Makarovaia ornata Moraza & Lindquist, 2015, new species
Creators: name: Moraza, María L.
name: Lindquist, Evert E.
Publisher: name: Zenodo
Year: 2015
Subjects: Biodiversity
Taxonomy
Animalia
Arthropoda
Arachnida
Mesostigmata
Ascidae
Makarovaia
Makarovaia ornata

description: Makarovaia ornata new species
(Figures 1–40, 98–100)
Diagnosis. Adult female and male: Dorsal shield reticulated marginally, with punctate central region, and with 18 pairs of smooth, similar, moderately short setae, except minute J5 barbed basally (Fig. 4). Peritremes extending to posterior margin of coxae II in female and anterior border of coxae III in male. Adult female: Sternal shield lateral remnants reticulated, as long as epigynal shield; epigynal shield drop-shaped, posteriorly rounded, with elongate reticula; anal shield wider than long. Peritrematal shield fragmented, with anterior free platelet bearing gp1 at level of coxae II. Fixed cheliceral digit with 9–11 teeth along masticatory surface, movable digit with five or six teeth. Adult male: Sternitigenital shield fully reticulated, with five pairs of setae; anal shield and opisthogastric setae as in female. Chelicera with paraxial arthrodial process at base of movable digit finely ciliate; fixed digit with eight teeth; movable digit with two minute teeth; spermatodactyl as long as middle article (including fixed digit) of cheliceral shaft. Nymphs: Opisthosoma normally developed, not reduced in form and size; legs IV extended posterolaterally, inserted normally, far from posterior margin of opisthosoma.
Description. Adult female. Idiosomatic dorum. Idiosoma oval, 775–850 long, 450–500 wide; dorsal shield 462–500 long, 309–335 at its greatest width at level of setae r3 (n = 7); shield with nearly parallel sides, reticulated laterally, changing to transverse lines anteriorly between setae j1 and j3, and densely punctate elsewhere; glandular poroids at level of setae Z3 conspicuously captured by horn-like extensions of shield. Dorsal shield with 18 pairs of smooth, similar, moderately short setae, except basally spiculate, minute J5 (Fig. 2); setal lengths: j1 (15–20), j3–j6, z2, z4 (17–23), z5, s4 (16–18), s1 (10–13), r2 (12), r3 (17), J4 (18–20), Z2, Z3, Z4 (14–16), Z5 (20–25), clunal J5 (4–6) (Figs 1, 4). Lateral soft cuticle with setae s5, s6, r5, S1–S5, R1–R6 and several UR’s 17–21 long. Peritrematal shields free from dorsal shield anteriorly.
Idiosomatic venter. Tritosternum with base variably but similarly as long (25–30) as wide (20–28), with laciniae free for about 90 percent of total length (98–112 excluding base), lacineae scarcely pilose (Fig. 11). Presternal area weakly transversely lineate (Fig. 5). Sternal shield with lateral elements reticulate and densely punctate; desclerotised region between elements weakly lineate; lateral element with anterior margin lobulate at base of sternal setae st1, posterior margin irregularly convex posterior to iv2, paraxial lateral margin with small notch between setae st1–st2, and adaxial lateral margin with elongated, free, narrow endopodal extension between coxae I–II bearing gvb, and with well developed, small endopodal extension between coxae II–III; narrowest width of “entire” sternal shield between legs II (87–112), of which surfaces of both lateral sclerites constitute about 40%; single sclerite length 109–128, narrowest width between legs II 15–25; sternal shield setae st1 (33–37) slightly shorter than st2 (37–43) (Fig. 100). Setae st3 (35–39) and st4 (30–40) on soft cuticle, gland pore gv1 and poroids iv3 absent. Endopodal strips well developed alongside coxae III and IV. Epigynal shield ornamented with elongate reticula, 130–142 long, 44–52 at widest region at level of setae st5, 18–23 at narrowest middle region, with rounded posterior margin; setae st5 similarly as long as other sternal setae (33–41). Anal shield reticulated, with broadly rounded anterior margin; greatest width (104–122) at level of gv3, about 1.3 its mid-length (84–95); gland pores gv3 at level of posterior border of anus; postanal seta (19–22) as long as para-anal setae (19–22); posterior shield margin throughout with thick cribrum (10–15). Soft cuticle with about nine pairs of opisthogastric setae of similar length, JV1 (19–25), JV2 (18–20), JV3–JV5, ZV2 (16–18) slightly thinner than other ventral setae, ZV3–ZV4 (18–19); these setae flanked by several pairs of posteriormost setae of UR - and R -series. Anterior fragment of peritrematal shield bearing gp1 between coxae I and II; main posterior part of shield consolidated with exopodal strip curving behind coxa IV, and bearing poroid ip2 and gland pore gp 2 in the middle, and poroids ips and ip3 and gland pore gp3 behind stigma; peritremes reaching posterior margin of coxae II (121–135) (Fig. 5). Spermathecal apparatus with small, curved tubular proximal section (curved length ca. 25, tubular width 2), widened distally (width nearly 10), discernible within body between coxae III and IV (Fig. 3), solenostome located slightly paraxially from between bases of coxae III and IV.
Gnathosoma. Gnathotectum (Fig. 2) with dorsal face convex, sparsely lineate, with prominent convex, serrated anterior margin. Cheliceral shaft, excluding basal section 116–130 long, with moderate sized digits (Fig. 7); dorsal face of fixed digit with basally widened dorsal seta (Fig. 6); fixed digit with offset subapical tooth by apical hook, with 9–11 teeth along masticatory surface; movable digit (45–50) with five or six teeth. Corniculus moderately short (21–27), stout (width 9–11 where inserted), with acuminate paraxial process (17–22), extending beyond main tip of corniculus (Fig. 10); corniculi inserted at level of ca. 20 µm posterior to insertions of setae hp1. Internal malae wide basally, bifid, tapered, long (50–64 from base of hp1) extending beyond tip of setae hp1 and with surfaces finely pilose; salivary styli tapered (Fig. 9). Labrum long, (60–68 from base of setae hp1), extending to palptibia. Subcapitulum with eight multidenticulate (6–8 denticles each) rows of deutosternal denticles. Subcapitulum with setae hp1 conspicuously thicker and longer (44–52) than other setae; hp2 (17–22) and pc (23–30) shorter and thinner than hp3 (30–40). Palpus length (145–155); palptrochanter with inner seta longer (28–32) than outer seta (12–20); palpfemoral seta al and palpgenual setae al–1 and al–2 relatively short (11–12) with tapered tips (Figs 8 a–c).
Legs. At level from femur to tibia, respectively, legs I as wide as legs IV; widths of femur and tibia: leg I 55 –76, 46–60; leg II 65 –90, 50–69: leg III 65 –85, 55–64; leg IV 60 –75, 48–60. Legs I to IV (Figs 12–15) each clearly shorter than dorsal shield length; legs I slightly shorter than legs II–IV, lengths, excluding pretarsi: I 360–380, II 372–405, III 374–410, IV 379–430. Leg I tarsus (91–102) about 1.7–1.8 longer than each of the femur (57–63), genu (52–60), and tibia (53–58). Tarsus I with nearly truncate distal margin, pretarsus (15–23 to base of claws) with large claws (19–23) and pulvillus; tarsus I without markedly elongated setae apically (Fig. 19). Legs II to IV with tarsus about 1.8–2.1 times as long as tibia. Tarsi II–IV with pair of apical setal processes ad -1, pd -1 (10–14) inconspicuous, shorter than length of pretarsus to base of claws (18–27); tarsal ventral projection truncate apically, with small central point (11–13) (Figs 16, 17); pretarsi with paradactyli inconspicuous as narrow sclerotised shafts flanking, and about as long as claws (19–23). Tarsi II–IV with ventral setae av -1, pv -1 similar in length to attenuated setae al -2, pl -1, and at least twice as long as other ventral setae (as in Fig. 17); av -2, pv2 similar in shorter length and thicker shape to al -1 (18), and mv with bulbose base and slightly shorter. Coxae I–IV reticulate on ventral surfaces; coxal setae simple, attenuate (Fig. 5), coxa I seta av (20–22), pv (16–18); coxae II–III av (ca. 25), pv (ca. 22); coxae I–IV with serrate posterior distal margin, and a second ventral serrate ridge parallel to that on coxa I; trochanter I with a dorsal serrate ridge; trochanters I–IV with few denticles on basal margin. Legs I to IV with chaetotactic formulae of segments as described for genus. Leg segments with several setae on small tubercles and thickened, spine-like setae as described for genus (Figs 12–15). Seta pv -2 on trochanters I–IV and av - 1 on trochanters II and IV (Figs 12–15) attenuate but not markedly elongated.
Adult male. Idiosomatic dorsum. Idiosoma oval, ca. 600–675 long, 450–500 wide; dorsal shield 465–600 long, 350–437 wide at level of setae r3 (n = 4); shield with ornamentation and complement of pore-like structures similar to that of female, but slightly more oval in shape, and slightly more widened anterolaterally to include anterior fragment of peritrematal shield bearing gp1 above coxae I and II. Dorsal shield with 18 pairs of setae; form and lengths of setae as in female, including reduced size and form of J5 (4–6). Lateral soft cuticle thickened as in female, usually with four pairs of setae UR. Peritrematal shields free from dorsal shield anteriorly (Fig. 21).
Idiosomatic venter. Tritosternum much as in female, with shorter base (length 14–20, width 18–24) and laciniae free for about 90% of total length (82–96). Presternal ornamentation as in female. Sternitigenital shield conspicuously reticulate (Fig. 25). Anterior margin of sternitigenital shield lobed on either side of genital opening (Fig. 25); total shield length 318–337, narrowest widths 99–112 and 90–109 at levels between coxae II and coxae IV respectively; lengths setae st1 (28–34), st2–st3 (35–37), st4 (34), and st5 shorter (26–28). Anal shield 123–130 wide at level of gv3, and 104–112 long, ornamented as in female; postanal seta (14–18) nearly as long as para-anal setae (16–19). Soft cuticle with nine pairs of opisthogastric setae similar in length (13–19) and shape as in female. Form of peritrematal and exopodal shields posteriorly as in female, peritremes 149 long (Fig. 25).
Gnathosoma. Gnathotectum similar in form to female (Fig. 22), 90 long, 118 wide. Cheliceral shaft, excluding basal section, 117–125 long; fixed digit with offset subapical tooth by apical hook followed by eight or nine moderate sized teeth, pilus dentilis indicated only by alveolar remnant; movable digit (39–40) with apical hook and two small teeth; spermatodactyl long (112–125), ca. 7 wide at its base and ca. 4 at its apical portion, extending posteroventrally from base of cheliceral shaft (Figs 23, 24); paraxial arthrodial process finely ciliate (20). Corniculi (25–27) more widely spaced basally, with inner acuminate process (22–23) thinner basally than in female (Fig. 27); internal malae as in female. Subcapitulum (Fig. 28) with seven rows of deutosternal denticles, form and relative lengths of setae much as in female, e.g. hp1 (41–45) prominent, hp2 (13–15) shortest. Palpi (length ca. 125) similar in structure as in female.
Legs. Relative leg thicknesses and lengths, excluding pretarsi, similar to those of female: I 367–395, II 401–425, III 420–480, IV 442–480. Legs with segment length ratios much as in female, tibia and genu I 55 –62 and 57–62 respectively, but femur I longer (65–73). Tarsus I (77–90) bearing pretarsus (13–17 to base of claws), claws 20–25. Tarsi II–IV with pair of apical setal processes ad -1, pd -1 (14–15), apical ventral process under pretarsus truncate (13–15), apex without central point present in female (Fig. 18), pretarsus (20–27) and pretarsal structures formed as in female, claws 20–25. Scant male dimorphism discerned among leg setae, except for ventral setae and al -1 on tarsi II–IV more thickly spinelike with a bulbous base (Fig. 18).
Deutonymph. Idiosomatic dorsum. Idiosoma broadly oval, length 458– 512 female, 539 male, width 320– 365 female, 445 male, dorsal shielding not clearly delineated, without lateral incisions (Fig. 29); dorsal shielding similarly ornamented as in female in matured specimens, but with fainter rugose cuticle when young. Most dorsal setae short (7–20), clunals J5 barbed basally and minute (3–5); central setae j1 (9–14), j3 (12–17), j4, j5, j6 (15–17), J4 (11–15); z2 (13–14), z4, z5, s4 (12–20), s5–s6 (16), s1, r2–r3 (10–11), Z2 (16–19), Z3–Z4 (16), Z5 (15–20); other peripheral setae on soft cuticle S, R and UR (5–14).
Idiosomatic venter. Tritosternum similar in form and extent of laciniae fusion to that of adult female, base length (29–30), lacinia with 13–14% of length (68–74) fused. Presternal area bare. Sternal shield with four pairs sternal setae st1–st4 and two pairs of poroids (Fig. 30), length ca. 275, width 123–140 at level of coxae II; endopodal extensions distinct; sternal setae st1–st3 similar in length (16–30), st4 shorter (8–13); setae st5 on soft cuticle, minute (5–8). Opisthogastric region length, from insertion of st5 to posterior margin of idiosoma, ca. 0.3 idiosomatic length. Anal shield poorly sclerotised, with thick cribrum as in adults; greatest width (88–100) of shield at level of postanal seta similar to midline length 90–99; postanal seta (6–8) slightly shorter than para-anal setae (9–10). Opisthogastric soft cuticle with at least nine pairs of setae (JV1–JV5, ZV2–ZV5), JV1 (9–11) longer than JV2, and JV5, ZV2–ZV5 minute, spine-like (5–7) (Fig. 30). Metapodal plates absent. Peritremes short (35–50 including stigma), reaching to mid-level of coxae III. Peritrematal shield poorly developed and not including peritrematal poroids which are on soft cuticle.
Gnathosoma. Gnathotectum (100 long, 110 wide), chelicerae (in female deutonymph), ventral gnathosomatic structures and palpi (107–123 long) as in adult female. In male deutonymph, cheliceral shaft, excluding basal section, 120 long, with movable digit (37–39) flattened (12–14 wide) and with sperm duct primordium, one minute tooth and minute distal hook discernible (Fig. 40); cheliceral dorsal seta short (9). Female deutonymph cheliceral shaft 106, movable digit as in adult female (37–42).
Legs. Relative leg lengths, excluding pretarsi, similar to those of female: female DN, I 336–350, II 341–350, III 354–360, IV 361–362; male DN, I 357–362, II 377–393, III 371–395, IV 361–388. Tarsus I with pretarsus ca. 10–15 to base of claws (ca. 18). Tarsi II–IV with pair of apical setal processes ad -1, pd -1 (12–16) similar to length of pretarsus to base of claws (14–15); with apical ventral projection short (8–12), spade-shaped (Fig. 26). Leg I segment length ratios and legs I–IV coxal structures and relative lengths and form of leg setae as in adult female. Trochanters I–IV with seta pv -2 slightly attenuated.
Protonymph. Idiosomatic dorsum. Idiosoma ovoid in shape, 500–575 long, ca. 400–500 wide (n = 3); dorsum poorly sclerotised (Fig. 32), without clearly delineated shields and with rugose, faint reticulation laterally, similar to that of larva and young deutonymph. Dorsal podonotal setae slightly longer than opisthonotal setae: j1 (9–10), j3–j5 (8–13), j6 (20), z2, z4, z5, r2, r3 (17–18), J4, Z2–Z4, S3–S5 (10–11), clunals J5 minute, thorn-like, basally spiculate (3–4), and other marginal setae short (5–6).
Idiosomatic venter. Tritosternum similar in form to that in deutonymph and adult female, but with length of base (14–16) less than width (22–26), and laciniae free for about 75% of total length (110–116). Presternal area bare. Sternal shield (ca. 260–300 in length), faintly sclerotised, reticulate, with setae st1–st3 (15–17) and poroids iv1, iv2. Setae st5 minute (3–4), on soft cuticle between bases of legs IV (Fig. 33). Anal shield faintly ornate, slightly elongate in form as on deutonymph (length 85–110, width ca. 80–100); gland pores gv3 on posterolateral corners of shield, poroids ivp on adjacent soft cuticle; postanal seta (9–10) slightly shorter than para-anal setae (10–13); cribum not only similar in width (ca. 90) to anal shield as in subsequent instars but thicker, extending from posterior shield margin to insertion of postanal seta. Opisthogastric setae on soft cuticle short (3–4). Opisthogaster length, from setae st5 to posterior margin of idiosoma 160–185, ca. 0.3 of total idiosomatic length. Peritremes short (ca. 32–35, including stigma) extending anteriorly to posterior margins of coxae III; peritrematal plate hardly discernible, with peritrematal poroids and gland pores on soft cuticle (Fig. 33).
Gnathosoma. Gnathotectum similar in form to subsequent instars, but dorsal face unornamented. Cheliceral shaft, excluding basal section, 94–105 long; fixed digit with offset subapical tooth followed by row of seven teeth; movable digit (30–35) with four teeth and a ventral proximal protuberance (Fig. 39). Other gnathosomatic structures, including inner acuminate process of corniculus and seven rows of deutosternal denticles (four to six denticles each), similar to those in subsequent instars; corniculi inserted at level of ca. 10 µm posterior to insertions of setae hp1 (Fig. 37 b). Palpi (length 105–115) with normal protonymphal complement of setae.
Legs. Leg lengths, excluding pretarsi: I 306–325, II ca. 300–350, III 323–350, IV 320–350. Form of legs and their setae as in deutonymph except with normal protonymphal complement of setae (see genus description); tarsus I apically bearing pretarsus (10–16 to base of claws). Tarsi II–IV with apical setal processes ad -1, pd -1 (13–15) similar in length to pretarsi (11–15 to base of claws); with ventroapical process under pretarsus as in deutonymph (15–18), bluntly triangular. Form of leg setae much as in deutonymph, but with spinelike setae less developed.
Larva. Idiosomatic dorsum. Idiosoma oval, 475–580 long, 375–428 wide at level of setae j6 (n = 2); dorsum weakly sclerotised, without clearly delineated shielding but faintly lineated posterolaterally (Fig. 34). Dorsal podonotal setae slightly longer than opisthonotal setae; j1, j3 (12–13), j4, j5, j6 longest (23–28), z2, z4, s4, s6 (18–22); J2, J3, J4, Z3–Z5, S3–S5 (11–13), J5 minute (3).
Idiosomatic venter. Tritosternum with base as long (26) as wide (25); laciniae free for about 80% of total length (86 excluding base). Sternal region smooth, with few lines behind setae st1; setae st1–st3 similar in length (15–17) (Fig. 35). Anal shield mostly smooth, as long as wide (ca. 70); para-anal setae (19–22) clearly longer than postanal seta (12–14). Four pairs of opisthogastric setae, JV1, ZV2 (8–10), JV5 (7–8), JV2 longest (12–15).
Gnathosoma. Gnathotectum with anterior margin unevenly triangular and smooth (Fig. 36). Fixed digit of chelicera with offset tooth near apical hook, alveolar vestige of pilus dentilis, and a small second tooth 5 µm distant from tip; movable chela (25) bidentate, with small ventral process (Fig. 38). Palps 98–125 long. Internal malae with external branch thick, pilose, shorter than internal branch (Fig. 37 a). Deutosternum with seven rows of denticles, basal rows with one to three teeth, distal rows with seven to nine teeth. Hypostomatic setae hp1 (36–37) thick, longer than pc (18–20); corniculi (18) inserted at same transverse level as alveoli of setae hp1, with inner acuminate process shorter (8) relative to subsequent instars, its tip not reaching that of main corniculus tip; salivary stylets (19) blunter than in subsequent instars (Fig. 37 a). Lateral seta al -1 of palpal femur and genu slender, tapered, as in protonymph; palptarsal claw with dissimilar tines (20 the longer, 15 the shorter).
Legs. Leg lengths similar, excluding pretarsi: I (338–340), II (325), III (325–342). Form of legs and their setae generally as in protonymph. Tarsus I (88–91) twice as long as tibia I (45). Tarsi II and III apically with well defined ventral process (7–9) shorter than setal processes ad -1, pd -1 (15) and pretarsus (14–16) to base of claws; tarsi II–III with setae a v -1, pv -1, al -2, pl -1 similarly longer (14–15), more attenuate than other tarsal setae.
Type material. All material collected from one locality, with initials of collectors indicating Arthropods of La Selva parataxonomists (ALAS, see acknowledgments), Derek M. Johnson (DMJ), Evert E. Lindquist (EEL). COSTA RICA. Heredia Province, La Selva Biological station (10°26’1” N, 84°1’2” W, elevation 50–150 m). HOLOTYPE: adult female, ex adult Chelobasis sp. on leaf of Heliconia pogognantha, February 1994, coll. EEL & ALAS: deposited in Instituto Nacional de Biodiversidad (INBio) of Costa Rica, Santo Domingo de Heredia. PARATYPES: two adult females, seven adult males, one deutonymph, two protonymphs (one pharate deutonymph), one larva, with same data as holotype; one adult female, one adult male, ex Chelobasis sp., June 1997, coll. DMJ; five adult males, four deutonymphal males, one deutonymphal female, one protonymph, under elytra of Chelobasis perplexa Baly from rolled Heliconia leaf, July 1997, #739025a, coll. ALAS; two adult females, two adult males, one deutonymph (pharate adult female), one larva (pharate protonymph), same data as previous entry, #739025b, coll. ALAS; one adult female, same data as previous entry, #739025c, coll. ALAS; two adult females, two adult males, same data as previous entry, #739026, coll. ALAS.
Host identifications. The identities of the host beetles and background plant substrate given above are in accord with current molecular studies, which indicate that one haplotype of Chelobasis perplexa sensu stricto is found on Heliconia pogonantha in the La Selva area of Costa Rica (Carlos Garcia-Robledo, personal communication, June 2014).
Etymology. The specific name indicates the conspicuously reticulate ornamentation of the intercoxal shielding of adults, and also the adults being commonly ornamented with remarkable laboulbeniaceous fungal growth.
Remarks. As noted in the description and confirmed by views of specimens with pharate adults, deutonymphs of Makarovaia ornata are dimorphic in the form of their movable cheliceral digits. While the movable digit of the female deutonymph is identical in form with that of the female adult, that of the male deutonymph displays anlage of the adult sperm duct, and has a straighter masticatory surface bearing just one minute tooth, instead of a series of five well-formed teeth.
description: Published as part of Moraza, María L. & Lindquist, Evert E., 2015, Systematics and biology of mites associated with neotropical hispine beetles in unfurled leaves of Heliconia, with descriptions of two new genera of the family Melicharidae (Acari: Mesostigmata: Gamasina: Ascoidea), pp. 301-351 in Zootaxa 3931 (3) on pages 315-326, DOI: 10.11646/zootaxa.3931.3.1, http://zenodo.org/record/244189