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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

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"]}

A case of homonymy in the genus Zonurobia is corrected by proposing Zonurobia circularis limpopoensis as a replacement name for Zonurobia circularis transvaalensis Lawrence, not Zonurobia transvaalensis Lawrence., Acarologia, 60, 557-558

Perscheloribates (Perscheloribates) parakontumensis n. sp. Zoobank: 0E24D8AB-6CBD-4AF0-BFDD-0A7CE11E7EE3 (Figures 4–6) Diagnosis — Body size: 348–365 × 182–199. Rostrum pointed. Prolamella and translamella absent. Rostral, lamellar and interlamellar setae long, setiform, barbed; ro shortest, in longest. Bothridial seta long, lanceolate, with long, setiform apex, barbed. Lateral keel-shaped ridge present. Notogastral setae short, setiform, roughened. Epimeral and anogenital setae setiform, roughened. Description — Measurements – Small species. Body length: 348 (holotype, female), 348– 365 (three paratypes, one male and two females); notogaster width: 182 (holotype), 182–199 (three paratypes). Integument – Body color light brown. Body surface punctate (visible under high magnification in dissected specimens). Lateral part of prodorsum slightly microgranulate. Prodorsum (Figs 4a, 5a) – Rostrum pointed. Lamella located dorsolaterally, about half of prodorsum (measured in lateral view). Sublamella thin, similar to lamella in length. Sublamellar porose area (4) rounded. Prolamella absent, but its basal part sometimes developed. Translamella absent. Rostral (53–61), lamellar (69–73) and interlamellar (77–82) setae setiform, barbed. Bothridial seta (65–73) lanceolate, barbed, with long stalk and shorter head having long setiform apex. Exobothridial seta (20–24) setiform, slightly barbed. Alveolar vestige of second exobothridial seta and lateral keel-shaped ridge present. Sejugal porose area not observed. Dorsophragma slightly elongated. Notogaster (Figs 4a, 5 a-c) – Anterior notogastral margin convex medially. Pteromorph triangular, broadly rounded distally. Ten pairs of notogastral setae (10–12) setiform, roughened. Four pairs of sacculi with small opening and drop-like chamber. Distance S1–S1 shorter than S2–S2. Opisthonotal gland opening and all lyrifissures distinct. Circumgastric scissure often not observed. Circumgastric sigillar band visible. Gnathosoma (Fig. 4b) – Similar to Perscheloribates paracuriosus n. sp. Subcapitulum longer than wide (82–90 × 65–73). Three pairs of subcapitular setae setiform, barbed; h (24– 28) longer than a (18–20) and m (10–12), m thinnest. Two pairs of adoral setae (10) setiform, barbed. Palp (49–53) with setation 0-2-1-3-9(+ω). Postpalpal seta (4) spiniform. Chelicera (90–94) with two setiform, barbed setae, cha (32–36) longer than chb (16–20). Trägårdh’s organ narrowly triangular. Epimeral and lateral podosomal regions (Figs 4b, 5a) – Epimeral setal formula 3-1-3-3. Setae setiform, roughened; 1b, 1c, 3b, 3c, 4c (16–24) longer than 1a, 2a, 3a, 4a, 4b (10–12). Setae 1c inserted laterally on pedotectum I. Pedotectum II quadrangular in ventral view, with one protruding tip. Discidium rounded distally. Circumpedal carina long, reaching pedotectum II. Anogenital region (Figs 4b, 5 a-c) – Four pairs of genital (8–10), one pair of aggenital (10–12), two pairs of anal (10–41) and three pairs of adanal (10–12) setae setiform, roughened. Adanal lyrifissure located close and parallel to anal plates. Marginal porose area complete, band-like. Preanal organ of typical, goblet-like form. Ovipositor elongated (134 × 28), blade (61) shorter than length of distal section (beyond middle fold; 73). Each of the three blades with four smooth setae, ψ 1 ≈ τ 1 (24) setiform, ψ 2 ≈ τa ≈ τb ≈ τc (12) thorn-like. Six coronal setae (2) spiniform. Legs (Figs 6a-d) – Claw of leg pretarsus sparsely barbed on dorsal side. Porose area on femora I–IV and on trochanters III, IV slightly visible; ventral porose area in basal part of tarsus and distal part of tibia not observed. Formulas of leg setation and solenidia: I (1-5-3-4-19) [1-2-2], II (1-5-2-4-15) [1-1-2], III (2-3-1-3-15) [1-1-0], IV (1-2-2-3-12) [0-1-0]; homology of setae and solenidia indicated in Table 2. Seta it present on all tarsi. Famulus of tarsus I short, erect, slightly swollen distally, inserted between solenidion ω 1 and seta ft”. Solenidion ω 1 on tarsus I, ω 1 and ω 2 on tarsus II and σ on genu III bacilliform, other solenidia setiform. Material examined — Holotype (female) and three paratypes (one male and two females): U.S.A., Michigan, Emmet Co., Wilderness State Park, 45°44’N, 84°55’W, phoretic on specimens of Phellopsis obcordata (Kirby, 1873) (Coleoptera, Zopheridae), May 1981 (J. Kukor). Host specimen in UMMZ bearing the voucher label, “Mites removed, B.M. OConnor #81-0604-001.” Mites were found on 3 specimens of P. obcordata collected from a fruting body of the polypore fungus, Porodaedalea pini (Brot.) Murrill, 1905 (= Fomes pini, = Phellinus pini), growing on red pine, Pinus resinosa Alton, 1789. Mites were found in pits on the dorsal surface of the elytra. The habitat consists of around 4050 hectares of dense coniferous and mixed hardwood forest on the northeast shore of Lake Michigan. The host beetle is associated with several species of polypore fungi in dense boreal forests in Eastern North America (Evans 2014), suggesting that the normal habitat of this mite species is either on or in the fungal fruiting bodies or in the mycelium growing within the wood of the host tree. Type deposition — The holotype and two paratypes are deposited in the collection of the University of Michigan Museum of Zoology, Ann Arbor, MI, USA; one paratype is deposited in the collection of the Tyumen State University Museum of Zoology, Tyumen, Russia. All specimens are stored in ethanol with a drop of glycerol. Note: See Table 1 for explanation. Etymology — The species name parakontumensis refers to the similarity between the new species and Perscheloribates kontumensis Ermilov & Frolov, 2019. Remarks — In the presence of pointed rostrum, lanceolate bothridial head with long setiform apex and developed notogastral setae, and in the absence of prolamella, the new species is similar to Perscheloribates kontumensis Ermilov & Frolov, 2019 from Vietnam (see Ermilov & Frolov 2019a), but differs from the latter by the presence of simple notogastral setae (versus with flexible tip in P. kontumensis) and the absence of body sculpturing (versus simple in P. kontumensis) and setae it on leg tarsi I–III (versus absent in P. kontumensis). General remarks As previously noted (Norton 1980; Ermilov 2019; Ermilov & Frolov 2019a, b), some oribatid mites are actively phoretic on insects, having morphological adaptations for attachment to the host (for example, gripping setae of beetles between the rostrum of the aspis and the anterior portion of the genital plates in ptyctimous mites, or specifically curved leg claws for holding onto setae of the host as in Siculobata (Paraleius)) (Norton 1980). However, other oribatid species (including P. paracuriosus n. sp. and P. parakontumensis n. sp.) have no visible adaptations, so mostly hold the host’s surface by hiding in various depressions and grooves of the host body and using the force of the pretarsal claws for attaching to the surface. Beetles of the family Passalidae are one of the more common insect hosts actively used by oribatid mites for phoresy (Ermilov 2019), however, phoretic oribatid mites had not been previously reported from A. lamellatus, nor from any beetles of the family Zopheridae. Hence, our findings (P. paracuriosus n. sp. on A. lamellatus and P. parakontumensis n. sp. on P. obcordata) are the first records of the use of A. lamellatus and zopherid beetles by oribatid mites for phoresy.

Subgenus Perscheloribates (Perscheloribates) Hammer, 1973 Type species Perscheloribates clavatus Hammer, 1973, Published as part of Ermilov, Sergey G. & OConnor, Barry M., 2020, New Perscheloribates species (Acari, Oribatida, Scheloribatidae) phoretic on beetles (Insecta, Coleoptera), pp. 289-300 in Acarologia 60 (2) on page 290, DOI: 10.24349/acarologia/20204368, http://zenodo.org/record/4487761, {"references":["Hammer M. 1973. Oribatids from Tongatapu and Eua, the Tonga Islands, and from Upolu, Western Samoa. Det Kong. Dansk. Vidensk. Selsk. Biol. Skr., 20 (3): 1 - 70."]}

Two new species of oribatid mites of the genus Perscheloribates (Oribatida, Scheloribatidae), phoretic on passalid and zopherid beetles (Coleoptera), are described from the Philippines and U.S.A., based on adults. Perscheloribates paracuriosus n. sp. differs from Perscheloribates curiosus Ermilov, 2016 by the length and morphology of notogastral and adanal setae, the presence of lateral keel-shaped ridge, and the absence of custodium. Perscheloribates parakontumensis n. sp. differs from Perscheloribates kontumensis Ermilov and Frolov, 2019 by the morphology of notogastral setae and the absence of body sculpturing and setae it on leg tarsi I–III., Acarologia, 60, 289-300