Nanohystricidae n. fam., an unusual, plesiomorphic enarthronote mite family endemic to New Zealand (Acari, Oribatida) Author Norton, Roy A. Author Fuangarworn, Marut text Zootaxa 2015 4027 2 151 204 journal article 10.11646/zootaxa.4027.2.1 058b8bab-084b-4a45-8d74-29446fed1877 1175-5326 238764 B2D0A15A-16BE-4311-B9F1-ADC27341D40C Nanohystrix hammerae n. sp. Adult ( Figs 1–17 ). General attributes. Unusually large mites: total length 968–1213 Μm (smallest contracted male, largest distended female, respectively, not including setae). Average total length of females (n = 10) 1095 Μm (range 1029–1213 ), average maximum width 697 Μm (range 653–774), average maximum height 636 Μm (range 605–677); average total length of males (n = 10) 1036 Μm (range 968–1137), average maximum width 622 Μm (range 593–653), average maximum height 584 Μm (range 532–665). Sejugal band of soft cuticle ca. 100 Μm wide dorsally, ca. 20 Μm ventrally (exposed or hidden according to degree of distention and prodorsal flexing). Body shiny in reflected light. Cuticle mostly of three types . (a) Sclerotized cuticle conspicuously porose ( Fig. 7A, B ), brownish-orange in recently collected mature adults ( Fig. 1D ) but pale, brownish-yellow and somewhat translucent (eggs and food boluses visible within uncleared females) after long preservation ( Fig. 1A, E–H ); tubercles of notogastral setae appearing slightly darker than surrounding cuticle; legs often slightly paler than body. (b) Soft cuticle of most articulations: colorless, thin, highly flexible, usually with surface granules (probably cerotegument) of various density and shape, in places forming thick vermiform pattern ( Fig. 11A, D, E ). (c) Pale cuticle of apparently intermediate flexibility: with very faint pigmentation, fine and inconspicuous porosity, and in places with fine, dense surface striation ( Figs 10D , 11F, G ). Setae hyaline, lustrous and glassy in reflected light ( Fig. 1 ), strongly birefringent in polarized transmitted light ( Fig. 8A, E ). Most setae characterized below as: acicular if stiff, straight or slightly curved, tapering to thick point, heavily barbed (unless noted otherwise) almost to end; and attenuate if with long, taper to thin, smooth tip, with at least modest distal curvature and apparent flexibility. Barbs usually strongly developed and birefringent, like setal shafts. Prodorsum . Outline of main aspis semi-elliptical in dorsal aspect ( Fig. 2B ), seen flat about 1.2 times wider than long (e.g. 430 x 370 Μm). In lateral view, plane of aspis usually at 50–70° angle from horizontal ( Figs 3A , 5A, B ), maximum seen ca. 80°, minimum ca. 10° (great rotation allowed by ventrosejugal articulation and broad soft cuticle of dorsosejugal, disjugal and pleural regions). Aspis cuticle finely porose throughout ( Fig. 7A, B ) and uniformly colored except for eyes (see below); sigilla of cheliceral retractor and subcapitular levator muscles in several pairs of patches occupying most of posterior half ( Fig. 7A ). Aspis dorsally convex ( Fig. 1C ); anteriorly with slightly recurved rostral margin ( rm ; Fig. 6F ); margin continuing posteriorly on each side as sharply defined carina ( ca1 ), ending at lateral eye, thereby forming outline of aspis in dorsal aspect; with second carina ( ca2 ) descending from eye at right angle ( Figs 3A , 6A ). Lateral region between carinae concave ( Fig. 1C ; presumably relating to motion of leg I), subtriangular in lateral view, with irregular ventral margin merging with soft supracoxal region (i.e., no ‘prodorsal tectum’ present); with long, narrow extension forming supportive strut ( str ) with distal condyle that articulates with subcapitulum ( Figs 3A , 7D, G ). Rostral tectum very narrow, crescent-shaped; undulating vertically to accommodate chelicerae ( Fig. 6B ); reflected ventral part (rostrophragma) without noticeable solid limb; about 60 Μm deep medially, effacing laterally and ending at level midway between rostral and lamellar setae. Tectum anteromedially with low, well-circumscribed, pale rostral bulge precisely dorsal to median eye, ca. 50 µm wide ( rb; Figs 6C , 7H, I ). Prodorsum abruptly constricted posteriorly, forming near-vertical ‘occipital’ wall ( ow ; Figs 2B , 3A , 5C ), ca. 50 Μm high, edge of constriction bordered by carina ca2 (laterally) and setal pairs in , bo and xs (dorsally); wall with sharp posterior bend to form nuchal constriction ( nc ) that extends posteriorly for ca. 30 Μm before joining soft sejugal cuticle ( Figs 3A , 5C , 6A ). Nuchal constriction comprising posterior fifth of aspis; hidden in fully contracted specimens, in which occipital wall abuts notogaster, protecting sejugal articulation ( Figs 4B , 5D ). Median eye ( oc.m ; Fig. 7H, J ) about 50 µm diameter, located on rostrophragma directly under pale, dome-like rostral bulge ( rb ); with strongly convex but rather thin-walled cornea. No pigment observed in median eye, but sclerotized cuticle of rostrophragma thick and pigmented posterior to eye (in region of constriction st ). Paired lateral eye ( oc.l ) smaller, about 25 µm diameter, each immediately ventral to seta xi at junction of carinae ca1 , ca2 ; dark pigmentation visible beneath hemispherical cuticular cornea ( Fig. 7D ), lost with clearing of specimens ( Fig. 7A, C, G ). Prodorsal setae of various forms ( Figs 2B , 3A , 4B , 6A ), larger ones inserting on conspicuous tubercles. Rostral seta ( ro ) very thin, attenuate, ca. 170 Μm long, straight except for fine tip, projecting anteriorly, with only several inconspicuous barbs; pair slightly divergent, inserted close together (ca. 25–30 Μm mutual distance) on rostral bulge. Lamellar seta ( le ) inserted slightly proximal to mid-length of prodorsum, erect but moderately curved; relatively thick, attenuate, ca. 420 Μm long, with strong barbs in three longitudinal rows for much of length; pair widely spaced (ca. 255 Μm mutual distance). Remaining four pairs of prodorsal setae aligned at edge of occipital wall; all nearly erect, with slight posterior direction. Interlamellar ( in ) and upper bothridial setae ( xs ) larger than le and bearing up to five rows of barbs in proximal region: in about 570 Μm long, moderately attenuate; xs about 480 Μm, acicular, with strong barbs along whole length. Lower exobothridial seta ( xi ) small (about 50 Μm); acicular, with minute barbs visible only at high magnification. Bothridial seta ( bo , = ‘sensillus’) thinner than setae on either side ( in , xs ); about 500 Μm long, tapering to thin flexible tip, with 2–4 rows of barbs along whole length. Bothridium ( Figs 6D , 7E, F ) with simple conical form, no proximal curvature; outer half lined with approximately 20 concentric closely-spaced, fine circular ridges (some merging), which separate even finer honeycomb pattern; inner half abruptly narrower, with coarser ridges directed internally. Bothridial rim projecting slightly from surface; posterolaterally with tongue-like flange ( fl ; Fig. 6D ). Notogaster. Length about three times that of prodorsal aspis (excluding nuchal constriction); anterior width similar to that of prodorsum, about half that of maximum. Strongly but unevenly convex in both dorsal and lateral aspects ( Figs 1 , 2B , 3A ); maximum width and height at about level of setal row f ; curvature steeper in posterior third (more marked in fully distended individuals). Notogaster without marginal tectum; weakly folded laterally in anterior third (humeral crease; h.cr ), forming pleural concavity (more defined in contracted individuals; Figs 1D , 4D ); concavity effaces posterior to setal row e , pleural region becoming slightly convex posteriorly. General cuticle rather uniform and featureless, except as noted below; sclerotized cuticle with pores finer and less conspicuous than those of prodorsum. Well-spaced, minute surface granules in soft bordering cuticle, continuing onto anterior ( Fig. 6A , arrow) and lateral notogastral surface, becoming less dense in middle region of notogaster and absent posteriorly. Narrow transverse band of fine reticulation present in region of setal pairs c1 and c2 ( Fig. 6E , arrow). Notogaster and adanal plates fused posterolaterally, commencing at about level of seta ad2 ( Fig. 3A ; see Remark 9); narrow separating cuticle anterior to ad2 pale, finely porose, hidden in fold in contracted individuals ( Fig. 10B ). Dorsal curvature interrupted by five transverse formations ( Figs 2–5 ). (a) Shallow, simple transverse groove ( t.gr. ) immediately posterior to setal pairs c1 and c2 . (b) Deeper, broader transverse sulcus ( t.sul ) immediately posterior to setal row d . (c) Transverse type-S scissure bearing setal pairs e1 and e2 . (d) Transverse type-S scissure bearing setal pairs f1 , f2 and h2 . (e) Transverse type-S scissure bearing setal pair h1 . Setae of row d on modest tubercles within narrowly elliptical, irregularly-bordered region of lighter-colored cuticle ( Figs 1D , 2B ); central region of ellipse (around pair d1 ) pale, lateral regions around pair d2 intermediate in color. Edges of pale cuticle of type-S scissures generally regular anteriorly but irregular posteriorly. Erectile setae of type-S scissures each on large tubercle with associated muscles (see Discussion). Scissures differ in: setal spacing, presence and form of side extensions from posterior slope of tubercle, presence of incursions (from sclerite behind scissure) or excursions (of pale cuticle into posterior sclerite). Anterior scissure ( Fig. 2C ) with setae of row e almost evenly spaced; tubercles of pair e1 with medial extensions fusing to form narrow bridge; with small incursion of sclerotization between e1 and e2 (*), sometimes appearing like separate, intercalary sclerite. Middle scissure ( Fig. 2D ) with mixed setation, including two pairs of row f setae and seta h2 ; pair f1 widely separated; central third of scissure narrow, without setae or noticeable incursions or intercalary sclerites; tubercles of f1 and f2 each with basal extensions, separated by narrow articulation; tubercle of f2 also extended laterally, reaching near tubercle of h2 ; tubercle of h2 without extension; scissure with irregular triangular region of pale cuticle extending posteriorly from near setae f1 and f2 . Posterior type-S scissure ( Fig. 2E ) with only setal pair h1 ; each tubercle with short medial extension. With 19 pairs of glassy notogastral setae (including row ad ); some ( e1 , e2 , h2 ) extremely long, some ( f1 , f2 , h1 , p1 ) merging posteriorly in brush-like effect when held horizontally ( Figs 1A , 4A ). Long setae of various forms: acicular setae and proximal regions of large attenuate setae with four (sometimes five) straight rows of dense barbs, equally spaced around seta ( Fig. 4E ), forming imbricate ridges where barbs long and closely spaced ( Fig. 4G ); attenuate setae usually with two or three rows of more widely spaced barbs, at least in distal half ( Fig. 4F ). Setal shapes and representative lengths (to show relativity in average female; all significantly variable) as follows: c1 (170 Μm) finely attenuate; c2 (230 Μm) narrowly acicular; c3 (750 Μm) attenuate); cp (950 Μm) attenuate; d1 (970 Μm) attenuate; d2 (320 Μm) narrowly acicular, but curved; e1 (1900 Μm) attenuate; e2 (1750 Μm) attenuate; f1 (1200 Μm) acicular; f2 (1000 Μm) acicular; h1 (880 Μm) acicular; h2 (1350 Μm) attenuate; h3 (430 Μm) narrowly acicular; p1 (790 Μm) acicular; p2 (1300 Μm) attenuate; p3 (330 Μm) narrowly acicular; ad1 (1250 Μm) attenuate; ad2 (370 Μm) attenuate; ad3 (200 Μm) attenuate. Six large pairs, including e1 , e2 , f1 , f2 , h1 and h2 , inserting on intercalary sclerites within scissures (described above), apparently erectile. Several setal pairs with unusual locations ( Figs 2B , 3A , 9B ): c3 high on pleural surface, aligned with setal row d and immediately anterior to cp ; f1 somewhat displaced laterally; h2 uniquely incorporated as third erectile setal pair in middle transverse scissure; h3 more anterior and dorsal than usual in oribatid mites; p2 dorsoventrally aligned with p3 or even anterior to it. Typical five pairs of notogastral proprioceptors present, but in unusual locations ( Fig. 3A ); each minute, ca. 6– 7 µm, cupular in form, round or slightly elliptical when seen face-on ( Fig. 11G ). Two pairs located within scissures: im in pale or intermediately pigmented cuticle at lateral extreme of anterior scissure, closely posterodorsal to seta cp ; ip within middle scissure, between setae f2 and h2 , anterior to lateral extension of f2 . Three pairs in lateral region: ia and im on sclerotized cuticle ca. 30–50 µm from ventral edge of notogaster, ia about at level of cp or c3 , ih at level of middle scissure; ips in pale cuticle near anterior end of separation between main notogaster and adanal region ( Fig. 11F, G ). Five pairs of conspicuous cuticular disks present, two large, three small ( Figs 2B , 3A ; see Remark 2), discernable in both transmitted and reflected light by lack of color ( Figs 1D , 8B, C ); cuticle finely porose. Small disks all in anterodorsal region, usually slightly elliptical: cd1 smallest (ca. 15 µm length), about 30 µm posterior to seta c1 ; cd2 (ca. 30 µm) about 40–50 µm ventral to seta c2 ; cd3 (ca. 25 µm) about 60–70 µm posterior to seta c2 . Larger two disks circular to slightly elliptical (ca. 55–60 µm); cd4 located midway between erectile setae f2 and h1 at end of pale triangular extension from middle scissure; cd5 immediately posterolateral to seta h1 . Two linear files of tendon insertions present anterolaterally ( ti ; Figs 2B , 3A ). One file of 6–10 minute, uniform, darkly sclerotized, closely spaced internal projections directed posteriorly from near disk cd3 , each projection serving as insertion for one thin tendon ( Fig. 8C ). Second, less distinct row of insertions, perpendicular to first, directed dorsomedially in transverse sulcus behind seta d2 ; only most lateral few insertions of second row with internal projections. Venter. Coxisternum comprising about one-third of ventral surface ( Figs 9B , 10B, C ); nearly quadrate overall, with trochanter of each leg articulating at about same distance from midline. Muscle sigilla weakly defined in transmitted light. Paired plates of epimeres I and II strongly convex and independent both from other member of pair and from adjacent epimeral plates; soft cuticle medially separating plates of epimere I very broad, that of epimere II very narrow ( Fig. 11A ). By contrast, epimeres III and IV without soft medial cuticle (but with line of cuticular weakness visible in transmitted light; Fig. 9B ); epimeres fused to each other in medial half, forming single unit. Central fused region of unit relatively flat, but lateral half of each epimere discrete, cylindrical. Epimere III articulating with epimere II through narrow ventrosejugal articulation having weak transverse striation; posterior edge of epimere IV strongly curved, sharply defined, slightly overhanging genital plates and hiding broad articulation. Without coxisternal apodemes; epimere IV laterally with thickened internal ridge supporting articulation with leg trochanter. Coxisternal setation 3-1-3-4; no variation noted. Setae 1a , 2a , 3a and 4a straight, tapered only distally, conspicuously barbed ( Fig. 10E ); others attenuate to various degrees, with fewer barbs or nearly glabrous. Setae range in size from about 85 Μm ( 3a ) to 150 Μm ( 1b ). FIGURE 1. A–H Nanohystrix hammerae n. sp. , reflected light stereomicroscope images of adults (genders unknown; multiyear preservation except D). A, dorsal aspect of specimen with legs I, II angled posteriorly; B, ventral aspect of same; C, anterior third in posterodorsal aspect; D, dorsolateral aspect of recently collected specimen; E, dorsal aspect of notogaster; F, lateral aspect with legs I in walking position; G, lateral aspect (anterior half) of specimen with right leg I elevated; H, posterior aspect showing granules (arrows) of paired digestive caeca, seen by transparency. Abbreviations: h.cr (humeral crease); other labels indicate setae (see text). All composites of 10–20 layers. Scale bars: all 100 Μm. FIGURE 2. A–E Nanohystrix hammerae n. sp. , adult female, dorsal aspect. A, habitus of fully distended specimen (legs and some setae incompletely drawn for overall clarity); B, enlargement of same (larger setae abbreviated, see Fig. 9A); C, left side of anterior notogastral scissure (arrow at midline, setae indicated only by alveolus on basal tubercles); D, same view of middle scissure, showing extensive pale cuticle and disk cd4 ; E, entire posterior scissure and cuticular disks cd5 (posterodorsal aspect). Abbreviations: bng (border of notogaster), cd1–5 (cuticular disks), ia , im , ip (lyrifissures), nc (nuchal constriction), ti (tendon insertions); other labels indicate setae (see text). Scale bars: 100 Μm (C–E to same scale). FIGURE 3. A–D Nanohystrix hammerae n. sp. A, lateral aspect of fully distended female (legs omitted, some setae abbreviated or represented only by basal tubercles); B, sam e, genital region (with plate and genital papillae) and fully extended ovipositor, anterior to right; C, ovipositor, end-view (anterior to top); D, ventrolateral aspect of male genital region, showing spermatopositor in almost lateral aspect. Abbreviations: ca1 , ca2 (prodorsal carinae), cd1–5 (cuticular disks), cpc (podocephalic canal), ia , im , ip (lyrifissures), nc (nuchal constriction), ti (tendon insertions); other labels indicate setae (see text). Scale bars: 100 Μm (A, B); 50 Μm (C, D). FIGURE 4. A–G Nanohystrix hammerae n. sp. , scanning electron micrographs (genders unknown; most with debris artifacts). A, dorsal overview of contracted specimen with erectile setae depressed and legs I, II directed laterally; B, anterodorsal aspect of same (prodorsum fractured centrally), insert includes enlargement of cuticular disk cd1 , indicated by black arrow in main figure; C, dorsolateral aspect of hysterosoma of partially distended specimen with setae erected; D, same, enlarged, short arrows on cuticular disks cd5 (insert is enlargement of right disk cd5 showing exudate); E, base of seta e2 ; F, basal portion of seta d1 ; G, basal portion of seta f1 . Abbreviations; h.cr (humeral crease); other labels indicate setae (see text). Scale bars 100 Μm (A–D); 10 Μm (E–G). FIGURE 5. A–G Nanohystrix hammerae n. sp. , transmitted light compound microscope images of uncleared adult females, coarsely sectioned near sagittal plane (all lateral views, anterior to right). A, overview of fully distended specimen, with marked sections showing location of indicated close-ups; B, anterior third of fully contracted specimen, with close-up indicated; C, dorsal sejugal region, showing occipital wall and nuchal constriction of prodorsum, and extrinsic retractor muscles of chelicera; D, dorsal sejugal region and anterior region of notogaster, showing prodorsal retractor muscles ( pr.mu ; broken in C) inserted near border of nuchal constriction; E, posterior slope of hysterosoma in region of third scissure—at two focal planes, showing postanal suspensor muscle ( po.mu ) and deeper plane (right) with cuticular disk cd5 (with underlying fibers, fib ) and seta h1 (with depressor muscle, h1.mu ); F, section of middle scissure (more lateral than indicated in A) showing base of seta f1 and associated depressor muscle ( f1.mu ); G, ventrosejugal region, showing narrow ventral part of sejugal articulation and broad soft cuticle of epimere I. Other abbreviations: bng (anterior border of notogaster), cr.mu (cheliceral retractor muscles); cw (caecum wall), ep1–3 (plates of epimeres 1–3), nc (nuchal constriction), t.gr (transverse groove), sej (sejugal articulation), t.sul (transverse sulcus). All composites of 5–20 layers. Scale bars: 100 Μm (A, B); 25 Μm (C–G). FIGURE 6. A–G Nanohystrix hammerae n. sp. , scanning electron micrographs of adults (most with small surface debris artifacts). A, sejugal region of partly distended specimen, dorsolateral aspect (anterior to left)—arrow indicates minute surface granules (probably cerotegument); B, anterior third, ventral aspect, showing rostrum, gnathosoma and epimeres I–III; C, same, ventrolateral aspect, with chelicerae spread; D, bothridium and base of bothridial seta; E, portion of sejugal region of contracted specimen, anterodorsal view (cf. left side of Fig. 4B), showing reticulation (arrow) on notogaster; F, enlargement of C, right center; G, prodorsum, lateral aspect (slightly anterodorsal). Abbreviations: ca1 , ca2 (prodorsal carinae), cpc (podocephalic canal), Ch (chelicera), dc (dorsal carina of chelicera), ep1–3 (plates of epimeres 1–3), fl (flange of bothridial wall), Ge (gena), Ls (labrum), nc (nuchal constriction), oc.l (lateral eye), rb (rostral bulge), rm (rostral margin), sub.t (subcapitular tubercle); other labels indicate setae (see text). Scale bars: 50 Μm (B, C, G); 25 Μm (A, F); 10 Μm (E); 5 Μm (D). FIGURE 7. A–J Nanohystrix hammerae n. sp. , transmitted light compound microscope images of adults (cleared unless noted). A, right posterior region of prodorsum, dorsal aspect, showing muscle sigilla; B, close-up of general cuticular porosity; C, lateral eye, dorsal view; D, partial lateral aspect of prodorsum and supracoxal region, anterior to right—uncleared specimen showing pigment in lateral eye (insert shows eye from posterodorsal aspect); E, bothridium and base of bothridial seta, lateral aspect with focus near edge of cup; F, same, with focus at center of cup; G, similar to D, but after clearing; H, optical section of hyaline rostral bulge and median eye, lateral aspect, anterior to right (cf. Grandjean 1958b, his Fig. 2); I, anterior view of central rostrum and cheliceral bases, focused on rostral bulge; J, same, focused deeper, on median eye. All composites of 5–20 layers. Abbreviations: bo (bothridium), bt (base of rostral tectum), ca1 (prodorsal carina), Ch (chelicera), cpc (podocephalic canal), oc.l (lateral eye), oc.m (median eye), mu.sl (subcapitular levator muscle), rb (rostral bulge), pK (prodorsal condyle), st (constriction of tectum depth), str (supporting strut), sub.t (subcapitular tubercle); other labels indicate setae (see text). Scale bars: 20 Μm (A, D, G–J); 10 Μm (B); 5 Μm (C, E, F). FIGURE 8. A–E Nanohystrix hammerae n. sp. , transmitted light compound microscope images of adults (A, C uncleared). A, Anterior portion of notogaster dissected from body (midline indicated by thin black line; prodorsal retractor muscles torn anteriorly) under strong DIC illumination to show muscles; B, left cuticular disk cd4 and pale extension from middle scissure (posterodorsal view, anterior to right); C, region of cuticular disk cd3 showing tendon insertion files and proximal parts of thin tendons (lateral aspect, anterior to right); D, portion of seta f1 , transmitted bright-field illumination; E, same, but with polarized light. A–C composites of 5-15 layers. Abbreviations: cd1, cd3 (cuticular disks), e1.mu , f1.mu (muscles acting on respective setae), pr.mu (dorsal components of prodorsal retractor muscles), ti (tendon insertions), t.sul (transverse sulcus, with concave wall appearing brown in transmitted light; other labels indicate setae (see text). Scale bars 20 Μm (A); 10 Μm (B–E). Supracoxal seta eI ( Fig. 11B ) minute, spiniform (ca. 13 Μm), apparently immobile (no clear alveolus); inconspicuously located on dorsal face of epimere I at edge of sclerotization. Anterodorsal corner of epimere II with sharp spine or cluster of up to four smaller spines ( Fig. 11C ). Podocephalic canal ( cpc ; Figs 3A , 7D ) typical in form, running from supracoxal region of epimere I across soft pleural cuticle, then onto subcapitulum near articulation with prodorsum; often hidden in cuticular fold ( Fig. 7G ). FIGURE 9. A–B Nanohystrix hammerae n. sp. , adult female. A, several of the large setae that are abbreviated in B and in Fig. 2B; B, ventral aspect, legs and some setae only partially shown. Abbreviations: ips (lyrifissure); other labels indicate setae (see text). Scale bar: 100 Μm. Genital plates about 1.2 times longer than broad (seen flat), with gently curving outer margin; pale inner margin often reflexed ( Fig. 10A ); cuticle connecting genital and anal plates mostly soft, sharply folded, reminiscent of shirt-collar, medially with small patch of sclerotized cuticle (not illustrated) observed only with dissection. With nine pairs of smooth, attenuate genital setae (ca. 70–85 Μm long) arranged in two longitudinal rows: six setae aligned with medial margin, anterior pair directed ventromedially and others sharply posteroventrally; three other pairs near lateral margin of plate. Aggenital region soft, without evidence of plate or aggenital setae. Cuticle of genital vestibule soft, granulate; genital papillae of simple form (ca. 50 Μm diameter), all three pairs similar ( Figs 3B , 9B ), with lightly pigmented cap. FIGURE 10. A–E Nanohystrix hammerae n. sp. , scanning electron micrographs of adults, ventral aspect (asterisks denote artifacts; most images with unmarked small surface debris). A, anogenital region, peranal valves slightly opened; B, habitus of fully contracted specimen with legs I, II laterally directed, insert showing posterior region of more distended specimen with weaker imbrication of adanal region (cf. arrows); C, coxisternal region; D, peranal seta and soft, finely striated cuticle lateral to it; E, seta 3a . Labels indicate setae (see text). Scale bars: 100 Μm (B); 50 Μm (A, C); 10 Μm (D, E). Ovipositor with typical, double-walled tubular form and plicate cuticle ( Figs 3B, C , 12A, B ); about 500–650 Μm when fully extended, half that when folded inside body. Distal lobes normal in form: paired anterior (dorsal) lobes and larger single posterior (ventral) lobe, the latter with convex, slightly sclerotized inner face that coapts with anterior lobes in retracted ovipositors (cf. Fig. 12A, B ). With normal complement of nine pairs of simple, eupathidial setae, but with unusual distribution (see Remark 6). Three pairs of coronal setae ( k ; 40–50 Μm); two pairs within primary fold constriction, one well below on anterior side. Each anterior lobe with three setae: τ1 (ca. 70 Μm) largest and most distal; τa (ca. 40 Μm) most proximal, at limit of plicate cuticle; τb (ca. 40 Μm) at intermediate level and more laterally positioned. Posterior lobe with three pairs of setae: ψ1 (ca. 75 Μm) most distal; ψa and ψb (each ca. 35 Μm) at same level, near limit of plicate cuticle. FIGURE 11. A–G Nanohystrix hammerae n. sp. adults, transmitted light compound microscope images. A, mid-region of epimeres I and II, ventral aspect, showing granular soft cuticle; B, right supracoxal seta eI , anterolateral aspect; C, dorsal aspect (anterior to top) of left supracoxal region of epimeres I and II, showing spinous process at anterior border of epimere II (inserts show variation in process); D soft lateral cuticle above epimere I, showing cerotegument granules, some elongated, vermiform; E, same, enlargement of lower right corner, showing underlying fine striation and porosity; F, portion of incursion of pale cuticle lateral to adanal region of notogaster (cf. lower right side of Fig. 9B), ventral aspect, anterior to top; G, close-up of cupule ips ; H, shallow internal view of dissected (partial) anogenital region, showing preanal apodeme and genital plate muscles; I, same, but more internal focal plane, showing transverse compressor muscles. Abbreviations: ep1 , ep2 (epimeral plates), g.mu (genital plate muscles), ips (lyrifissure), pa (base of preanal seta), pr.a (preanal apodeme), pra.m (transverse preanal muscles); other labels indicate setae (see text). All composites of 5–20 layers. Scale bars: 50 Μm (H, I); 20 Μm (A, D); 10 Μm (C); 5 Μm (B, E–G). Spermatopositor ( Figs 3D , 12C ) very short (ca. 50–60 Μm), conical, but with weakly striate lobes discernable; ovate in cross-section at base (ca. 50 x 60 Μm). With eight pairs of small eupathidial setae (20–25 Μm), all tapered but blunt-ended; each anterior lobe with three at base ( k) and three distally (τ); posterior lobe with two pairs of setae (ψ). FIGURE 12. A–F Nanohystrix hammerae n. sp. adults, transmitted light compound microscope images except B (SEM). A, distal quarter of extended ovipositor with lobes spread, lateral aspect, anterior to right; B, similar, but distal lobes only slightly spread; C, spermatopositor, ventral aspect; D, lateral aspect of genu I solenidion σ1 and coupled seta d ; E, dorsolateral aspect of tibia I seta d (coupled solenidion removed); F, distal end of tarsus IV of male, showing potential for bending. All labels relate to setae (see text). Except B, all composites of 5–20 layers. Scale bars: 25 Μm (A–C, F); 10 Μm (D, E). Plates of anal region collectively forming vague trapezoid in contracted individuals, widest posteriorly ( Fig. 10A, B ). Seen in distended individuals, anal plates widely separated from peranal plates and adanal region of notogaster ( Fig. 9B ), mostly by pale, porose cuticle with inconspicuously linear pattern. Each plate about four times longer than wide (ca. 280 x 70 Μm); tapered anteriorly to about one-third maximum width. Anal setae aligned along midline of plate; all attenuate, barbed, increasing in size posteriorly ( an3 157 Μm, an2 270 Μm, an1 970 Μm); an2 closer to an3 than to an1 . Peranal plates extremely narrow, strap-like (ca. 300 x 15 Μm); medial edge soft, unpigmented, often flared outward. Plate pair fused anteriorly to form small ‘preanal’ apodeme for muscle attachment ( Fig.11 H, I); similar small ‘postanal’ apodeme posteriorly for attachment of suspensor muscles (see Discussion). Single pair of peranal setae small ( pa , ca. 60 Μm; Fig. 10D ), attenuate, weakly barbed; inserted on lateral edge near anterior end of plate. Gnathosoma ( Figs 13–15 ). Subcapitulum and chelicerae usually oriented vertically or nearly so ( Figs 3A , 5A ), even slightly posteroventrally in some individuals. Subcapitulum relatively small but unusually wide at base (ca. 1.2 times wider than long), abruptly narrowed by almost 50% at level of palp insertion ( Fig. 15A ). Stenarthric, ( Fig. 13A ): triangular mentum well delineated by pair of oblique labiogenal articulations ( lg ) closely approaching each other anteromedially, near base of lateral lips. Mentum ( H ) paler than genae; with one pair of attenuate, sparsely barbed setae ( h , ca. 80 Μm). Gena ( G ) without dorsal rasp-like region of denticles; each gena with three weakly barbed ventral setae: posterior two ( n , ca. 90 Μm; m , 80 Μm) nearly aligned with h , distal seta ( a , 90 Μm) about at level with anterior corner of mentum. Lateral lips ( LL ) relatively large, fully exposed in ventral aspect, each clearly separated from gena by postadoral articulation ( d ); distal halves tapering to narrow adjacent tips. Each lip with triangular sclerite on ventral face and three pairs of adoral setae, each having few, minute barbs; distal seta or1 (ca. 30 Μm) inserted in soft cuticle immediately distal to tip of sclerite, setal base distinctly flared, without normal alveolus; setae or2 and or3 (ca. 55 Μm) inserted on sclerite, at mid-length of lip, on small tubercles and in normal alveoli, in nearly transverse row. Dorsal surface with several longitudinal rows of extremely minute cilia or denticles, close to medial edge in distal half of lip. Rutellum ( Ru ) atelobasic; no rutellar brush; narrow, nearly uniform in width; distally rather chisel-like, with three aligned distal teeth. Rutellum basally incorporated smoothly with elongated manubrial zone ( Mn ) of gena; manubrial-rutellar unit angled rather sharply anteromediad, with rutellar tips nearly meeting just anterior to lateral lips ( Fig. 14B, D, E ); distal limit of manubrium ( c ) well-marked in transmitted polarized light due to strong birefringence of rutellum ( Fig. 15B ); abaxial fissure ( αf ) marking base of manubrium visible in both SEM ( Fig. 14B ) and transmitted light ( Fig. 15B ). Dorsal face of subcapitulum without capitular apodeme; posterior margin supported by pair of narrow transverse bands of sclerotization interrupted by U-shaped medial notch ( m.no ; Fig. 15A , insert). With flattened, sometimes pointed tubercle ( sub.t ) on each dorsolateral corner of subcapitulum, at articulation with prodorsal condyle pK ( Fig. 7G ). Postpalpal seta ( ep; ca. 8–10 Μm) spiniform, erect, inserted without alveolus (apparently immobile) just anterior to tubercle ( Fig. 15D ). Labrum ( Ls ; Fig 15B, C ) broad, with slightly raised medial ridge; distally rounded, extending almost to level of rutellar tips (if not elevated). Distal quarter of labrum with complex but slightly variable structure: with three transverse rows of dense spicules spanning plane of symmetry on dorsal face and fourth, most ventral row usually interrupted medially by larger distal spines, usually one pair ( Figs 14F , 15C ); ventral surface usually with several nipple-like lobes and proximal folds ( Fig. 14E ). FIGURE 13. A–E Nanohystrix hammerae n. sp. , gnathosomal elements of adult. A, central region of subcapitulum, ventral aspect; B, right palp, abaxial aspect (integumental granules shown only on basal segment); C, distal portion of palp tarsus with enlarged section of fused eupathidia; D, left chelicera, adaxial aspect; E. Right chelicera, abaxial aspect. Abbreviations: c.fd (carina of fixed digit), c.md (carina of movable digit), d (postadoral articulation), lg (labiogenal articulation), G (gena), H (mentum), LL (lateral lip), Ru (rutellum), sc (scale-like flap), tr.t (trochanteral tectum), Tr (trochanter), vb (vesicular band of chelicera); other labels indicate setae (see text). Scale bars: 50 Μm (D, E); 25 Μm (A, B); 10 Μm (C). FIGURE 14. A–I Nanohystrix hammerae n. sp. , scanning electron micrographs of adult gnathosomal and leg elements (all with small surface debris artifacts). A, overview of proterosoma, ventral aspect; B, subcapitulum and chelicerae, ventral aspect; C, lateral aspect of gnathosoma and rostrum; D, enlargement of top-central region of B; E, enlargement of labrum, ventral aspect; F, distal part of labrum, anterolateral aspect; G, distal region of palp, lateral aspect; H, leg apotele and distal part of tarsus, near lateral aspect; I, famulus of tarsus I, lateral aspect, with insert showing open tip of hollow barb. Abbreviations: αf (abaxial fissure), c.fd (carina of fixed digit), Ch (chelicera); lg (labiogenal articulation), Ls (labrum); Ru (rutellum); other labels indicate setiform organs (see text). Scale bars: 50 Μm (A); 20Μm (B, C); 10 Μm (G, H); 5 Μm (D–F, I). FIGURE 15. A–I Nanohystrix hammerae n. sp. adults, transmitted light compound microscope images of dissected parts of gnathosoma. A, dorsal aspect of subcapitulum and palps with enlarged insert showing medial notch in posterior margin ( m.no ); B, distal subcapitulum in strong DIC, dorsal aspect, showing birefringence of rutellum; C, same, but with focus on tip of labrum; D, right posterolateral corner of subcapitulum, lateral aspect (anterior to right), showing postpalpal seta and tubercle; E, chelicera, ventral aspect; F. chelicera, adaxial aspect, with insert showing lamellated organ; G, enlargement of ventrodistal region of cheliceral trochanter (cf. E), showing marginal spines and tectum with hyaline fringe (black arrow); H, proximodorsal region of chelicera showing porose cuticle with spines; I, chelicera in strong DIC illumination, focused internally to show levator tendon of movable digit and associated muscles. All composites of 5–20 layers. Abbreviations: αf (abaxial fissure), c (junction of manubrium with birefringent rutellum), ep (postpalpal seta), l.or (lamellated organ), Ls (labrum), l.mu (levator muscle), l.te (levator tendon of movable digit), Mn (manubrium), m.no (medial notch), m.sp (marginal spines), Ru (rutellum), sub.t (subcapitular tubercle), Tr (trochanter) tr.t (trochanteral tectum), vb (vesicular band). Scale bars: 50 Μm (E, F, I); 25 Μm (A); 10 Μm (B–D, G, H). FIGURE 16. A–G Nanohystrix hammerae n. sp. , adult legs, abaxial aspect. A, right leg I, tarsus partially shown; B, right tarsus I; C, enlargement of tarsus I famulus ( e ); D, tibia I solenidion φ1 with minute coupled seta d ; E, right leg II, tarsus partially shown; F, right tarsus; G, two variations of tarsus II famulus ( e ). Labels all indicate setiform organs (see text). Scale bars: 50 Μm (A, B, D–F); 5 Μm (C, G). Palp relatively large and conspicuous ( Figs 4B , 14A , 15A . Simple in form ( Fig. 13B ); five freely articulating segments with typical proportions: trochanter, genu and tibia short; femur about twice length of genu; tarsus elongated, about equal to femur and genu combined. Tarsus finely striated distally; with dorsal lyrifissure proximally. Setal formula (trochanter to tarsus) 0- 2 -1-3-13 (solenidion not included; tarsus count includes fused distal eupathidia as two); probable homologies indicated on Fig. 13B, C . All setae of femur, genu and tibia, and several proximal setae of tarsus, attenuate and noticeably barbed; more distal setae with fewer barbs, or none. Tarsal solenidion ω finely tapered, flexible; inserted unusually far distally, in terminal quarter of segment, extending slightly beyond most distal tarsal setae when straight. Homology of tarsal setation incompletely known (see Remark 10); most tarsal setae with soft cuticle of alveolus teardrop-shaped, pointed distally. Six distal tarsal setae eupathidial, including probable acm and ultimal ( ul ) pair ( Figs 13C , 14G ); ul' and su fused in basal 2/3, but individual setae distinguishable, spread noticeably in distal third. Variations: one specimen with femoral seta sup absent; another with seta inf doubled (represented by two closely-spaced, unusually small setae). Chelicera usually oriented vertically; chelate-dentate in form, with typical outline in lateral view ( Fig. 13D, E ). Cheliceral frame not encroaching onto principal segment; cuticle of latter distinctly porose proximal to chela. Trochanter ( Tr ) distinct, relatively large, well sclerotized ventrally; soft dorsally, merging with and losing distinctness from soft granular cuticle of cheliceral frame. Trochanter with tongue-like, well sclerotized tectum ( tr.t ; Fig. 15E–G ) ventrally, overhanging articulation with principal segment. Tectum with hyaline cuticular coating, extending distally as thin, colorless fringe with minute cilia; several large spines more basally on abaxial edge of tectum, distal to articulation with principal segment. Principal segment strongly asymmetrical, giving broad, flat-faced appearance to anterior of mite ( Fig. 6B, C, G ). Adaxial face almost flat, forming coaptive surface when chelicerae close together ( Fig. 6B ), accentuated dorsally by carina ( dc ; Fig. 6C ) bearing several short spines; abaxial face strongly convex, increasingly so as chelicera broadens proximally to maximum width, close to proximal end ( Fig. 15E ). Abaxial face of principal segment with few features: proximal third with about dozen minute, sharp spicules, many at end of fine, sharply defined carinae angling posterodorsad ( Figs 7J , 13E , 15H ); fixed digit with low carina ( c.fd. ) near middle ( Figs 13E , 14B ); movable digit with short carina ( c.md ) proximally, which coapts to edge of fixed digit if chela closed. Adaxial face of principal segment ( Fig. 13D ) deeply excavated in proximal half, accommodating asymmetrical insertion of trochanter; with about half-dozen spicules in distal half. Without Trägårdh’s organ, but with attached, irregular, ‘puffy’ (perhaps inflatable; Fig. 15E ) vesicular band of soft cuticle ( vb ) running distally from trochanter to end at strong spine (ca. 15 Μm) on mid-face of fixed digit. Small, freely projecting scale-like flap ( sc ) present, with ciliate margin, arising at about level of articulation between segments. Lamellated organ ( l.or. ; Fig. 15F , insert) present close to flap (see Remark 5). Fixed digit of chela strongly birefringent distal to seta chb ; with five teeth (including terminal tooth and small tooth high on adaxial face). Movable digit birefringent in distal half; with four teeth (including terminal and very weak subterminal tooth). Principal segment with two attenuate setae, both aligned near midline, just abaxial to dorsal carina ( Fig. 13D, E ): cha (ca. 65 Μm) clearly barbed, inserted at base of fixed digit; chb (ca. 40 Μm) nearly smooth, inserted at mid-level of fixed digit. Pronounced, often pigmented tendon ( ts ; Fig. 15I ) present, attaching bands of massive levator muscle to movable digit; articulation of digit with distinct onchophysis op' ( Fig. 13D ). Legs. ( Figs 16 , 17 ). Relatively long: leg IV longest, about as long as body (varies with degree of body distension); approximate proportions of others (relative to IV): 0.8 (I), 0.7 (II), 0.8 (III). Legs III and IV with normal, posterolateral direction in all specimens. Legs I and II unusually mobile (inferred from highly variable positions in preserved specimens); with normal anterolateral orientation in some specimens ( Fig. 1D, F ) but in others directed laterally, posterolaterally, or even posterodorsally (at nearly same angle as legs III and IV; Figs 1A, B, G , 10B ). Leg I, and to small extent leg II, rotatable crab-like; i.e. ventral surface twisted forward in some specimens ( Fig. 4B ). Surface of most leg segments nearly smooth, but tarsi finely striated distally ( Fig. 14H ); cuticle finely porose but without regions of larger pores (porose areas) or other surface patterns. Each tarsus with small proximodorsal lyrifissure. Pretarsus of all legs similar, tridactylous with empodial claw slightly longer and about twice width of lateral claws (14H); claws without denticles; ambulacrum without separate condylophores. Except for trochanters I and II and all tarsi, leg segments roughly tubular and similar in size ( Figs 10B , 16 , 17 ). Trochanters I and II relatively short, less than half length of respective femur. Tarsi much elongated; tarsus I about 6–7 times longer than proximal width, tapering gradually from base to tip; tarsus IV about 11–12 times longer than proximal width, with most tapering in proximal quarter, distal to which segment almost isodiametric, very thin and somewhat flexible (based on variations in preservation; Fig. 12F ); form of tarsi II and III progressively intermediate between that of I and IV. Leg setation, expressed as formula (I-II-III-IV; famuli and minute setae d included, solenidia excluded): trochanters (1-1-2-2); femora (6-6-4-4); genua (5-5-4-4); tibiae (6-6-4-4); tarsi (20-18-17-15). Setal homologies given in Figs 16 , 17 and Table 1. Noted variations include two tarsi I lacking seta it' and one tarsus II lacking seta tc ' (20 of each leg examined). Most setae moderately to strongly barbed, exceptions being distal setae of tarsi, on which barbs usually small and sparse as illustrated, and smooth eupathidial setae and regressive seta d of genu and tibia I (see below). As in body setae, with birefringent core extending into all barbs. Alveoli of tarsal setae dropshaped, pointed distally. Six setae eupathidial on tarsus I, including s , m" and pairs ( it ) and ( p ). Famulus ( e ) of tarsus I small (ca. 20 Μm) but conspicuous, being most proximodorsal setiform organ of segment and erect on small tubercle ( Figs 14I , 16C ); with attenuate main tip and usually three simple or forked barbs clustered near mid-length (rarely fourth barb present); tip and barbs all open distally ( Fig. 14I , insert). Famulus of tarsus II more distal, usually about mid-length on tarsus, smaller (ca. 10–12 Μm), strongly angled toward surface; variable in formattenuate to spiniform, sometimes with one small distal barb giving forked appearance ( Fig. 16F, G ). TABLE 1. Ontogeny of setae (Roman letters) and solenidia (Greek letters) in Nanohystrix hammerae n. sp. 1 1Setae and solenidia are first added at the indicated instar (La, Pn, Dn, Tn, Ad = larva, protonymph, deutonymph, tritonymph, adult, respectively) and remain present in subsequent instars; dash indicates no addition. Setae in parentheses represent pseudosymmetrical pairs, brackets indicate physical change; for seta d , subscripts m , n indicate that the seta is minute or normally formed, respectively.

Trochanter Femur	Genu	Tibia	Tarsus
Leg I
La	- d, bv"	(l), σ1, σ2	d, c", l' (v), φ	(ft), (pl), (pv), (tc), (a), (u), (p), s, m", e, ω1
Pn	v' -	d m, v"	-	(it), m', ω2
Dn-Tn*	- (l) (v)	v', σ3	l"	ω3
Ad	- -	-	-	-
Leg II
La	- d, bv", bx'	d m, (l), σ	d, c", l' (v), φ	(ft), (pl), (pv), (tc), (a), (u), (p), s, e, ω1
Pn	v' -	v" [d n]	-	-
Dn-Tn*	- (l), v"	v'	l"	(it), ω2
Ad	- -	-	-	-
Leg III
La	- d, ev'	d m, l', σ	d, l', (v), φ	(ft), pl', (pv), (tc), (a), (u), (p), s
Pn	v' -	v" [d n]	-	-
Dn-Tn*	l' l', v"	v'	-	(it), pl"
Ad	-	-	-	-
Leg IV
Pn	- -	-	v"	ft", (pv), (u), (p)
Dn-Tn*	v', l' d, ev', l', v"	d, l', (v), σ	d, l', v', φ	ft', (tc), (pl), (a), s
Ad	-	-	-	-

* Setae in these rows are present in the tritonymph, but as no deutonymphs were studied they may have been added in either the deuto- or tritonymph. Disjunctions of tarsal setae as follows (' = anterior member more distal; " = posterior member more distal; null = without noticeable disjunction): pair ( ft ) slightly ' on I, null on II, slightly " on III, IV; ( tc ) strongly ' on I, II, weakly ' on III, weakly " on IV; ( it ) ' on I, II, almost null on III; ( p ) null on all legs; ( u ) null on I–III, " on IV; ( a ) strongly ' on I, weakly ' or null on II, null to strongly " on III, IV; ( pl ) strongly ' on I–IV; ( pv ) strongly " on I–IV; ( m ) strongly " on I. See Remark 16 for variation in disjunctions and Remark 17 for setal pair ( m ). Solenidial formulas: genua (3-1-1-1); tibiae (1-1-1-1); tarsi (3-2-0-0). Solenidia of tarsus I and all genua long, finely tapered (‘piliform’ in terminology of Grandjean 1935b ), similar in length to most leg setae. Solenidia of tibia and tarsus II distally tapered, but not finely so (‘ceratiform’). Tibia I solenidion φ longest ( Fig. 16D ), flagellate (‘tactile’), about 2/3 length of leg I. Seta d and respective solenidia with various relationships on genua and tibiae ( Figs 16 , 17 ): seta d of genu I immediately adaxial to σ3, coupled in partially merged alveoli, minute (ca. 30–35 Μm), finely attenuate ( Fig. 12D ); d present but distantly proximal to solenidion on genua II–IV; d of tibia I similar to that of genu, closely coupled adaxial to flagellate φ ( Figs 12E , 16D ); d of tibia II well posterior to φ; d of tibiae III and IV closely coupled to φ but proximal to solenidion and in separate alveolus ( Fig. 17A, C ). Solenidia ω1 and ω3 of tarsus I inserted near dorsal midline, ω1 loosely grouped with famulus and seta ft" in basal quarter and ω3 located at about one-third segment length; ω2 with unusual distal position near level of iteral setae, almost ventral on abaxial face of segment. FIGURE 17. A–C Nanohystrix hammerae n. sp. , adult legs, abaxial aspect. A, left leg III, tarsus partially shown; B, left tarsus III; C, left leg IV (slightly twisted at base). Labels all indicate setiform organs (see text). Scale bars: 100 Μm. FIGURE 18. A–K Nanohystrix hammerae n. sp. juveniles: larva (A –I), protonymph (J, K). A, venter, legs partially shown; B, right leg I, abaxial aspect, tarsus simplified; C, right tarsus I, abaxial aspect, slightly rotated dorsally; D, right genu and tibia I, dorsal aspect; E, right leg II, abaxial aspect, tarsus simplified; F, right genu II, dorsal aspect; G, right tarsus II, abaxial aspect; H, left leg III, abaxial aspect, tarsus simplified; I, left tarsus III, abaxial aspect; J, right genu I, dorsal aspect (cf. D); K, right genu II, dorsal aspect (cf. F). Tibial seta d broken in H, I. Abbreviations: cd5 (cuticular disk), ih (lyrifissure), t.e. (vestige of leg IV); other labels indicate setiform organs (see text). Scale bars: 50 Μm (A, B, E, H to same scale); 25 Μm (C, D, F, G, I–K to same scale, except famulus enlargements in C, G). FIGURE 19. A–E Nanohystrix hammerae n. sp. tritonymph (appendages and many setae incomplete). A, dorsal aspect; B, close-up of right humeral region of hysterosoma; C. lateral aspect; D, close-up of proterosomal coxisternum region (cf. C), some epimeral setae represented only by alveoli; E, ventral view of hysterosoma. Abbreviations: cd1–5 (cuticular disks), cpc (podocephalic canal), ia , im , ip , ih, ips (lyrifissures), sp2 (dorsal spine of epimere 2), Pf (palp femur), pK (prodorsal condyle), Tr1 , Tr2 (trochanters of legs 1, 2); other labels indicate setae (see text). Scale bars: 100 Μm (A, C, E); 20 Μm (B, D). Ontogeny ( Figs 18–20 ) . Facies, size. All juvenile instars with general appearance of adult, including almost vertically oriented prodorsum, similar proportions of body regions, and presence of extremely long notogastral setae having proportions mostly similar to those of adult ( Fig. 20A–E ). Differ from adult in being mostly pale, with soft cuticle covered in fine granules; sclerotization and pigmentation more localized and more weakly developed than in adult, but still with fine pores. Body length (n = 1; not including setae or chelicerae) of larva 335 µm, protonymph 560 µm, tritonymph 900 µm (deutonymph not measured). FIGURE 20. A–I Nanohystrix hammerae n. sp. juveniles, stereo- (A–C) and compound (D–I) light microscope images. A, larva with all legs in walking position, lateral aspect; B, larva, anterior half in dorsal aspect, with right legs I, II; C, deutonymph with legs I and II laterally directed, dorsal aspect; D, tritonymph, left anterolateral region of hysterosoma showing muscles, cf. Fig. 8A; E, tritonymph, lateral aspect (filled with food boluses and gregarine parasites, see Fig. 22G),with insert showing enlargement of prodorsal strut; F, tritonymph, seta e1 and cuticular disk cd4 , dorsolateral aspect, anterior to left; G, protonymph, distal part of tarsus III, lateral aspect; H, tritonymph, distal part of tarsus III, dorsal aspect; I, protonymph, left leg IV, abaxial aspect (seta ft" torn away). Abbreviations: cd4 (cuticular disk), e1.mu (depressor muscles of seta e1 ); pr.mu (prodorsal retractor muscles), str (prodorsal strut); other labels indicate setae (see text). D–I composites of 5–20 layers. Scale bars: 100 Μm (A B, E); 25 Μm (D, F–I). Scale not determined for C, probably similar to A. Prodorsum. Aspis pale, sclerotization hardly discernable in early instars, gradually becoming more noticeable and lightly pigmented in deuto- and tritonymph (cf. Fig. 20A, C, E ). Structured as in adult, except vertical occipital region gradually decreasing in degree of sclerotization from top, and without nuchal constriction ( Fig. 19A, C ). Without lateral carina ca2 ; sclerotization in region below ca1 hardly discernable, except for strut ( st = ‘lateral nervure’) ending in strong condyle pK ( Fig. 20E , insert). Eyes as in adult except pigmentation not observed in lateral eyes (possibly lost in specimen preparation). Prodorsal setae generally similar in form and distribution to those of adult. Gastronotic region . Simple in structure, without general sclerotization or transverse scissures; noticeable sclerotization limited to tubercles at base of individual setae ( Fig. 19A–C ); in nymphs only setae c3 and cp on same sclerite (not examined in larva). Soft cuticle expansive and finely granulate. Cuticular disks well delineated: in tritonymph similar in size and distribution to those of adult; not studied in other instars, except posterior two large disks present in larva with same relative position as in adult. Distribution and form of setae generally similar to those of adult, except f1 , h2 , h3 , p2 with more normal positions relative to surrounding setae (not laterally or anteriorly displaced; cf. Figs 2B , 3A ); seta e1 proportionally longer in larva ( Fig. 20A ) than adult (2 times vs. 1.6 times body length, respectively). Setae e1 , e2 , f1 , f2 , h1 , h2 apparently erectile (held horizontally in larva but erect in studied nymphs; cf. Fig 20A, E ); tritonymph with muscles attaching to tubercles of presumed erectile setae in same pattern as in adults ( Fig. 20D ; see Discussion). Files of sclerotized tendon insertions absent. Coxisternal region. Incompletely studied. Larva ( Fig. 18A ) with weakly defined epimeral plates I–III similar in form to those of adult: epimere I with wide separation between paired plates, epimere II with narrow separation, epimere III without clear separation. Vestige of leg IV ( t.e. ) present posterior to epimere III, in form of dome-like swelling (see Remark 12). Coxisternal setation (I to III) 3-1-2; seta 1c modified as scale covering Claparède’s organ; 1a and 3a shortest, 3b longest; most with noticeable barbs, those of 3a minute. Tritonymph ( Fig. 19E ) with epimeres structured as in adult but more weakly sclerotized; setation as in adult (3-1-3-4). Supracoxal region as in adult, except seta eI proportionally larger in larva ( Fig. 18B ); spine of epimere 2 present in tritonymph ( sp2 ; Fig. 19D ; not studied in other instars). Anogenital region . Ontogeny of genital papillae normal; larva without features in presumptive genital region; protonymph with single pair of genital setae on valves; tritonymph ( Fig. 19E ) with seven pairs, three in lateral row, four in medial row (posterior two absent); not examined in deutonymph. Larva ( Fig. 18A ) with four pairs of pseudanal setae: p2–p4 similar to epimeral setae in size but with fewer barbs, p4 shortest; p1 large, diverging distally, longer than pseudanal valves; seta p4 (inguinal seta) lost in protonymph. Larva without inguinal seta of h - row; h3 well formed. Cupule ih close to pseudanal valve in larva. Anal region of tritonymph similar to adult except for lack of noticeable sclerotization; peranal segment fully formed as well-delineated paraproctal valves; with small seta pa and small preanal and postanal apodemes extending inward from ends of peranal valves and with similar attached muscles. Gnathosoma. Subcapitulum stenarthric in all juvenile instars; seta n of gena added in deuto- or tritonymph; adoral seta or3 added in protonymph. Palp seta inf added in deuto- or tritonymph; setation of other segments similar in all post-larval instars, except in protonymph (and probably larva) terminal eupathidia ul', su not fused as far distad as in adult. Chelicera as in adult except for generally lesser sclerotization. Legs . Pretarsi of all legs monodactylous in larva and protonymph ( Figs 18 , 20G, I ), tridactylous in tritonymph ( Fig. 20H ), not recorded for deutonymph (see Remark 13). Setal ontogeny (unknown for deutonymph) given in Table 1. Larva with femur II seta bx' present (see Remark 21); famulus I with single barb. Genu ontogeny unusual (see Remarks 23–25): seta d absent from genu I of larva ( Fig. 18D ), forming as minute seta in protonymph ( Fig. 18J ), but distant from any solenidion until formation of σ 3 in deuto- or tritonymph; d of genua II and III minute in larva (Fig, 18F), becoming normal, barbed in protonymph ( Fig. 18K ) but proportionally smaller than in adult, reaching full adult proportion by tritonymph. Only proral setae ( p ) eupathidial in larva; s becoming eupathidial in protonymph; m" and iteral pair ( it ) normal in protonymph, becoming eupathidial in deuto- or tritonymph. Leg IV of protonymph ( Fig. 20I ) with lyrifissure absent (otherwise present on all tarsi), tibia with v" present, tarsus with typical set of seven setae: ( p ), ( u ), ( pv ) and ft" ; other segments glabrous (see Remark 22). On leg I, solenidion ω2 of tarsus appearing in protonymph at same distal location as in adult; genu solenidion σ3 forming in deuto- or tritonymph, coupled to seta d . Observed variations all on leg II of tritonymph (n = 3 specimens ): (1) seta l" absent unilaterally from tibia of one specimen; (2) famulus proximal to setae ( ft ) in one specimen. Material examined . Type series. Holotype : adult female, deposited in NZAC ; New Zealand , North Island, Waikato District, nr. Limestone Downs, ~ 37°27'S , 174°46'E (Waikato Wind Farm, C-block); C.H. Watts, col., 28- I-2010 , ex: pitfall trap on floor of native forest. Paratypes : 51 adults with same collection data as holotype except dates 21-XII-2009 through 28-I-2010 (13 deposited in NZAC , four in ANIC , four in CNC , 15 in CUMNH , 15 in RAN). Preserved in alcohol, except 10 paratypes dissected on slides (in RAN). Other studied material. Northland Region . Four adults (in NZAC ) from Te Paki Ecological District; Radar Bush, ~ 9.5km, SE of Cape Reinga, ca. 34°28'S , 172°46'E ; O. Ball, col., IX-2006 to III-2007 , ex: pitfall traps in native shrubland. Three adults (in FM), one tritonymph (partly studied, lost), one deutonymph (lost), one protonymph and one larva (partly studied, lost) from Waipoua Forest (Toronui Track), 150 m a.s.l.; A. Newton and M. Thayer, col., 13-IV-1980 , ex: decaying nikau palm leaf bases and litter. One adult (in CNC ) from Waipoua Kauri Forest; L. Masner, col., 12-XII-1983 , ex: sweeping low vegetation in kauri forest. Auckland Region. Twentysix adults ( 21 in NZAC , five in RAN) from vicinity of Laingholm, ~ 36°57'S , 174°38'E ; J.T. Pusateri col., dates ranging from 19-V-2007 to 23-I-2008 , ex: pitfall traps in semi-native forest. One adult from Huapai Scientific Reserve; 6-XI-2009 , S.E. Thorpe, col.; ex: leaf litter in native forest. Waikato Region. Three tritonymphs from type locality (one in CUMNH , two in RAN); 28-I-2010 . Fourteen adults from Te Akau, Waikato Wind Farm H-block ( 11 in NZAC , three in RAN), ~ 37°40'S , 174°50'E ; C.H. Watts, col., 21-VII-09 and 13-I-2010 , ex: pitfall traps in native forest. Etymology . The genitive species name honors the memory of Dutch soil biologist Marie Hammer ( 1907– 2002 ). Her pioneering studies greatly expanded the geographic and taxonomic horizons of our knowledge of oribatid mites, especially those of the southern hemisphere ( Balogh 1985 ). In particular, her three large publications on New Zealand oribatid mites ( Hammer 1966 , 1967 , 1968 ) provide a firm foundation for continuing studies of this interesting fauna.