setae st1-st5 of dissimilar lengths as in female, st1-st3 (27 – 30), st4 shortest (20 – 23) and st5 longest (30 – 35). Ventrianal shield strongly reticulate as in female, its greatest width (282 – 305) larger than its length (178 – 220, including cribrum ca. 9 – 11 long, 75 – 86 wide); shield with seven pairs of opisthogastric setae; lateral margins irregular or eroded near where setae ZV4 on adjacent soft cuticle; setae JV1 (30 – 32), JV2 (37 – 42), JV3 (37 – 44), ZV1 (27 – 30), ZV2 (35 – 38), ZV3 (28 – 33), ZV4 (35 – 40), and JV5 the longest (50 – 59) inserted in bluntly angled lateral corners; postanal seta smooth (40 – 48), about twice as long as paranal setae (19 – 25). Soft opisthogastric cuticle with two pairs of setae, including ZV5 aligned behind R6, and one pair of lyrifissures, ivp (Fig. 5E). Gnathosoma — Adult female. Gnathotectum anterior margin broadly truncate, variably with two lateral, short, smooth prongs or with few denticles, sometimes with smaller denticles between lateral prongs (Fig. 3C). Cheliceral shaft, excluding basal section, 136 – 148 long, with slender digits (Figs 3E, F); dorsal seta blunt (8 – 11); fixed digit with tiny offset subapical tooth and row of 19-24 very fine teeth (Fig. 3E), pilus dentilus minute (1 – 2) at level of 4 th apical tooth; movable digit (62 – 72) tridentate. Subcapitulum rows of deutosternal denticles each with numerous (ca 25-40) fine denticles, those of most anterior row slightly coarser (ca 25-30), 2 nd- 5 th rows with ca 35-40 teeth, 6 th and sometimes 7 th rows slightly wider and slightly more denticulate (Fig. 4D); subcapitular setae hp1 (30 – 37) slightly shorter than capitular setae pc (36 – 42), hp3 slightly longest (40 – 47), much longer than hp2 (23 – 27). Salivary styli and corniculi as in figure 4C. Palp length (193 – 202); apical margin of palptrochanter without spines anterolaterally; palptrochanter setae dissimilar in length, v1 45 – 63 and v2 34 – 42 (Figs 4E, F). Adult male. Gnathotectum as in female (Fig. 5C). Cheliceral shaft, excluding short basal section, 110 – 120 long (Fig. 5F); fixed digit with offset subapical tooth followed by pilus dentilis behind hollow area and row of 10-16 small-sized teeth (Figs. 5F, G); movable digit (42 – 45) unidentate; spermatodactyl 40 – 57 long, 6 – 7 wide at base, slightly wider (8) at medial region, extending anteriorly ca 35 beyond apex of digit, with narrowed neck and small head apically (Figs 5F, H). Corniculi slightly more widely spaced and arched inwardly than in female, 25 – 30 long, and flanked medially by pair of short (7 – 8) spiculate processes (Figs 5I, 11B); internal malae and labrum as in female. Subcapitulum with seven or rarely eight rows of deutosternal denticles formed much as in female; hypostomatic setae hp1 not attenuated (21 – 27), slightly shorter than capitular setae pc (22 – 32), hp3 (30 – 37) slightly longest, hp2 shortest (15 – 20) (Fig. 5I). Palpi (length 173 – 188) similar in structure, form of setae as in female; palptrochanter with inner seta longer (38 – 40) than outer seta (20 – 23) but not markedly whip-like (Fig. 5I). Legs — Adult female. Legs I (669 – 715) nearly as long as legs IV (678 – 725), and slightly longer than dorsal shield; other leg lengths (excluding pretarsi): II 519 – 529, III 538 – 545 (Figs 6 A-E). Leg I pretarsus with biarticulated shaft (38 – 45 to base of claws), claws 15 – 17 long; tarsus (173 – 182) about 1.4-1.5 longer than tibia; other length ratios, femur (137 – 145): genu (110 – 117): tibia (112 – 124) about 1.2: 0.95: 1.0. Legs II-IV with tarsus/tibia length ratio about 2.0-2.2 (Figs 6C, D). Chaetotactic formulae of leg segments as given for species-group. Tarsi II-IV with ventroapical process (15 – 17) truncate-spiculate apically (Figs 13C, 14B), and with setae (d-1) (25 – 30) shorter than pretarsus to base of claws (33 – 44). Tarsi II-IV with paradactyli nearly symmetrical, weakly developed, anterior element barely extending beyond tip of claw (Figs 13G, 14B). Tarsi II (Fig. 13C), III with setae (l-1) (44 – 48) shorter than (v-l) (50 – 65); dorsal seta md (77 – 85) longest of tarsal setae, (d-2) barbed, less attenuated (50 – 62). Tarsus IV with setae av-1 (70 – 75) slightly longer than pv-1 (60 – 65), both nearly twice as long as similarly slender (l-1) (38 – 42) (Fig. 14B). Adult male. Legs I slightly thicker than those of female, and longer than dorsal shield; relative lengths similar to those of female, I 690 – 705, II 475 – 525, III 493 – 507, IV longest, 710 – 735. Legs with segment length ratios as in female. Tarsus I with long pretarsus (25 – 35 to base of claws), claws 15 long. Tarsi II-IV with spiculated form of ventroapical process (Fig. 13D), relative lengths of dorsoapical setal processes (d-1) (25 – 28) and pretarsal structures as in female; tarsi II-III with setae (v-1) (45 – 55) symmetrically longer than slender (l-1) (33 – 44), increasingly so on III; tarsus IV with av-1 (70 – 75) and pv-1 (55 – 67) asymmetrically longer than (l-1) (32 – 35) (Fig. 13D). Legs II with dorsal setae of genu, tibia, basitarsus and (d-2) of telotarsus (16, 23, 26, 27), distinctively shorter than in female and bluntly spine-like (Figs 12A, 13D); genu III and tibiae III-IV seta pv and tibia IV seta av dimorphically hooked, blunt (Fig. 12B). Length and form of other setae slightly thicker than in female. Deutonymph. None available to study for comparative purposes. Protonymph. General description, based on one specimen, without measurements, for speciesgroup perspective. Dorsal idiosoma (Fig. 7C) with setae generally nearly aciculate, but with some setae acquiring slightly tricarinate or barbed form as in adult. Podonotal region holotrichous, with 15 pairs of setae (five pairs added to larval complement, including j2, s5 amidst 11 pairs on podonotal shield and r2, r3, r5 along with larval s6 on lateral soft cuticle); j2 inserted in nearly transverse alignment with j1. Opisthonotal region holotrichous, with 15 pairs of setae (five pairs added to larval complement, including J1, Z1, Z2, S2, R1, all on interscutal soft cuticle; pygidial shield expanded so as to include eight pairs of setae, including J3, Z5, S4, S5 (all larval setae). Ventral idiosoma (Fig. 7D) with all setae aciculate, with no postlarval podosomal additions except st5. Sternal poroids iv1, iv2 present, iv3 absent, and larval funicules suppressed. Anal shield oval, with paranal setae similar in length or slightly shorter than postanal seta; rudiments of euanal setae suppressed. Opisthogastric soft cuticle with four or five pairs of setae, with asymmetrical addition of ZV5, which usually deutonymphal; JV5 relatively larger than in larva, and inserted more posteriorly, in transverse alignment with postanal seta. Gnathosoma — Fixed cheliceral digit with even row of 12-13 teeth; movable digit tridentate. Gnathotectum similar in form to adult. Subcapitulum with four pairs of setae, hypostomatic h1-h3 and capitular pc. Palpus with normal protonymphal complement of setae, including one on trochanter (Lindquist & Evans 1965); palptarsus with hyaline flap developed beside apotele as in adult. Legs — Legs I-IV with claws, pretarsi and shape of setae similar to adult female; tarsi II-IV with ventroapical process similar in form, serrated apically, as adult. Legs I, II, III, IV with normal, holotrichous protonymphal chaetotaxy (Lindquist & Evans 1965), except genu IV with accelerated presence of usually deutonymphal seta pl-1, and femur IV with accelerated alveolar indication of seta v-1 near basal lyriform fissure: coxae 2, 2, 2, 1; trochanters 4, 4, 4, 4; femora 10 (2 2/1 2/2 1), 8 (1 2/1 2/1 1), 5 (1 2/1 1/0 0), 4 (1 2/0 1/0 0); genua 8 (1 2/1 2/1 1), 6 (1 2/0 2/0 1), 6 (1 2/0 2/0 1), 6 (1 2/0 2/0 1); tibiae 8 (1 2/1 2/1 1), 7 (1 1/1 2/1 1), 7 (1 1/1 2/1 1), 7 (1 1/1 2/1 1); tarsi II-IV each with 17 (md added to larval complement). Tarsus IV with seta av-1 distinctively erect, longer and slightly thicker than all other tarsal setae (Fig. 12G), in contrast to its form on tarsi II, III of protonymph, and on tarsus IV of adult. Larva. General description, based on two unborn specimens, without measurements, for species-group perspective. Dorsal idiosoma (Fig. 7A) with all setae aciculate. Podonotal shield holotrichous with typical nine pairs of setae j (1, j3-j6, z2, z4, z5, s4); setae s6 on soft cuticle behind posterolateral margin of shield. Opisthonotal region holotrichous with 10 pairs of setae (J2-J5, Z3-Z5, S3-S5); pygidial shield margins mostly indiscernible on specimens at hand, but its surface with typical pair of arcuate lines near insertions of setae J4, and with typical complement of four pairs of setae, including tiny J4, J5 and prominent Z3, Z4. Soft interscutal cuticle with two pairs of mesonotal scutellae and typical six pairs of setae, including moderately long J2, J3, S3, S5, prominent Z5 (in ventrolateral position), and shorter S4. Ventral idiosoma (Fig. 7B) with all setae aciculate. Sternal shield outline not distinguishable; setae st1-st3 similarly short, st4, st5 absent, but the typical pair of subcutaneous structures (‘funicules’ sensu Athias-Henriot 1971a) present posteromediad bases of legs III. Anal shield margins indistinguishable; anal valves with rudimentary pair of euanal setae; paranal setae slightly longer than postanal seta. Opisthogastric soft cuticle with typical four pairs of similarly short setae (JV1, JV2, JV5, ZV2). Gnathosoma — Fixed cheliceral digit with even row of 10-11 teeth; movable digit bidentate. Gnathotectum triramous, branches shorter than in adult. Subcapitulum with two pairs of hypostomatic setae, h1 longer than h2. Palpus with normal larval complement of setae (Lindquist & Evans 1965); palptarsus with hyaline flap not evident beside apotele. Legs — Legs I-III with claws, pretarsi and shape of setae similar to adult female; tarsi II, III with ventroapical process similar in form, serrated apically, as adult, and with setae d- (1) more strongly developed, slightl

Lasioseius duobtusisetis n. sp. (Figures 10, 12 E-F, 13I, 14C) Zoobank: F8B79273-9C72-4212-816C-8BC080C933CE Diagnosis. Adult female: Sternal shield weakly lineate along lateral margins, smooth medially, its posterolateral corners not emarginated; poroids iv3 and setae st4 together on slender metasternal platelets; epigynal shield partially reticulated laterally, its lateral margins gently concave, narrowed anterior to setae st5; ventrianal shield with posterolateral margins on either side of anus nearly parallel, and with four pairs of opisthogastric setae, JV1 on shield; metapodal plates well separated on each side. Dorsal setae r6 on soft cuticle; setae Z5, S5 thickened, spinelike with blunt tips, Z4, S4 somewhat thickened but attenuated. Adult attributes: Gnathotectum anterior margin triramous, denticulate. Fixed chela with file of ca 20 teeth in female. Sternal setae st1 inserted on anterior edge of sternal shield. Tarsi II-IV with ventral apical process bluntly pointed, smooth apically; tarsi II-III with setae (v- 1), (l-1) symmetrically stout; tarsus IV with apical seta av-1 longer, stouter than pv-1, (l-1), and pretarsus with anterior paradactylus longer than posterior one. Description Idiosomatic dorsum ��� Adult female. Dorsal shield 540 ��� 595 long, 293 ��� 319 wide at level of r3, 320 ��� 345 at level of r5 (n=4), lightly reticulated over most of surface, but nearly smooth in dorsal hexagonal area (delimited by setae j5-z5-j6), not punctate behind level of setae Z4, but finely transversely striate between Z5 behind J5; shield with 37 pairs of setae, with lengths of most podonotal setae and J1-J3 not reaching bases of following setae; Z5, S5 thickened, spinelike with blunt tips, Z4, S4 somewhat thickened but attenuated (Figs. 10A, B): j1 (34 ��� 42), j2 (31 ��� 40), j3 (39 ��� 43), j4 (28 ��� 34), j5 (30 ��� 37), j6 (38 ��� 50), z1 (24 ��� 27), z2 (33 ��� 40), z3 (40 ��� 43), z4 (40 ��� 50), z5 (30 ��� 38), z6 (37 ��� 40), s1 (29 ��� 35), s2 (38 ��� 50), s3 (43 ��� 50), s4 (40 ��� 51), s5 (45 ��� 52), s6 (50 ��� 57), r2 (40 ��� 50), r3 erect (38 ��� 42), r4 (44 ��� 57), r5 (47 ��� 57), J1 (33 ��� 38), J2 (33 ��� 42), J3 (43 ��� 52), J4 (40 ��� 57), J5 minute and barbed (7 ��� 9), Z1 (53 ��� 62), Z2 (58 ��� 62), Z3 (62 ��� 70), Z4 (80 ��� 87), Z5 blunt (85 ��� 103), S1 (48 ��� 59), S2 (55 ��� 67), S3 (58 ��� 69), S4 (63 ��� 75), S5 blunt (75 ��� 90). Lateral soft cuticle with eight pairs of smooth setae, including r6 (45 ��� 60), R 1 (35 ��� 52), R 2- R 6 shorter (28 ��� 42) and ZV5 posteriorly (40) (Fig. 10C). Idiosomatic venter ��� Adult female. Tritosternum base 26 ��� 35 long, 15 ��� 16 wide at its base, with paired laciniae fused for about 10 ��� 17 of their total length (112 ��� 118). Presternal region with circumscribed pair of weakly granulated areas with jigsawed lineation, sometimes flanked anteriorly by a pair of narrow slightly sclerotized strips. Sternal shield 115 ��� 127 long, with slightly concave posterior margin, 98 ��� 109 wide, its surface nearly smooth medially, weakly lineate along lateral margins; sternal setae st1-st3 attenuate, of similar length (21 ��� 25) (Fig. 10C). Sternal poroids iv3 and setae st4 (20 ��� 22) inserted together on small metasternal platelets (21 ��� 26 long). Endopodal strips narrow between coxae III and IV. Epigynal shield with lightly reticulated lateral areas separated by smooth area medially, its narrowly rounded hyaline anterior margin reaching or slightly overlapping posterior edge of sternal shield, its lateral margins slightly concave, narrowed anterior to setae st5 (20 ��� 24), and its posterolateral corners slightly emarginated behind st5: shield 178 ��� 197 long, 82 ��� 102 at widest level at posterior angulate corners, 70 ��� 75 at level of st5, and 54 ��� 62 at narrowest width between legs IV. Opisthosomatic venter with two pairs of well separated metapodal platelets, primary pair oval, 30 ��� 40 long, 15 ��� 19 wide, secondary pair small, greatest dimension transverse, 18 ��� 25. Ventrianal shield inversely subtriangular with rounded anterior corners, with posterolateral margins on either side of anus nearly parallel; surface entirely sculptured, with primarily transverse lineation changing to reticulation behind setae JV2-ZV2, except anal region conspicuously punctate on either side of postanal seta; shield wider (216 ��� 235) than long, including cribrum (176 ��� 197), 113 ��� 132 wide at anal region; shield with four pairs of ventral setae and glandular pores gv3: JV1 (19 ��� 22), JV2 (22 ��� 25), JV3 (23 ��� 27), ZV2 (23 ��� 29); with paranal setae (17 ��� 20) shorter than postanal seta (36 ��� 45). Soft cuticle with five pairs of simple opisthogastric setae, ZV1 (22 ��� 24) anterior to ventrianal shield, ZV3 (22 ��� 26), ZV4 (25 ��� 33), JV5 markedly thicker (53 ��� 57) flanking the shield, and ZV5 somewhat aligned behind R 6; four pairs of poroids (including ivp). Peritrematal-exopodal shield lineate posteriorly behind leg IV; peritreme extending to level between insertions of setae s1 and z1 (Fig. 10C). Spermathecal apparatus with cup-like calyx 40 long including embolus and 22 wide at distal margin (Fig. 10 F). Gnathosoma ��� Gnathotectum anterior margin triramous, with three short denticulate prongs, lateral margin slightly serrate (Fig. 10D). Cheliceral shaft, excluding basal section, 161 ��� 182 long, with slender digits; dorsal seta blunt (10); fixed digit with tiny, slightly offset subapical tooth and row of 20-28 very fine teeth (Fig. 10G), pilus dentilus minute (3) at level of 4 th or 5 th apical tooth; movable digit (70 ��� 80) tridentate. Subcapitulum rows of deutosternal denticles each with numerous (ca 20-30) fine denticles, 6 th and sometimes 7 th slightly widest and with ca 35 and 30 teeth, respectively; subcapitular setae hp1 (41 ��� 45) nearly as long as capitular setae pc (48 ��� 50), hp3 shorter (30 ��� 32), but longer than hp2 (13 ��� 17) (Fig. 10E). Palp length (160 ��� 165); apical margin of palptrochanter with a single spine anterolaterally; palptrochanter setae dissimilar in length, v1 42 ��� 45 and v2 27 ��� 36 (Fig. 10E). Legs ��� Legs I (503 ��� 562) slightly longer than legs IV (488 ��� 546), and shorter than dorsal shield; other leg lengths (excluding pretarsi): II 390 ��� 445, III 380 ��� 442. Leg I pretarsus with biarticulated shaft (25 ��� 27 to base of claws), claws 15 long; tarsus (142 ��� 167) about 1.8 longer than tibia; other length ratios, femur (88 ��� 105): genu (73 ��� 82): tibia (80 ��� 95), about 1.1: 0.9: 1.0. Legs II-III with tarsus/tibia length ratio about 1.8-2.2, leg IV with that ratio about 1.8-1.9. Chaetotactic formulae of leg segments as given for species-group. Tarsi II-IV with ventroapical process (8 ��� 10) bluntly pointed, smooth apically (Figs 13I, 14C), and with setae (d-1) (13 ��� 18) nearly as long as pretarsus to base of claws (15 ��� 18). Tarsi II-III with paradactyli reaching tips of tarsal claws, but tarsus IV with anterior paradactylus about 1.5 longer (Fig. 14C). Tarsi II and III with setae (l- 1) and (v-1) symmetrically stout, pl-1 (18) and pv-1 (18 ��� 25) slightly longer than al-1 (15) and av-1 (17 ��� 20), respectively; md (43 ��� 50) reaching bases of claws; setae (d-2) symmetrically attenuate (45 ��� 53). Tarsus IV with apical seta av-1 longer (28 ��� 45), stouter than pv-1 (16 ��� 23); setae (l-1) symmetrically similar in length (13 ��� 15), md (55 ��� 61) and (d-2) (60 ��� 72) attenuate (Fig. 14C). ADULT MALE. Unknown. Type material ��� All specimens collected at the La Selva Biological Station, Heredia Province, Costa Rica (10��26��N, 84��01��W, elevation 50-150 m): HOLOTYPE: adult female, (CRI 002, 739028), 8 July 1997, ex adult Chelobasis perplexa Baly, no host plant data, coll. M Paniagua. PARATYPES: 1 adult female, February 1994, associated with adult Chelobasis sp. on leaf of Heliconia pogonantha, coll. E.E. Lindquist & ALAS; 1 adult female, 15 April 1994, ex adult Chrysomelidae, no host plant data, coll. M. Paniagua; 1 adult female, 12 June 1997, ex adult Cephaloleia aequilata Uhmann (misidentification, see below) on leaf of Calathea, coll. ALAS; 1 adult female, 12 June 1997, ex adult Cephaloleia erichsonii Baly on leaf of Calathea, coll. ALAS. Host identifications ��� With one exception, the identities of the associated beetles given above are in accord with current molecular studies of their field diet records, which indicate that one haplotype of Chelobasis perplexa sensu stricto is on Heliconia pogonantha. Cephalolaeia erichsonii is a host generalist. Cephaloleia aequilata does not occur at La Selva, and specimens identified as such from there are, in all cases checked by DNA barcodes, C. dilaticollis Baly, which is a host generalist, often on Heliconia, Calathea and other host plants (Carlos Garcia-Robledo, personal communications 2017; Garcia-Robledo et al. 2017). Etymology ��� The specific name is a Latinized adjectival combination of ���duo-obtusi-setis���, referring to two pairs of characteristically blunt-tipped setae Z5 and S5 on the dorsal shield. Remarks ��� One female asymmetrically lacks one of setae s2. One female asymmetrically has seta JV4 inserted on the ventrianal shield; otherwise, this setal pair is symmetrically absent, characteristically in all species described of this group. One female at hand, from under elytra of Cephaloleia erichsonii, Corcovado National Park, May 1997, coll. D. Johnson, has been tentatively identified as Lasioseius duobtusisetis, but is excluded from the type material. It differs primarily from other females by having lengths of most podonotal setae and J1-J3 longer, reaching beyond bases of following setae; yet setae Z5, S5 are thickened, spinelike with blunt tips, and Z4, S4 somewhat thickened but attenuated, which are distinctive of this species. Two aspects of the sparse collection data are notable, with two records of association with beetles of the genus Chelobasis, and two other records with beetles of two species of Cephaloleia from leaves of a different genus of host plant, Calathea. Tentatively, subject to further studies, we conclude that, at La Selva, Lasioseius duobtusisetis is associated regularly with generalist forms of Chelobasis perplexa and Cephaloleia erichsonii on leaves of various species of Heliconia (and C. erichsonii also on various species of Calathea and Goeppertia), and also to some extent with another generalist form, Cephaloleia dilaticollis on a similar variety of host plant genera (Staines & Garcia-Robledo 2014, Garcia-Robledo et al. 2017)., Published as part of Moraza, Maria L. & Lindquist, Evert E., 2018, A new species-group with new species of the genus Lasioseius (Acari: Mesostigmata: Blattisociidae) associated with Neotropical hispine beetles in furled leaves ofHeliconia, pp. 62-98 in Acarologia 58 (1) on pages 88-93, DOI: 10.24349/acarologia/20184227, http://zenodo.org/record/4487537, {"references":["Garcia-Robledo C., Horvitz C. C., Kress J. W., Carvajal-Acosta N., Erwin T. L., Staines C. L. (2017) - Experimental assemblage of novel plant-herbivore interactions: ecological host shifts after 40 million years of isolation - Biotropica, 49 (6): 803 - 810. doi: 10.1111 / btp. 12464","Staines C. L., Garcia-Robledo C. 2014 - The genus Cephaloleia Chevrolat, 1836 (Coleoptera: Chrysomelidae: Cassidinae) - ZooKeys, 436: 1 - 355. doi: 10.3897 / zookeys. 436.5766"]}

Lasioseius serripes n. sp. (Figures 3-7, 11B, 12A, B, G, 13 C-D, 14B) Zoobank: F1A4EE0C-0031-410D-B993-20CDC3D08D71 Diagnosis ��� Adult female: Sternal shield reticulated over entire surface, its posterolateral corners emarginated lateral to setae st3; poroids iv3 and setae st4 on soft cuticle; epigynal shield fully reticulated, its lateral margins parallel anterior to setae st5; ventrianal shield with posterolateral margins on either side of anus divergent, and with three pairs of opisthogastric setae, JV1 on soft cuticle; metapodal plates connected or coalesced on each side. Dorsal setae r6 usually on soft cuticle; Z5, S5 attenuated like adjacent setae. Adult male: Sternitigenital shield reticulated over entire surface. Leg II with dorsal setae dimorphically short, blunt on genu, tibia, tarsus. Tibiae III-IV with seta pv dimorphically hooked, blunt. Adult attributes: Gnathotectum anterior margin truncate, irregularly denticulate. Fixed chela with file of ca 20 teeth in female, ca 18 in male. Sternal setae st1 inserted on anterior edge of sternal shield. Tarsi II-IV with ventroapical process truncate-spiculate apically; tarsi II-III with seta av-1 slightly longer and thicker at base than pv-1 and both slightly longer than l- (1); tarsus IV with apical setae (v-1) symmetrically thicker than, and about twice as long as d- (1); pretarsal paradactyli small, symmetrical. Description Idiosomatic dorsum ��� Adult female: Dorsal shield 651 ��� 695 long, 337 ��� 385 wide at level of r3, 368 ��� 397 wide at level of r5 (n=9), lightly reticulated over entire surface, more faintly so behind setae Z4, where also faintly punctate. Shield with 37 pairs of setae, all acutely pointed (Figs. 3A, B), with few barbs, except J5 short (15 ��� 17), clearly barbed; J1-J4 about as long as longitudinal intervals between their bases, Z1-Z4 and S2-S4 longer than intervals between their bases; r3 erect, similarly as long as r2, r4: j1 (41 ��� 50), j2 (45 ��� 55), j3 (60 ��� 70), j4 (43 ��� 50), j5 (48 ��� 55), j6 (58 ��� 63), z1 (20 ��� 29), z2 (50 ��� 55), z3 (52 ��� 60), z4 (64 ��� 70), z5 (47 ��� 55), z6 (60 ��� 70), s1 (48 ��� 55), s2 (50 ��� 60), s3 (53 ��� 65), s4 (64 ��� 70), s5 (63 ��� 75), s6 (57 ��� 72), r2 (50 ��� 60), r3 (54 ��� 62), r4 (55 ��� 62), r5 (56 ��� 67), J1 (45 ��� 55), J2-J4 (48 ��� 62), Z1-Z3 (71 ��� 97), Z4 (87 ��� 102), Z5 (70 ��� 88), S1 (57 ��� 70), S2 (66 ��� 77), S3 (64 ��� 75), S4 (66 ��� 78), S5 (64 ��� 75). Lateral soft cuticle with eight pairs of smooth setae, including r6 (52 ��� 62), R1 (48 ��� 55), R2-R6 (44 ��� 52) and ZV5 (43 ��� 47) posteriorly. Adult male. Dorsal shield 557 ��� 587 long, 332 ��� 357 wide at level of setae r3, 325 ��� 353 wide at level of r6 (n = 5); shield ornamentation, complement of pore-like structures and setation as in female except slightly more expanded laterally so as to become united with peritrematal shields at level of setae s2 and bear marginal setae r6 and sometimes R1 (Fig. 5A). Dorsal shield with 37-38 pairs of setae, their form and relative lengths as in female, except bases of Z4, Z5, S4, S5 swollen sub-basally. Setae j1 (28 ��� 33), j2 (40 ��� 43), j3 (53 ��� 60), j4 (38 ��� 40), j5 (38 ��� 47), j6 (48 ��� 54), z1 (17 ��� 20), z2, z3 (45 ��� 50), z4 (55 ��� 60), z5 (38 ��� 44), z6 (50), s1 (46 ��� 48), s2 (52 ��� 56), s3 (50 ��� 61), s4 (58 ��� 65), s5 (58 ��� 65), s6 (55 ��� 58), r2-r4 (48 ��� 53), r5 (57 ��� 60), r6 (50 ��� 52), J1-J4 (40 ��� 47), J5 (9 ��� 12), Z1-Z3 (66 ��� 75), Z5 (61 ��� 72), S1 (50 ��� 52), S2 (55 ��� 57), S3-S5 (57 ��� 66), Z4 the longest opisthonotal setae (75 ��� 86). Lateral soft cuticle with usually six pairs of marginal setae (40 ��� 47). Form of peritrematal shields as in female. Idiosomatic venter ��� Adult female. Tritosternum base 28 ��� 33 long, 16 ��� 20 wide at its base, with paired laciniae fused along basal 15 ��� 22 of entire length (95 ��� 115). Presternal region with circumscribed pair of weakly lineated granulated areas adhered to anterior margin of sternal shield. Sternal shield 100 ��� 112 long, with irregularly concave posterior margin, 118 ��� 125 wide, well reticulated over entire surface, midsurface with narrower reticula; sternal setae of similar length, st1 (38 ��� 42), st2 (33 ��� 39), st3 (35 ��� 42). Sternal poroids iv3 and setae st4 (26 ��� 32) inserted on soft cuticle. Endopodal strips narrow between coxae III and IV. Epigynal shield well reticulated over entire surface, with narrower reticula anterior to insertion of st5 (36 ��� 42), its broadly rounded hyaline anterior margin not reaching posterior edge of sternal shield, its lateral margins nearly parallel anterior to setae st5, and its posterolateral corners slightly to clearly emarginated behind st5; shield 188 ��� 210 long, 95 ��� 107 at widest level at posterior angulate margin, 62 ��� 70 at narrowest width between legs IV. Opisthosomatic venter with two pairs of metapodal platelets connected or coalesced on each side, primary pair ovate, 25 ��� 30 long, 13 ��� 17 wide, secondary pair with greatest dimension transverse, 13 ��� 22; one adjacent set of small, narrow platelets near anterior margin of ventrianal shield, lateral to JV1. Ventrianal shield inversely subtriangular with rounded anterior corners, with posterolateral margins on either side of anus divergent anteriorly; surface strongly reticulate except faintly in anal region where also faintly punctate on either side of postanal seta; shield wider (253 ��� 295) than long (179 ��� 205), including cribrum (8 ��� 12); shield with three pairs of ventral setae and sometimes pores gv3: JV2 (33 ��� 40), JV3 (35 ��� 45), ZV2 (34 ��� 40); with paranal setae (20 ��� 27) shorter than postanal seta (49 ��� 62). Soft cuticle with six pairs of simple opisthogastric setae, JV1 and ZV1 (26 ��� 33) anterior to ventrianal shield, ZV3 (32 ��� 40), ZV4 (40 ��� 47), JV5 (55 ��� 65) flanking the shield, and ZV5 (43 ��� 47) aligned behind R6; three pairs of poroids (including ivp) and sometimes gland pore gv3 on soft opisthogastric cuticle. Peritrematal-exopodal shield with lines extending from stigma to irregularly rounded posterior margin (Fig. 4A); peritreme extending to level slightly anterior to insertion of setae s1 (Fig. 3A). Spermathecal apparatus with cup-like calyx ca. 30 ��� 35 long from atrium at base, 23 ��� 26 wide at distal margin (Fig. 3D). Adult male. Tritosternum much as in female, but with base biarticulate, longer (21 ��� 24) than wide (13 ��� 17), and laciniae (78 ��� 88) fused along basal 10 ��� 12 of entire length (Fig. 5C). Presternal region weakly granulated, with or without pair of weakly sclerotized transverse strips anteriorly. Sternitigenital shield length 245 ��� 283 from anterior margin of genital opening, widths 104 ��� 110 at level between coxae II, 92 ��� 100 between coxae III, and 58 ��� 71 between coxae IV; shield nearly fully reticulated, but bare near posterior margin where endopodal apices freely truncated beside coxae IV; shield medial region more weakly ornamented but punctate between setae iv1 and st3, and with poroids iv3 appearing as distinctively rounded pits; setae st1 on anterior margin of sclerotized region of shield; setae st1-st5 of dissimilar lengths as in female, st1-st3 (27 ��� 30), st4 shortest (20 ��� 23) and st5 longest (30 ��� 35). Ventrianal shield strongly reticulate as in female, its greatest width (282 ��� 305) larger than its length (178 ��� 220, including cribrum ca. 9 ��� 11 long, 75 ��� 86 wide); shield with seven pairs of opisthogastric setae; lateral margins irregular or eroded near where setae ZV4 on adjacent soft cuticle; setae JV1 (30 ��� 32), JV2 (37 ��� 42), JV3 (37 ��� 44), ZV1 (27 ��� 30), ZV2 (35 ��� 38), ZV3 (28 ��� 33), ZV4 (35 ��� 40), and JV5 the longest (50 ��� 59) inserted in bluntly angled lateral corners; postanal seta smooth (40 ��� 48), about twice as long as paranal setae (19 ��� 25). Soft opisthogastric cuticle with two pairs of setae, including ZV5 aligned behind R6, and one pair of lyrifissures, ivp (Fig. 5E). Gnathosoma ��� Adult female. Gnathotectum anterior margin broadly truncate, variably with two lateral, short, smooth prongs or with few denticles, sometimes with smaller denticles between lateral prongs (Fig. 3C). Cheliceral shaft, excluding basal section, 136 ��� 148 long, with slender digits (Figs 3E, F); dorsal seta blunt (8 ��� 11); fixed digit with tiny offset subapical tooth and row of 19-24 very fine teeth (Fig. 3E), pilus dentilus minute (1 ��� 2) at level of 4 th apical tooth; movable digit (62 ��� 72) tridentate. Subcapitulum rows of deutosternal denticles each with numerous (ca 25-40) fine denticles, those of most anterior row slightly coarser (ca 25-30), 2 nd- 5 th rows with ca 35-40 teeth, 6 th and sometimes 7 th rows slightly wider and slightly more denticulate (Fig. 4D); subcapitular setae hp1 (30 ��� 37) slightly shorter than capitular setae pc (36 ��� 42), hp3 slightly longest (40 ��� 47), much longer than hp2 (23 ��� 27). Salivary styli and corniculi as in figure 4C. Palp length (193 ��� 202); apical margin of palptrochanter without spines anterolaterally; palptrochanter setae dissimilar in length, v1 45 ��� 63 and v2 34 ��� 42 (Figs 4E, F). Adult male. Gnathotectum as in female (Fig. 5C). Cheliceral shaft, excluding short basal section, 110 ��� 120 long (Fig. 5F); fixed digit with offset subapical tooth followed by pilus dentilis behind hollow area and row of 10-16 small-sized teeth (Figs. 5F, G); movable digit (42 ��� 45) unidentate; spermatodactyl 40 ��� 57 long, 6 ��� 7 wide at base, slightly wider (8) at medial region, extending anteriorly ca 35 beyond apex of digit, with narrowed neck and small head apically (Figs 5F, H). Corniculi slightly more widely spaced and arched inwardly than in female, 25 ��� 30 long, and flanked medially by pair of short (7 ��� 8) spiculate processes (Figs 5I, 11B); internal malae and labrum as in female. Subcapitulum with seven or rarely eight rows of deutosternal denticles formed much as in female; hypostomatic setae hp1 not attenuated (21 ��� 27), slightly shorter than capitular setae pc (22 ��� 32), hp3 (30 ��� 37) slightly longest, hp2 shortest (15 ��� 20) (Fig. 5I). Palpi (length 173 ��� 188) similar in structure, form of setae as in female; palptrochanter with inner seta longer (38 ��� 40) than outer seta (20 ��� 23) but not markedly whip-like (Fig. 5I). Legs ��� Adult female. Legs I (669 ��� 715) nearly as long as legs IV (678 ��� 725), and slightly longer than dorsal shield; other leg lengths (excluding pretarsi): II 519 ��� 529, III 538 ��� 545 (Figs 6 A-E). Leg I pretarsus with biarticulated shaft (38 ��� 45 to base of claws), claws 15 ��� 17 long; tarsus (173 ��� 182) about 1.4-1.5 longer than tibia; other length ratios, femur (137 ��� 145): genu (110 ��� 117): tibia (112 ��� 124) about 1.2: 0.95: 1.0. Legs II-IV with tarsus/tibia length ratio about 2.0-2.2 (Figs 6C, D). Chaetotactic formulae of leg segments as given for species-group. Tarsi II-IV with ventroapical process (15 ��� 17) truncate-spiculate apically (Figs 13C, 14B), and with setae (d-1) (25 ��� 30) shorter than pretarsus to base of claws (33 ��� 44). Tarsi II-IV with paradactyli nearly symmetrical, weakly developed, anterior element barely extending beyond tip of claw (Figs 13G, 14B). Tarsi II (Fig. 13C), III with setae (l-1) (44 ��� 48) shorter than (v-l) (50 ��� 65); dorsal seta md (77 ��� 85) longest of tarsal setae, (d-2) barbed, less attenuated (50 ��� 62). Tarsus IV with setae av-1 (70 ��� 75) slightly longer than pv-1 (60 ��� 65), both nearly twice as long as similarly slender (l-1) (38 ��� 42) (Fig. 14B). Adult male. Legs I slightly thicker than those of female, and longer than dorsal shield; relative lengths similar to those of female, I 690 ��� 705, II 475 ��� 525, III 493 ��� 507, IV longest, 710 ��� 735. Legs with segment length ratios as in female. Tarsus I with long pretarsus (25 ��� 35 to base of claws), claws 15 long. Tarsi II-IV with spiculated form of ventroapical process (Fig. 13D), relative lengths of dorsoapical setal processes (d-1) (25 ��� 28) and pretarsal structures as in female; tarsi II-III with setae (v-1) (45 ��� 55) symmetrically longer than slender (l-1) (33 ��� 44), increasingly so on III; tarsus IV with av-1 (70 ��� 75) and pv-1 (55 ��� 67) asymmetrically longer than (l-1) (32 ��� 35) (Fig. 13D). Legs II with dorsal setae of genu, tibia, basitarsus and (d-2) of telotarsus (16, 23, 26, 27), distinctively shorter than in female and bluntly spine-like (Figs 12A, 13D); genu III and tibiae III-IV seta pv and tibia IV seta av dimorphically hooked, blunt (Fig. 12B). Length and form of other setae slightly thicker than in female. Deutonymph. None available to study for comparative purposes. Protonymph. General description, based on one specimen, without measurements, for speciesgroup perspective. Dorsal idiosoma (Fig. 7C) with setae generally nearly aciculate, but with some setae acquiring slightly tricarinate or barbed form as in adult. Podonotal region holotrichous, with 15 pairs of setae (five pairs added to larval complement, including j2, s5 amidst 11 pairs on podonotal shield and r2, r3, r5 along with larval s6 on lateral soft cuticle); j2 inserted in nearly transverse alignment with j1. Opisthonotal region holotrichous, with 15 pairs of setae (five pairs added to larval complement, including J1, Z1, Z2, S2, R1, all on interscutal soft cuticle; pygidial shield expanded so as to include eight pairs of setae, including J3, Z5, S4, S5 (all larval setae). Ventral idiosoma (Fig. 7D) with all setae aciculate, with no postlarval podosomal additions except st5. Sternal poroids iv1, iv2 present, iv3 absent, and larval funicules suppressed. Anal shield oval, with paranal setae similar in length or slightly shorter than postanal seta; rudiments of euanal setae suppressed. Opisthogastric soft cuticle with four or five pairs of setae, with asymmetrical addition of ZV5, which usually deutonymphal; JV5 relatively larger than in larva, and inserted more posteriorly, in transverse alignment with postanal seta. Gnathosoma ��� Fixed cheliceral digit with even row of 12-13 teeth; movable digit tridentate. Gnathotectum similar in form to adult. Subcapitulum with four pairs of setae, hypostomatic h1-h3 and capitular pc. Palpus with normal protonymphal complement of setae, including one on trochanter (Lindquist & Evans 1965); palptarsus with hyaline flap developed beside apotele as in adult. Legs ��� Legs I-IV with claws, pretarsi and shape of setae similar to adult female; tarsi II-IV with ventroapical process similar in form, serrated apically, as adult. Legs I, II, III, IV with normal, holotrichous protonymphal chaetotaxy (Lindquist & Evans 1965), except genu IV with accelerated presence of usually deutonymphal seta pl-1, and femur IV with accelerated alveolar indication of seta v-1 near basal lyriform fissure: coxae 2, 2, 2, 1; trochanters 4, 4, 4, 4; femora 10 (2 2/1 2/2 1), 8 (1 2/1 2/1 1), 5 (1 2/1 1/0 0), 4 (1 2/0 1/0 0); genua 8 (1 2/1 2/1 1), 6 (1 2/0 2/0 1), 6 (1 2/0 2/0 1), 6 (1 2/0 2/0 1); tibiae 8 (1 2/1 2/1 1), 7 (1 1/1 2/1 1), 7 (1 1/1 2/1 1), 7 (1 1/1 2/1 1); tarsi II-IV each with 17 (md added to larval complement). Tarsus IV with seta av-1 distinctively erect, longer and slightly thicker than all other tarsal setae (Fig. 12G), in contrast to its form on tarsi II, III of protonymph, and on tarsus IV of adult. Larva. General description, based on two unborn specimens, without measurements, for species-group perspective. Dorsal idiosoma (Fig. 7A) with all setae aciculate. Podonotal shield holotrichous with typical nine pairs of setae j (1, j3-j6, z2, z4, z5, s4); setae s6 on soft cuticle behind posterolateral margin of shield. Opisthonotal region holotrichous with 10 pairs of setae (J2-J5, Z3-Z5, S3-S5); pygidial shield margins mostly indiscernible on specimens at hand, but its surface with typical pair of arcuate lines near insertions of setae J4, and with typical complement of four pairs of setae, including tiny J4, J5 and prominent Z3, Z4. Soft interscutal cuticle with two pairs of mesonotal scutellae and typical six pairs of setae, including moderately long J2, J3, S3, S5, prominent Z5 (in ventrolateral position), and shorter S4. Ventral idiosoma (Fig. 7B) with all setae aciculate. Sternal shield outline not distinguishable; setae st1-st3 similarly short, st4, st5 absent, but the typical pair of subcutaneous structures (���funicules��� sensu Athias-Henriot 1971a) present posteromediad bases of legs III. Anal shield margins indistinguishable; anal valves with rudimentary pair of euanal setae; paranal setae slightly longer than postanal seta. Opisthogastric soft cuticle with typical four pairs of similarly short setae (JV1, JV2, JV5, ZV2). Gnathosoma ��� Fixed cheliceral digit with even row of 10-11 teeth; movable digit bidentate. Gnathotectum triramous, branches shorter than in adult. Subcapitulum with two pairs of hypostomatic setae, h1 longer than h2. Palpus with normal larval complement of setae (Lindquist & Evans 1965); palptarsus with hyaline flap not evident beside apotele. Legs ��� Legs I-III with claws, pretarsi and shape of setae similar to adult female; tarsi II, III with ventroapical process similar in form, serrated apically, as adult, and with setae d- (1) more strongly developed, slightl, Published as part of Moraza, Maria L. & Lindquist, Evert E., 2018, A new species-group with new species of the genus Lasioseius (Acari: Mesostigmata: Blattisociidae) associated with Neotropical hispine beetles in furled leaves ofHeliconia, pp. 62-98 in Acarologia 58 (1) on pages 74-83, DOI: 10.24349/acarologia/20184227, http://zenodo.org/record/4487537, {"references":["Lindquist E. E., Evans G. O. 1965 - Taxonomic concepts in the Ascidae, with a modified setal nomenclature for the idiosoma of the Gamasina (Acarina: Mesostigmata) - Mem. Entomol. Soc. Canada, 47: 1 - 64. doi: 10.4039 / entm 9747 fv","Athias-Henriot C. 1971 a - Un progres dans la connaissance de la composition metamerique des gamasides: leur sigillotaxie idiosomale (Arachnida) - Bull. Soc. zool. France, 96. 73 - 85.","Staines C. L., Garcia-Robledo C. 2014 - The genus Cephaloleia Chevrolat, 1836 (Coleoptera: Chrysomelidae: Cassidinae) - ZooKeys, 436: 1 - 355. doi: 10.3897 / zookeys. 436.5766","Garcia-Robledo C., Horvitz C. C., Kress J. W., Carvajal-Acosta N., Erwin T. L., Staines C. L. (2017) - Experimental assemblage of novel plant-herbivore interactions: ecological host shifts after 40 million years of isolation - Biotropica, 49 (6): 803 - 810. doi: 10.1111 / btp. 12464","Evans G. O. 1963 - Observations on the chaetotaxy of the legs in the free-living Gamasina (Acari: Mesostigmata) - Bull. br. Mus. nat. Hist. Zool., 10: 275 - 303. doi: 10.5962 / bhl. part. 20528"]}

Hispiniphis new genus (Figs 41���92, 101��� 105) Type species. Hispiniphis parva new species. Genus based on adult female, adult male, nymphal and larval material representing six species, of which one is newly described here, the others to be described subsequently. Diagnosis. Adult Hispiniphis are distinguished from those of Makarovaia in having the dorsal shield less extensive anterolaterally, leaving at least setae z 2, all s, and all r setae on adjacent soft cuticle, which has a thicker, micropunctate appearance distinct from the normally striate ventral soft cuticle. Adults have less strongly sclerotised and ornamented shields on the venter, including a smaller, inversely subtriangular anal shield, slightly longer than wide. Male stenitigenital shield incompletely formed, deeply excavated posteriorly, leaving setae st 5 on soft cuticle or on a separated platelet medially. Hypostome of nymphs and adults short, corniculi inserted at level only slightly behind insertions of hypostomatic setae hp 1. Movable cheliceral digit of female and nymphs tridentate, and with pointed process discernible on midventral face. Male chelicerae lacking a digitiform process at level where movable digit hinged to fixed digit, and with spermatodactyl projecting freely anteriorly. In addition to leg chaetotactic reductions noted for genus-group description, femur I and tibia I lacking dorsal seta ad - 3; genu II lacking lateral seta al - 2; genua II and III retaining ventral seta av. Tarsus I with ventrodistal acuminate process extending under pretarsus; tarsi II���IV with ventral projection under pretarsus seeming articulated, acuminate in immature instars and adults. Protonymph and to lesser extent deutonymph with opisthosoma reduced in form, size, sometimes flanked closely by posteriorly-directed legs IV. Description. Idiosomatic dorsum . Adult female (Fig. 42). Dorsal shield moderately well sclerotised, with reticulate ornamentation; shield relatively smaller in size, compared to overall dorsal surface area, not extensive enough to bear setae s 1, z 2, r 2, r 3 which remain with s 4, s 5, s 6, r 5 on surrounding soft cuticle; soft cuticle thickly striate, micropunctate, giving ornamented effect in contrast with opisthogastric soft cuticle (Fig. 103, 104). Dorsal shield with a maximum complement of 13-20 pairs of setae, including seven to 12 podonotal (j 1, j 3 ���j 6, z 4, z 5, z 6, sometimes j 1 off shield, sometimes j 4 duplicate) and six to 11 opisthonotal pairs (J 1 +, J 3, J 4 +, J 5, Z 2 ���Z 5, sometimes Z 2, Z 3, and Z 5 duplicate) (Fig. 88); dorsal setae usually similar in form, somewhat variable in length, except for J 5 always minute. Dorsal shield with complement of 13���15 pairs of discernible pore-like structures (three to five podonotal, 10 opisthonotal), of which five to seven pairs (two to four podonotal, three or four opisthonotal) superficially appear secretory (gland pores) and seven to nine pairs (one or two podonotal, six or seven opisthonotal) non-secretory (poroids). Soft lateral cuticle with 20���45 pairs of lateral, marginal and submarginal setae, including z 2, all s, all r, sometimes Z 2, and all S, R, and UR, and at least six pairs of marginal poroids, including idRp; few to many of setae on soft cuticle may be duplicate. Adult male ( Fig. 56). Dorsal shield usually similar in size, ornamentation and setation to that of female; sometimes capturing one to three more pairs of setae than in female; form and relative lengths of shield setae as in female. Idiosomatic venter. Adult female (Fig. 45). Tritosternum with laciniae fused for about half of their length, without basal elaborations; laciniae nearly devoid of pilosity (Fig. 47). Ventral shields variably poorly sclerotised. Presternal region smooth or faintly lineated, without platelets (Fig. 45). Sternal shield fragmented, its form and setation as noted for genus-group description; sternal area with varying faint ornamentation; shield fragment sometimes with bases of setae st 1, st 2 circumscribed by sclerotised thickenings, while unsclerotised medial region sometimes faintly ornamented but not striated with soft cuticle. Epigynal shield lightly sclerotised, unornamented, narrow, with hyaline anterior margin hardly widened between legs III, and with posterior margin hardly widened but strongly rounded or bluntly pointed (Fig. 48). Opisthosomatic venter lacking metapodal platelets, with soft striated cuticle bearing four to six pairs of opisthogastric setae (JV 1, JV 2, ZV 2, JV 5, sometimes JV 3, JV 4) and four pairs of poroids, flanked laterally and posteriorly by further thickened, ornate cuticle variably bearing several pairs of R - or UR - setae. Anal shield small, well sclerotised, smooth or reticulated, pyriform or inversely subtriangular, slightly longer than wide; para-anal setae inserted at mid-level of anus, shorter than postanal seta; shield with pair of gland pores gv 3 on lateral edges at level of insertions of para-anal setae, and with cribrum along tapered posterior margin (Fig. 49). Peritrematal shield weakly sclerotised, extending without interruption to about midlevel of coxae I; shield with gland pore gp 1 located at anterior extension at level of coxa I (Fig. 43); exopodal strip continuous alongside peritrematal shield only to level of coxae III, extensions between bases of coxae II���III rudimentary or absent. Peritremes thick, reaching at most to level of anterior margin of coxae II (Figs 43, 45). Spermathecal apparatus sometimes with small section of tubular piece discernibly sclerotised, slightly widened (Fig. 55). Adult male (Fig. 60). Form of tritosternum and presternal area as in female. Sternitigenital shield anterior margin well defined at level of genital opening; shield lightly sclerotised, united with endopodal strips between coxae I���II, II���III, III���IV; shield with four pairs of setae, st 1 -st 4, and two pairs of poroids; shield incompletely formed, deeply excavated posteriorly, leaving setae st 5 on soft cuticle or on a separated platelet medially (Figs 60, 62). Form and extent of exopodal and peritrematal structures much as in female (Fig. 60). Opisthogastric region much as in female, with same complement of poroids but only four or five pairs of opisthogastric setae (JV 1, JV 2, ZV 2, JV 5, sometimes JV 3). All ventral idiosomatic setae smooth, similar in shape and length as female. Anal shield similar in form and structure as in female. Gnathosoma. Female and male. Gnathotectum with anterior margin convex, denticulate (Figs 52, 57). Chelicerae generally as described for the genus-group description. Fixed cheliceral digit sparsely multidentate (offset subapical tooth and six to nine teeth), with pilus dentilis reduced to alveolar remnant or indiscernible; movable digit of female tridentate, with a small pointed process on mid-ventral face (Fig. 44). Movable cheliceral digit of male with one small tooth, and lacking small process on ventral face; spermatodactyl digit-like, variously sinuous, directed anteriorly, not recurved basally (Figs 58���59); male chela lacking any arthrodial process on paraxial surface at base of movable digit. Corniculi of female and male similar in form, short, stout, inserted at level of insertions of setae hp 1 (Figs 51, 53, 61); paraxial process at base of corniculus slightly shorter than corniculus (Fig. 51); bifid internal malae elongate, extending nearly to or beyond apex of palpgenu (Fig. 53); apex of labrum extending to palptibia. Subcapitulum with a pair of pore-like structures at level of insertion of corniculi (Fig. 53). Subcapitular setae smooth, with hp 1 clearly longer and thicker than other pairs. Deutosternum with seven moderately narrow, multidenticulate rows of denticles, connected by lateral margins; basal two rows not narrowed, sometimes more finely multidenticulate. Palpal setation and form of setae as described for genus-group description (Fig. 54 a���c); palptrochanter with inner seta somewhat longer than external setae (54 a). Legs. Female and male. Legs moderately short, similar in length to dorsal shield but clearly shorter than idiosoma. Legs I and IV similar in length and thickness to legs II, III (Figs 63���70). Tarsus I with sensilla s inconspicuous, short and acute (Fig. 72 a), and with ventrodistal acuminate process extending under pretarsus (Fig. 72 b). Pretarsus of legs I to IV with slender claws and large, rounded pulvillus; those of legs II���IV with short, inconspicuous paradactyli formed as narrow sclerotised shafts flanking claws (Fig. 74). Legs II to IV with tarsus (excluding pretarsus) about twice as long as tibia. Tarsi II���IV with apical setal processes ad - 1, pd - 1 inconspicuous, shorter than short pretarsi, and with apical ventral process short, triangular, more retrorse in male than in female (Figs 73, 74). Setation and its ontogeny on segments of legs I to IV deficient from full complement of Melicharidae (as presented by Lindquist & Evans 1965 for Melicharini), coxae, 2 - 2 - 2 - 1; trochanters, 5 - 5 - 5 - 5; femora, 11 (2 2 / 1 2 / 2 2) ��� 9 or 10 (1 2 or 3 / 1 2 / 2 1) ��� 6 (1 2 / 1 2 /0 0) ��� 6 (1 2 / 1 2 /0 0); genua, 11 (2 3 / 1 2 / 1 2) ��� 8 (1 3 / 1 2 /0 1) ��� 7 (1 2 / 1 2 /0 1) ��� 7 (1 2 /0 3 /0 1); tibiae, 10 (2 2 / 1 2 / 1 2) ��� 7 (1 1 / 1 2 / 1 1) ��� 7 (1 1 / 1 2 / 1 1) ��� 7 (1 1 / 1 2 / 1 1); trochanter I seta ad vestigial, only alveolus discernible. In addition to setal absences noted for family: femur I, tibia I, and sometimes femur II lacking seta ad - 3; genu II lacking seta al - 2; however, genua II���III retaining av, and genu IV retaining pd - 3. Several setae moderately thickened, spinelike: ad - 1, pd - 1, pd - 2 on femora I���II and ad 1, pd - 1 on femora III���IV; ad - 2 on femur IV. Seta av on femur III and pv - 2 on trochanters I���IV (on female and male) attenuated. Other leg setae simple, not strongly differentiated on either sex (male leg setal dimorphism not developed). Deutonymph. Idiosomatic dorsum. Discoidal form. Dorsal shield poorly sclerotised, without lateral incisions, lightly lineated, with ca. 35 pairs of similarly short setae except more posterior ones smaller: 17 pairs (j 1, j 3 ���j 6, z 2, z 4 ���z 6, s 1, s 2, s 4 ���s 6, r 2, r 3, r 5) on podonotal region, and a maximum 18 pairs (J 1, J 3, J 4, J 5, Z 2 ���Z 5, S 1 ���S 5, R 1 -R 6) on opisthonotal region (Figs 75); many of these setae displaced more marginally on dorsal shield than in adult. Ventrolateral soft cuticle nearly devoid of setae. Dorsal idiosomatic complement of poroids and gland-pores similar to that in female. Idiosomatic venter. Sternal region lightly sclerotised, unornamented, without endopodal extensions between coxae, with three pairs of platelets bearing sternal setae s 1 -st 3 and poroids iv 1, iv 2; setae st 4 -st 5 inserted on soft cuticle between coxae III and IV (Fig. 76); poroids iv 5 absent. Anal shield prominent, lightly sclerotised, unornamented, roughly rounded, with circumanal setae similar in length; adanal gland pores (gv 3) at level of paraanal setae; cribrum as in adult female. Opisthogaster with four to six pairs of ventral setae (the four larval JV 1, JV 2, JV 5, ZV 2, sometimes plus two minute JV 3, JV 4), sometimes not flanked by further setae on soft cuticle (Fig. 76). Peritrematal shields and peritremes shortened, paedomorphic, as in protonymphal instar, developed only alongside level between coxae III���IV (Fig. 76). Peritrematal gland pore gp 2 and poroid ip 2 on soft cuticle between levels of coxae II and III; poststigmatic poroid ips adherent to stigma on posterior edge of peritrematal shield, and poststigmatic gland pore gp 3 and poroid ip 3 on soft cuticle. Rim of exopodal plate behind coxa IV inconspicuous or indiscernible, with gland pore gv 2 not discernible. Gnathosoma. Gnathotectum, chelicerae and other mouthpart structures, corniculi and adjacent structures as in adult female; subcapitular and palpal structures similar to those in adult female. Legs. Pretarsal structures, and chaetotaxy of legs I���IV as in adult, including these additions to protonymphal setal complement: av - 2 on trochanters I���IV; al - 2 and pl - 2 on genu I, ad - 3 on genua I���II, pd - 3 on genu IV, pl - 1 on genu IV, av - 1 on genua II���IV, pv - 1 on genu II, al - 2 and pl - 2 on tibia I. Form and shape of leg setae similar to those on adult female, except seta pl on femur, genu, tibia, basitarsus of leg IV strongly developed and recurved/bent. Legs IV directed posteriorly, as in protonymph. Tarsi II���IV with form of setae and acuminate form of apical ventral process as in protonymph. Protonymph. Idiosomatic dorsum . Discoidal to ovoidal form. Dorsal shielding weakly sclerotised, with or without delineation between podonotal and pygidial shields, and with faint lateral, mostly podonotal ornamentation; mesonotal scutellae not evident (Figs 77, 89); dorsum maximally with 24 to 33 pairs of similarly short setae except more posterior ones smaller, J 5 minute (protonymphal setae s 5, r 2, r 3, r 5, Z 2, S 2, R 1, sometimes J 1 added to larval complement, j 2 absent); setae J 3, J 5, Z 4, Z 5 sometimes on pygidial shield if delineated (Fig. 89). Sometimes as many as eight pairs of dorsal deutonymphal setae (s 1, z 6, S 1, R 2 -R 6) may accelerate their ontogeny to this instar (Figs 77, 78). Body dorsum with complement of 11 pairs of discernible pore-like structures, of which six non-secretory (poroids) and five superficially appear to be secretory (gland pores). Idiosomatic venter. Sternal region with two or three pairs of weakly sclerotised platelets (without endopodal elements) bearing setae st 1, st 2 and poroids iv 1, iv 2 (iv 2 added to larval complement); setae st 3 on either a third pair of weak platelets or on soft cuticle; setae st 5 rudimentary, paragenital poroids indiscernible. Opisthogastric region reduced to small area behind coxae IV. Anal shield faintly sclerotised, unornamented, yet prominent, about as wide as long, with postanal seta longer than para-anal setae; para-anal setae inserted at mid-level of anal opening, roughly in transverse line with adanal gland pores; cribrum similar in development as in deutonymph. Opisthogaster with four or six pairs of ventral setae; JV 1, JV 2, sometimes ZV 2 anterior to anal shield; larval complement sometimes augmented by normally deutonymphal setae JV 3, JV 4 on posterior margin, along with larval JV 5 (Figs 78, 90). Peritrematal region on each side with poroid and gland pore (ip 2, gp 2) on soft cuticle at mid-level of coxae III, anterior to apex of weakly formed peritreme between coxae III and IV, and with poststigmatic poroid and gland pore (ip 3, gp 3) on soft cuticle behind peritreme. Exopodal rim and gland pore gv 2 behind coxa IV not discernible. Gnathosoma. Form of gnathotectum, corniculus and adjacent paraxial process, elongated internal malae (Fig. 85), and other gnathosomatic structures similar to those in deutonymph, except palpi with normal protonymphal complement of setae (see Evans 1964), including only one trochanter seta. Cheliceral movable digit bi- or tridentate, with mid-ventral projection rudimentary or indiscernible; fixed digit weakly multidenticulate, with offset subapical tooth and sparse (five or six) teeth along masticatory surface (Fig. 87). Legs. Legs I to IV with pretarsi, well-developed claws, and normal protonymphal complement of setae as described for Ascidae by Lindquist & Evans (1965) except for absence of second ventral seta on femur II: coxae, 2 - 2 - 2 - 1; trochanters 4 - 4 - 4 - 4 (1 0/ 2 1 on I, II, 1 1 / 2 0 on III, IV); femora 10 (2 2 / 1 2 / 1 2) ��� 7 (1 2 / 1 2 /0 1) ��� 5 (1 2 / 1 1 / 0 0) ��� 4 or 5 (1 2 /0 1 /0 0 or 1); genua 8 or 9 (1 2 or 3 / 1 2 / 1 1) ��� 6 or 7 (1 2 or 3 /0 2 /0 1) ��� 6 (1 2 /0 2 /0 1) ��� 5 or 6 (1 2 /0 2 /0 0 or 1); tibiae 8 (1 2 / 1 2 / 1 1) ��� 7 (1 1 / 1 2 / 1 1) ��� 7 (1 1 / 1 2 / 1 1) ��� 7 (1 1 / 1 2 / 1 1); normally deutonymphal seta pl - 1 sometimes accelerated in appearance on femur and genu IV, and ad - 3 similarly on genua I and II. Coxae I���II distal rims with serrated posteroventral strips. Leg setae generally simple. Legs IV directed posteriorly, with seta pl on femur, genu, tibia, and to less degree on tarsus IV slightly to clearly thicker and longer than other setae of segments (Fig. 79). Tarsi II���IV with setae ad - 1, pd - 1 small but as long as short pretarsus; md, al - 2, av - 1, pl - 1 and pl - 2 longer, thinner than other tarsal setae, except av - 2 similarly attenuate on tarsus IV; apical ventral tarsal process acutely triangular. Larva. Idiosomatic dorsum . Body with dorsal shielding weakly sclerotised, not clearly delimited, less discernibly ornamented than on nymphal instars; mesonotal scutellae indiscernible (Figs 80, 91). Body dorsum with 17 pairs of setae: nine podonotal pairs, and eight opisthonotal pairs (J 2); discernible pore-like structures include eight non-secretory poroids (three podonotal, five opisthonotal) and three apparently secretory gland pores (two podonotal, one opisthonotal). Peritrematal region on each side with gland pore gp 1 at level of setae j 3, gland pore gp 2 and poroid ip 2 at level above coxae II���III, and poststigmatic gland pore gp 3 and poroid ip 3 at level of anal shield anterior margin. Idiosomatic venter. Tritosternum normally developed, similar in form as in subsequent instars. Sternal region with two or three pairs of weakly (hardly discernible) sclerotised platelets bearing setae st 1, st 2 and poroids iv 1; setae st 3 on soft cuticle or on a third pair of platelets; sometimes protonymphal setae st 5 accelerately present between coxae III (Figs 81, 92); pair of subcutaneous structures sometimes discernible at level of posterior margins of coxae III. Anal shield faintly sclerotised, elongate or as wide as long; para-anal setae inserted at level slightly posterior to anal opening, and slightly longer than postanal seta; adanal gland pores (gv 3) at level of postanal seta; cribrum rudimentary or absent. Venter with six pa, Published as part of Moraza, Mar��a L. & Lindquist, Evert E., 2015, Systematics and biology of mites associated with neotropical hispine beetles in unfurled leaves of Heliconia, with descriptions of two new genera of the family Melicharidae (Acari: Mesostigmata: Gamasina: Ascoidea), pp. 301-351 in Zootaxa 3931 (3) on pages 326-330, DOI: 10.11646/zootaxa.3931.3.1, http://zenodo.org/record/244189, {"references":["Lindquist, E. E. & Evans, G. O. (1965) Taxonomic concepts in the Ascidae, with a modified setal nomenclature for the idiosoma of the Gamasina (Acarina: Mesostigmata). Memoirs of the Entomological Society of Canada, 47, 1 - 64.","Evans, G. O. (1964) Some observations on the chaetotaxy of the pedipalps in the Mesostigmata (Acari). Annals and Magazine of Natural History, Series 13, 6, 513 - 527.","Staines, C. L. (1996) The genus Cephaloleia (Coleotera: Chrysomelidae) in Central America and the West Indies. Revista Biologia Tropical, Special Number, 3, 3 - 87.","Garcia-Robledo, C. & Staines, C. L. (2008) Herbivory in gingers from latest Cretaceous to present: is the ichnogenus Cephaloleichnites (Hispinae, Coleoptera) a rolled-leaf beetle? Journal of Paleontology, 82, 1035 - 1037. http: // dx. doi. org / 10.1666 / 07 - 089.1","Staines, C. L. (2009) A review of the genus Chelobasis Gray, 1832 (Coleoptera: Chrysomelidae: Cassidinae). Zootaxa, 2033, 58 - 68.","Garcia-Robledo, C., Horvitz, C. C. & Staines, C. L. (2010) Larval morphology and development, host plants, adult longevity, sexual dimorphism and notes on natural history in Cephaloleia rolled-leaf beetles (Coleoptera: Chrysomelidae: Hispinae). Zootaxa, 2610, 50 - 68.","Staines, C. L. & Garcia-Robledo, C. (2014) The genus Cephaloleia Chevrolat, 1836 (Coleoptera: Chrysomelidae: Cassidinae). Zookeys, 436, 1 - 355. http: // dx. doi. org / 10.3897 / zookeys. 436.5766","Lindquist, E. E. & Moraza, M. L. (1999) Observations on homologies of idiosomal setae in Zerconidae (Acari: Mesostigmata), with modified notation for some posterior body setae. Acarologia, 39, 203 - 226."]}

Makarovaia new genus (Figs 1���40, 98��� 100) Type species. Makarovaia ornata new species. Genus based on adult female, male, nymphal and larval material representing one newly described species. Diagnosis. Adults of Makarovaia are distinguished from those of Hispiniphis in having the dorsal shield more extensive anterolaterally, so as to bear setae z 2, s 1, s 4, r 2, r 3, and in having more strongly sclerotised and ornamented shields on the venter, including a moderately large, obovate anal shield slightly wider than long. Male sternigenital shield fully formed, with endopodal extensions between leg bases, a convex posterior margin between bases of legs IV, and bearing all five pairs of sternal setae. Hypostome of nymphs and adults elongate, resulting in corniculi inserted relatively posteriorly, at level about halfway between insertions of hypostomatic setae hp 1 and hp 3. Movable cheliceral digit of female and female nymphs multidenticulate (���four teeth), and with bluntly pointed process distinct on midventral face. Male chelicera with ciliated, flexible, digitiform process paraxially at level where movable digit hinged to fixed digit, and with spermatodactyl abruptly bent downward near it base, directing the freely extending part posteroventrally. In addition to leg chaetotactic reductions noted for the genusgroup description, genua II and III lacking ventral setae (av absent); femora I���II and tibia I retaining seta ad - 3, genu II retaining al - 2. Tarsus I without ventrodistal acuminate process extending under pretarsus; tarsi II���IV with ventral projection under pretarsus acuminate in immature instars but truncate in adult. Nymphs with opisthosoma normal in fully-developed form, size, and not closely flanked by legs IV, which directed posterolaterally. Description. Idiosomatic dorsum. Adult female (Fig. 1). Dorsal shield well sclerotised, sufficiently expansive anterolaterally to bear setae s 1, z 2, r 2, r 3; surrounding soft cuticle thickly formed and ornamented, in distinction to simply striated ventral cuticle, as noted for family. Dorsal shield with maximum complement of 18 pairs of setae, including 12 podonotal (j 1, j 3 ���j 6, z 2, z 4, z 5, s 1, s 4, r 2, r 3) and six opisthonotal pairs (J 4, J 5, Z 2 ���Z 5); dorsal setae similar in form and general size except J 5 always minute. Dorsal shield with complement of 17 pairs of discernible pore-like structures (nine podonotal, eight opisthonotal), of which five pairs (three podonotal, two opisthonotal) superficially appear secretory (gland pores) and 12 pairs (six podonotal, six opisthonotal) non-secretory (poroids). Soft lateral cuticle with ca. 20���25 pairs of lateral and marginal setae (s 5, s 6, r 5, S 1 ���S 5, R 1 ���R 6, and six or more UR ���s), and at least six pairs of marginal poroids, including idRp. Adult male (Fig. 21). Dorsal shield similar in size, ornamentation and setation to that of female; form and relative lengths of setae on dorsal shield and dissimilar structure of surrounding soft cuticle as in female. Idiosomatic venter. Adult female (Figs 5���11). Tritosternum with laciniae free for most of length, without basal elaborations; laciniae nearly devoid of pilosity (Fig. 11). Ventral shields well sclerotised and ornamented. Presternal region faintly lineated, without platelets (Fig. 5). Sternal shield fragmented, form and setation as noted for genus-group description; sternal shield pieces reticulated, and unsclerotised medial region faintly longitudinally lineate or reticulate; setae st 3 and st 4 on smooth cuticular area on either side (perhaps unsclerotised sternal remnants). Epigynal shield well sclerotised and ornamented, narrow, with hyaline anterior margin not widened between legs III, but with posterior margin widened, strongly convex. Opisthosomatic venter lacking metapodal platelets, with soft cuticle normally striated but not further ornamented, and with ca. nine pairs of opisthogastric setae (JV 1 ���JV 5, ZV 2 ���ZV 5) and four pairs of poroids flanked by several pairs of R - or UR -setae. Anal shield moderately large, well sclerotised and ornamented, ovoid, wider than long; para-anal setae inserted near anterior level of anus, similar in length to postanal seta; shield with pair of gland pores gv 3 slightly within lateral margins posterior to level of insertions of para-anal setae, and with extensive cribrum along posterior margin. Peritrematal shield poorly developed, interrupted at level of coxae II and with a separate platelet bearing gland pore gp 1 at level between coxae I and II (Fig. 5); exopodal strip continuous alongside peritrematal shield only to level of coxae III, extensions between bases of coxae I���II, II���III absent. Peritremes wide, shortened, reaching at most to mid-level of coxa II. Spermathecal apparatus with small section of tubular piece discernibly sclerotised, its distal portion widened (Fig. 3). Adult male (Fig. 25). Form of presternal area as in female, tritosternum with laciniae similar in length to those in female. Sternitigenital shield anterior margin strongly defined at level of genital opening; shield well sclerotised, united with endopodal strips between coxae I���II, II���III, III���IV, with five pairs of setae and two pairs of poroids. Form and extent of exopodal and peritrematal structures much as in female, except fragment absent between coxae I���II, and peritremes slightly shorter. Opisthogastric region much as in female, with same pairs of opisthogastric setae and poroids, flanked by several (fewer) pairs of R - or UR -setae. All ventral idiosomatic setae smooth, similar in shape and length as in female. Anal shield much as in female, but with para-anal setae inserted at mid-level of anus. Gnathosoma. Female and male. Gnathotectum with anterior margin convex to nearly angular, denticulate (Figs 2, 22). Chelicerae generally as described for the genus-group description (Figs 6, 7, 23, 24). Fixed cheliceral digit multidentate, with pilus dentilis reduced to an alveolar remnant; movable digit of female multidentate (Fig. 7), with a small blunt process on mid-ventral face. Movable cheliceral digit of male bidentate, lacking pointed process on ventral face; spermatodactyl digit-like, abruptly curved ventrally near base and directed posteroventrally (Fig. 24); male chela with conspicuous ciliated arthrodial process on paraxial surface at base of movable digit (Fig. 24). Corniculi of female and male similar in form, short, stout, inserted at level midway between insertions of subcapitular setae hp 1 and hp 3, and with tip of paraxial acuminate process reaching slightly beyond that of main corniculus tip (Figs 10, 27, 28); bifid internal malae elongate, extending well beyond apex of palpgenu (Figs 10, 28); apex of labrum extending to mid-level or apex of palptibia. Subcapitulum with a pair of pore-like structures at level of insertion of corniculi (Fig. 10). Subcapitular setae hp 1 much longer and thicker than other pairs. Deutosternum with seven or eight moderately narrow rows of denticles, basal rows slightly narrower; rows similarly multidenticulate, connected by lateral margins. Palpal setation and form of setae as described for genusgroup description (Figs 8 a, b, c); palptrochanter with inner seta twice as long as external seta (Fig. 8 a). Legs. Female and male. Legs relatively short, clearly shorter than dorsal shield. Legs similar in size, II and III slightly thicker than legs I and IV (Figs 12���15). Distal posterior ventral and dorsal margins of coxae I���IV serrated, and coxa I with a serrated ridge on ventral face. Pretarsi of legs I to IV with slender claws and large, rounded pulvillus; those of legs II���IV with short, inconspicuous paradactyli (Fig. 16). Tarsus I with sensilla s inconspicuous, short and acute (Fig. 20). Legs II to IV with tarsus (excluding pretarsus) about twice as long as tibia. Tarsi II���IV with apical setal processes ad- 1, pd- 1 reduced, shorter than short pretarsi (Fig. 16), and with apical ventral process short, truncated, unhinged basally (Figs 17, 18). Setation and its ontogeny on segments of legs I to IV deficient from full complement of Melicharidae (as presented by Lindquist & Evans 1965 for Melicharini): coxae, 2 - 2 - 2 - 1; trochanters, 5 - 5 - 5 - 5; femora, 12 (2 3 / 1 2 / 2 2) ��� 10 (1 3 / 1 2 / 2 1) ��� 6 (1 2 / 1 1 /0 1) ��� 6 (1 2 / 1 1 /0 1); genua, 11 (2 3 / 1 2 / 1 2) ��� 8 (2 3 /0 2 /0 1) ��� 6 (1 2 /0 2 /0 1) ��� 6 (1 2 /0 2 /0 1); tibiae, 11 (2 3 / 1 2 / 1 2) ��� 7 (1 1 / 1 2 / 1 1) ��� 7 (1 1 / 1 2 / 1 1) ��� 7 (1 1 / 1 2 / 1 1); trochanter I seta ad vestigial, only alveolus discernible; femora III���IV with seta pd - 2 inserted relatively laterally, in pl position. In addition to setal absences noted for family: genua II���III lacking av, and genu IV lacking pd - 3; however, femora I���II and tibia I retaining seta ad - 3. Several setae moderately thickened, spinelike: pl on trochanters I and II, and al on trochanter IV; ad - 1, pd - 1, pd - 2 on femora I���II and ad 1, ad - 2, pd, pl on femora III���IV; some ventral setae on femora I���IV (on female and male); pv on genu and tibia I, and pv and to less extent av on tibiae II���IV. Seta pv - 2 on trochanters I���IV attenuate, thinner and longer than other setae. Male leg setal dimorphism limited to form of some ventral and lateral setae on tarsi II to IV. Tarsi II���IV in both sexes with setae al - 2, pl - 1, slightly longer, more attenuated than other tarsal setae; on female, ventral setae av - 1, pv - 1 similar to pl - 1, while al - 1, av - 2, pv - 2 and moreso mv moderately spine-like (Fig. 17); on male, al - 1, av - 1, pv - 1, av - 2, pv - 2 and mv thickened, similarly spine-like with swollen, bulbous base (Fig. 18); apical ventral tarsal process truncate-acuminate on female (Fig. 17) but truncate-straight on male (Fig. 18). Other leg setae simple, not strongly differentiated on either sex. Deutonymph. Idiosomatic dorsum . Dorsal shield extensive, poorly sclerotised, without lateral incisions, lightly lineated, with 18 pairs of similarly smooth, short setae except J 5 minute: 12 pairs (j 1, j 3 ���j 6, z 2, z 4, z 5, s 1, s 4, r 2, r 3) on podonotal region, and six pairs (J 4 -J 5, Z 2 ���Z 5) on opisthonotal region (Figs 29, 30). Lateral soft cuticle with s 5, s 6, r 5 on podonotal region and S 1 -S 5, R 1 -R 6 and variably few UR -setae on opisthonotal region. Dorsal idiosomatic complement of poroids and gland-pores similar to that in female. Idiosomatic venter. Sternal shield lightly sclerotised and ornamented, with endopodal extensions between coxae I���II, II���III and III���IV, with four pairs of setae st 1 -st 4 and poroids iv 1 and iv 2; poroids iv 3 absent; setae st 5 on soft cuticle together with poroids iv 5 between coxae IV (Fig. 30). Anal shield lightly sclerotised and ornamented, with circumanal setae short and similar in length, adanal gland pores (gv 3) at level between para-anal and postanal setae; cribrum large as in adult. Opisthogaster with nine pairs of ventral setae in deutonymph (the four larval JV 1, JV 2, JV 5, ZV 2 plus five minute deutonymphal JV 3, JV 4, ZV 3 ���ZV 5, flanked by a few either S - or R - or UR - setae) (Fig. 30). Peritrematal shields and peritremes shortened, paedomorphic, as in protonymphal instar, developed only between midlevels of coxae III-IV (Fig. 30). Peritrematal gland pore gp 2 and poroid ip 2 indistinct on soft cuticle; poststigmatic poroid ips adherent to stigma on posterior edge of peritrematal shield, but poststigmatic gland pore gp 3 and poroid ip 3 indistinct on soft cuticle in that area. Rim of exopodal plate behind coxa IV inconspicuous, with gland pore gv 2. Gnathosoma. Gnathotectum, chelicerae and other mouthpart structures, corniculi and adjacent structures as in adult female (Fig. 31), but fixed cheliceral digit less hooked apically; deutosterum with seven rows of denticles (two basal rows narrower and less arched than anterior ones), with structures otherwise as in adult; palpi similar to those in adult female, including similar form of al setae on palpfemur and palpgenu. Chelicerae of male deutonymph with movable digit thick, with apical hook and one distal minute tooth and formation of primordial sperm duct present (Fig. 40). Legs. Pretarsal structures and chaetotaxy of legs I���IV as in adult, including these additions to protonymphal setal complement: av - 2 on trochanters I-IV; ad - 3 on femur I, pd - 2 on femora I, III, IV, pv - 2 on femora I, II, al - 2 and pl - 2 on genu I, ad - 3 on genua I���II, pl - 1 on genu IV, al - 2 and pl - 2 on tibia I. Size and shape of leg setae similar to those on adult female. Trochanter I with postero-dorsal serrated ridge. Tarsi II���IV with form of setae and acuminate form of apical ventral process as in protonymph (Fig. 26). Protonymph. Idiosomatic dorsum . Body with dorsal shielding not clearly delimited (outlines of podonotal shield, mesonotal scutellae, pygidial shield not evident), but with faint lateral lineation, less than in deutonymph; dorsum maximally with 24 pairs of similarly smooth, short setae, except J 5 minute (protonymphal setae r 2, r 3, r 5, R 1, Z 2 added to larval complement; j 2, J 1, Z 1 absent, as on deutonymph and adult, but S 2 not discerned). Body dorsum with complement of 12 pairs of discernible pore-like structures, of which seven non-secretory (poroids) and five superficially appear to be secretory (gland pores); paravertical poroids idj 1 added to larval complement. Idiosomatic venter. Sternal shield weakly sclerotised and ornamented, with faint endopodal extensions between coxae I���II, and II���III, with three pairs of setae st 1 -st 3 and two pairs of poroids (iv 2 added to larval complement); setae st 5 on soft cuticle at level of posterior border of coxae IV (Fig. 33); paragenital poroids absent. Anal shield faintly sclerotised and ornamented, roughly as wide as long, with circumanal setae similarly short; para-anal setae inserted at level slightly posterior to anal opening; adanal gland pores (gv 3) at level of postanal seta; cribrum large, as in deutonymph. Opisthogaster with four to six pairs of ventral setae; larval complement sometimes augmented by normally deutonymphal setae JV 3, JV 4 (Fig. 33). Peritrematal region on each side with poroid and gland pore on soft cuticle alongside weakly formed peritreme between coxae III and IV, and with poststigmatic poroid and gland pore on soft cuticle behind peritreme (Fig. 33). Exopodal rim behind coxa IV not discernible, but gland pore gv 2 present on inguinal area. Gnathosoma. Form of gnathotectum, corniculus and its paraxial acuminate process, elongated internal malae, and other gnathosomatic structures similar to those in deutonymph, except palpi with normal protonymphal complement of setae (see Evans 1964), including only one trochanter seta; corniculi inserted clearly posterior to level of insertions of setae hp 1, and with tip of inner acuminate process reaching beyond that of main corniculus tip (Fig. 37 b). Cheliceral movable digit with three or usually four teeth and a small mid-ventral projection; fixed digit multidentate, with offset subapical tooth and sparse (five to seven), well-formed teeth along masticatory surface (Fig. 39). Legs. Legs I to IV with pretarsi endowed by well-developed claws and normal protonymphal complement of setae as described for Ascidae by Lindquist & Evans (1965): coxae, 2 - 2 - 2 - 1; trochanters 4 - 4 - 4 - 4 (1 0/ 2 1 on I, II, 1 1 / 2 0 on III, IV); femora 10 (2 2 / 1 2 / 1 2) ��� 8 (1 2 / 1 2 / 1 1) ��� 5 (1 2 / 1 1 /0 0) ��� 4 (1 2 /0 1 /0 0); genua 8 (1 2 / 1 2 / 1 1) ��� 6 (1 2 /0 2 /0 1) ��� 6 (1 2 /0 2 /0 1) ��� 6 (1 2 /0 2 /0 1); tibiae 8 (1 2 / 1 2 / 1 1) ��� 7 (1 1 / 1 2 / 1 1) ��� 7 (1 1 / 1 2 / 1 1) ��� 7 (1 1 / 1 2 / 1 1); seta pl - 1 accelerated in first presence in protonymph, rather than deutonymph, on genu IV. Coxae I���II distal rims with serrated posterodorsal strips. Leg setae generally simple, not markedly differentiated. Tarsi II���IV with setae ad - 1, pd - 1 small but as long as short pretarsus; al - 2, a v - 1, pv - 1, pl - 1 and md longer, thinner than other tarsal setae; apical ventral tarsal process present, bluntly triangular. Larva. Idiosomatic dorsum . Body with dorsal shielding not clearly delimited, and surface not discernibly ornamented as on nymphal instars; mesonotal scutellae indiscernible (Fig. 34). Body dorsum with 20 pairs of smooth setae: ten podonotal pairs, and ten opisthonotal pairs (J 1 absent, J 5 miniscule, S 3 ���S 5 inserted ventrolaterally); discernible pore-like structures include five (one podonotal, four opisthonotal) poroids, and one (opisthonotal) gland pore. Idiosomatic venter. Tritosternum normally developed, similar in form as in subsequent instars. Sternal shield not distinguishable, intercoxal region with setae st 1 ���st 3 and poroids iv 1; pair of subcutaneous structures (prevalent among larvae of Gamasina) indiscernible at level of posterior margins of coxae III. Anal shield faintly sclerotised, roughly as wide as long; para-anal setae inserted at level slightly posterior to anal opening, and longer than postanal seta; adanal gland pores (gv 3) at level of postanal seta; cribrum absent. Venter with four pairs of opisthogastric setae on soft cuticle anterolateral to anal shield, flanked by opisthonotals (S 3 ���S 5). Poststigmatic poroid ip 3 and gland pore gp 3 on soft cuticle near level of posterior margin of coxae III (Fig. 35). Gnathosoma. Gnathotectum with convex anterior margin less denticulate than in subsequent instars (Fig. 36). Cheliceral movable digit with two teeth and a small mid-ventral projection; dentition of fixed digit weakly developed, with offset subapical tooth and masticatory ridge with a weak tooth proximally (Fig. 38). Form of corniculus and its inner acuminate process much as in subsequent instars, but inserted at same transverse level as alveoli of setae hp 1, and with inner process shorter relative to subsequent instars, its tip not reaching that, Published as part of Moraza, Mar��a L. & Lindquist, Evert E., 2015, Systematics and biology of mites associated with neotropical hispine beetles in unfurled leaves of Heliconia, with descriptions of two new genera of the family Melicharidae (Acari: Mesostigmata: Gamasina: Ascoidea), pp. 301-351 in Zootaxa 3931 (3) on pages 311-315, DOI: 10.11646/zootaxa.3931.3.1, http://zenodo.org/record/244189, {"references":["Lindquist, E. E. & Evans, G. O. (1965) Taxonomic concepts in the Ascidae, with a modified setal nomenclature for the idiosoma of the Gamasina (Acarina: Mesostigmata). Memoirs of the Entomological Society of Canada, 47, 1 - 64.","Evans, G. O. (1964) Some observations on the chaetotaxy of the pedipalps in the Mesostigmata (Acari). Annals and Magazine of Natural History, Series 13, 6, 513 - 527.","Strong, D. R. (1977) Rolled-leaf hispine beetles and their Zingiberales host plants in Middle America. Biotropica, 9, 156 - 169. http: // dx. doi. org / 10.2307 / 2387878","Staines, C. L. (2009) A review of the genus Chelobasis Gray, 1832 (Coleoptera: Chrysomelidae: Cassidinae). Zootaxa, 2033, 58 - 68.","Evans, G. O. (1963) Observations on the chaetotaxy of the legs in the free-living Gamasina (Acari: Mesostigmata). Bulletin of the British Museum (Natural History). Zoology, 10, 275 - 303.","Micherdzinski, W. (1969) Die Familie Parasitidae Oudemans 1901 (Acarina, Mesostigmata). Zaklad Zoologii Systematycznej Polskiej Akademii Nauk. Krakow, Panstwowe Wydawnictwo Naukowe, 690 pp."]}