Spadiseius calyptrogynae, Lindquist, Evert E. & Moraza, Maria L., 2008

Lindquist, Evert E. & Moraza, Maria L., 2008, A new genus of flower-dwelling melicharid mites (Acari: Mesostigmata: Ascoidea) phoretic on bats and insects in Costa Rica and Brazil, Zootaxa 1685, pp. 1-37 : 9-21

Spadiseius calyptrogynae new species

( Figs. 1–6 , 10–34 , 38, 40 , 44 , 68, 69 )

Diagnosis. All instars: leg I with apical sclerotized rim of femur, genu, tibia usually smooth, not serrated, ventrally, those of other legs weakly to clearly serrated. Adult: peritremes of female and male of moderate thickness, and extending anteriorly well beyond humeral setae r3 to level of setae s1; hypostomatic setae hyp4 shorter than hyp1–hyp3. Adult female: marginal setae r2–r4 and sometimes r5 on edge of dorsal shield; subcapitulum with pair of gland-like structures between insertions of setae hyp1, and with setae hyp4 abruptly swollen basally. Adult male: dorsal shield with setae in central region short, simple, often inserted individually on small subcircular warts, and with setae j1 and a variable number of peripheral setae of z–Z, s–S and r– R series forming a corona of enlarged, thickened, apically capitate setae; presternal area with pair of small platelets; peritrematal shield with gland pore greatly enlarged at level between coxae II and III; spermatodactyl divided near base into a shorter pointed process and a longer, straight, apically thickened terminus; leg IV lacking spur on basitarsus. Deutonymph: coxa IV seta short, half as long as setae on coxae III; legs I–IV with femoral seta ad-1 thicker than adjacent setae, and clearly spatulate on IV but not on I–II and sometimes III. Protonymph: pygidial shield usually with poroid-bearing hump or tubercle laterad seta S5; legs I–IV with femoral seta ad-1 not spatulate but slightly thicker than adjacent setae. Larva: podonotal shield faintly lineated laterally; pygidial shield sparsely puncate, with medial projection of anterior margin not extending beyond level of setal alveoli J2; legs I–III with femoral seta ad-1 not spatulate but less tapered, slightly thicker than adjacent setae.

Description. ADULT FEMALE. Dorsal shield 440–540 long, 275–325 at its greatest width at level of setae r4 (n = 8); shield lightly evenly reticulate over nearly entire surface ( Fig. 18 ); podonotal region with longitudinal distance from alveoli of j1 to j5 slightly longer (about 1.1) than that from j5 to j6. Peritrematal shields uniting with dorsal shield at level between setae s1–s2; peritremes extending to level of setae s1. Body dorsum with approximately 52–54 pairs of setae, most inserted on dorsal shield, but variably r5 and consistenly R1– R6, and 0 to 3 UR– setae inserted on soft cuticle; dorsal shield with 3 to 5 extra pairs of setae amidst j, z and s series on podonotal region and several paired or asymmetrically unpaired setae amidst J series on opisthonotal region. Dorsal setae uniform in simple but slightly curved, scimitar-like form (most of them slightly swollen at base) and moderately short length (17–23), S5, Z5 slightly largest (23–25), J5 clearly smallest (10). Anterodorsal expansions of exopodal plates above legs I formed as lineated plates (not illustrated, difficult to discern) touching vertex of dorsal shield.

Tritosternum ( Fig. 4 ) with length of trapezoidal base subequal to its basal width (16–17); laciniae free for about 0.6 of total length (70–75 excluding base). Presternal area weakly sclerotized, insensibly fused with sternal shield, and with pair of small platelets amidst lineation. Sternal shield ( Fig. 19 ) unornamented or faintly lineated along lateral margins, weakly sclerotized over much of surface but faint or desclerotized posterior to level of second poroids, posterior margin faint, slightly concave; setae st1–st2 and first and second pairs of poroids inserted on sclerotized portion of shield, st3 inserted on desclerotized portion; st1–st3 similar in length (30–35), slightly longer than st4 (25–27) and st5 (20) on soft cuticle. Endopodal strips faintly formed alongside coxae III and IV. Genital shield unornamented, its lateral margins slightly concave at level of genital setae. Anal shield weakly reticulate-lineate, longer (90) than greatest width at level of anterior edge of anus (75); postanal seta (20–22) slightly longer than para-anal setae (17–18). Ten pairs of opisthogastric setae (JV1–JV5, ZV1–ZV5, excluding 1 or 2 pairs of adjacent UR– setae) on soft cuticle around anal shield, all similarly short (16–18) except for slightly larger JV5, ZV5 (22–24). Metapodal plates undivided, usually elongate triangular, pointed posteriorly, about 25 long, 12 wide. Spermathecal apparatus difficult to discern, with convoluted conduit and weakly sclerotized calyx ( Fig. 20 ).

Tectum with anterior margin smooth, sharply pointed medially (as in Fig. 7 ). Fixed chela with 2 or 3 minute teeth beside offset tooth subapically (as in Fig. 50 ); movable chela (length 40) usually weakly bidentate, the more apical tooth hardly discernible; antiaxial face of movable chela with sharply defined ridge along its basal half; ventral mucro slender, attenuate, surpassing basal half of chela (as in Figs. 8, 9 ). Corniculi with lateral edges convergent apically but with medial edges nearly parallel such that apices usually separate; internal malae slender, blade-like, shorter than corniculi, their lateral margins smooth ( Fig. 1 ). Deutosternum with 7 or 8 transverse rows of denticles, evenly multidenticulate (7–15 denticles), none distinctly widened. Subcapitulum with pair of gland-like structures between bases of anterior subcapitular setae ( Fig. 1 ); setae finely tapered, medial pair longest; capitular pair abruptly thickened basally, shorter (12–15) than anterior and medial pairs (27 and 35–40, respectively). Subcapitulum without ridge extending on either side laterally from bases of capitular setae. Palptrochanter setae slender, inner seta slightly longer than outer one; palpgenu with seta al-2 similar in spatulate form to palpfemoral seta al; apical sclerotized rim of palpfemur and palpgenu smooth ventrally ( Fig. 5 ).

Legs IV the longest (415–480, excluding pretarsus) of legs, about 0.9 as long as dorsal shield; other leg lengths: I 385–430, II 335–375, III 350–400. Tarsi II–IV with pair of apical setal processes (ad-1, pd-1) about one-half as long as length of pretarsi to base of claws ( Fig. 34 ). Leg II slightly thicker than legs III–IV, but thickness and form of its setae not differentiated from those of legs III–IV ( Figs. 31–33 ). Coxae I with several serrated ridges on ventral face, coxae II with 2 or 3 serrated ridges on posteroventral face, coxae III–IV unornamented (other than vertical line caused by small boss below each seta); seta on coxa IV short (11–12), half as long as setae on coxa III (22–24) ( Figs. 32, 33 ). Apical sclerotized rim of femur, genu, tibia usually smooth ventrally on leg I, but usually sparsely or irregularly serrated on legs II–IV. Leg IV with trochanter seta ad and femoral seta ad-2 stout, poorly tapered. Femora I–IV with dorsal seta ad-1 thickened, untapered, rod-like or slightly oar-like in contrast with adjacent simple setae, increasingly longer from I (15–16) to IV (25–27) ( Figs. 30–33 ); legs without elongated macrosetae. Setation of genua and tibiae I–IV as presented for genus.

ADULT MALE. Dorsal shield 445–480 long, 335–440 at its greatest width at level of setae r4 (n =10); podonotal region with longitudinal distance from alveoli of j1 to j5 slightly less (0.8–0.9) than that from j5 to j6; shield with uniformly short (7–10) setae, including some hypertrichy, on its central and posterior area, and with a corona of a variable number of enlarged (length to 135), apically capitate or oar-like setae peripherally ( Figs. 21 , 23 ); setae j1 consistently untapered, slightly oar-like. Dorsal shield ornamentation and form of peripheral setae polymorphic, with short setae inserted individually on circular wart-like thickenings correlated with more elongated peripheral setae (75–140) on some specimens ( Fig. 23 ), but with none or few of short setae on such thickenings and shield reticulate, correlated with less elongated peripheral setae (30–80) on others ( Figs. 21 ); whether elongated or not, S1 consistently shorter (0.5–0.7) than adjacent setae s6 and S2 ( Table 1 ). Peritrematal shields uniting with dorsal shield at level between bases of legs II and III, shield gland pore at that level greatly enlarged ( Figs. 24, 25 ); peritremes extending to level of setae s1. Dorsal shield complement of setae, poroids and gland-pores similar to that in female, marginals r5, R1–R2, R4–R6, poroid Rp and sometimes 1 or 2 UR setae variably on soft cuticle or on edge of shield; R3 sometimes absent.

Tritosternum much as in female, but with length of trapezoidal base (13–15) less than its basal width (20– 22), and laciniae free for about 0.8 of total length (65–68 excluding base). Presternal area with pair of small platelets ( Fig. 22 ). Sternogenital shield unornamented; of four or five pairs of setae on shield, st4 (50–60) clearly longer than st1–st2 (15–28) and st3 (20–25), st5 variably much longer (85–110) and often inserted near paragenital poroids on soft cuticle closely beside invaginated posterolateral corners of shield; posterior margin of shield strongly delineated, broadly rounded. Ventrianal shield unornamented, with weakly delineated anterior and lateral margins variable in outline; shield length (185–190) about 0.6–0.7 its greatest width (280–305) anteriorly where metapodal plates incorporated; anal region more clearly sclerotized, sometimes appearing like a distinct plate surrounded by desclerotized cuticle ( Fig. 22 ); ventrianal shield with circumanal setae and usually 7 pairs of opisthogastric setae (JV1–JV4, ZV1–ZV3); ZV1–ZV2 more attenuated and variably longer (15–50), and JV4 variably longer (25–80) than in female; ZV3 variably oarlike (25) or attenuate (63–90); postanal seta variably longer (25–40) than para-anal setae (16–25); JV5 usually clavate distally (30–60) and variably attenuate (ca 90), inserted on soft cuticle; ZV4–ZV5 absent. Form of peritrematal and exopodal shields posteriorly as in female.

Tectum as in female. Cheliceral shaft abruptly narrowed dorsally at base of fixed digit ( Fig. 27 ); fixed digit with a small offset tooth followed by 2–3 minute teeth at level of hyaline pilus dentilis, and with a large rounded tooth (or lobe) basally; spermatodactyl biramous, with a pointed hyaline process above the nearly straight, apically lobed main shaft which extends anteriorly about 28–35 beyond apex of chela ( Fig. 27 ). Corniculi more slender and widely spaced basally than in female; internal malae larger than in female, with fringed lateral margins and bifid apices ( Fig. 26 ). Subcapitulum with rows of deutosternal denticles slightly wider and more multidenticulate than in female. Subcapitulum lacking the pair of gland-like structures near setae hyp 1 in female, and with capitular setae short but not conspicuously swollen basally; other subcapitular and palpal structures similar to those in female.

Relative leg lengths, excluding pretarsi, similar to those of female, I (380–440), II (325–395), III (350– 420), IV (425–510). Coxa IV seta relatively short as in female; apical sclerotized rim of femur, genu, tibia usually smooth ventrally on legs I and sometimes on legs II, but usually serrated on legs III–IV. Tarsi II–IV with pair of apical setal processes (ad-1, pd-1) nearly half as long as length of pretarsi to base of claws. Legs II curved and thicker relative to legs III–IV than on female, and with setae av-1, av-2, av-3 (mv) modified to short, stout, bulbous spines ( Fig. 28 ); legs III–IV also slightly thicker relative to leg I than in female; basitarsus of legs III–IV without spur ( Fig. 29 ). Leg IV with trochanter seta ad and femoral seta ad-2 attenuate or clavate. Femur I with dorsal seta ad-1 variably rod-like (30) or oar-like (45) but slenderer than in female; ad-1 on femora II–IV variably oar-like (32–65) or attenuated (90–135), either way increasingly longer from II to IV. Legs I–IV with size and form of many dorsal and lateral setae variably polymorphic, oar-like or attenuated on genu, tibia and tarsus, and legs III–IV with some ventral setae variably attenuated on femur, genu, tibia and tarsus ( Figs. 68, 69 ).

DEUTONYMPH. Dorsal shield 405–450 long, 270–310 wide at level of lateral incisions by setae s6 (n = 4); shield lightly evenly reticulate over nearly entire surface ( Fig. 16 ); podonotal region with longitudinal distance from alveoli of j1 to j5 nearly equal (0.9) to that from j5 to j6. Dorsal shield with approximately 35 to 40 pairs of setae: about 17–18 pairs (j1–j6, z2–z6, s3–s6, and 2 to 4 pairs of extra setae) on podonotal region, and 18 pairs (J1–J5, Z1–Z5, S1–S5, and a few paired or asymmetrically unpaired setae amidst J series) on opisthonotal region. Dorsal setae mostly alike in simple form (most of them slightly swollen at base) and short length (10–15) in central area, s3–s6, S2–S3 slightly larger (20–23), S1 only half as long as adjacent setae s6 and S2, and S4, S5, Z5 poorly tapered, slightly barbed and largest (25–28). Dorsal setae on soft cuticle mostly 10–15 long, r3 and R1 slightly longest (15–20). Peritremes and their shields extending anteriorly to level of setae s1.

Tritosternum similar in form to that in adult female. Sternal shield smooth, with narrowly rounded posterior margin ( Fig. 17 ); setae st4 clearly shorter (8–10) than st1–st3 (25–27). Anal shield unornamented, length subequal to greatest width (90–95) at level of anterior edge of anus; postanal seta subequally as long as paraanal setae (15–17). Opisthogaster with 8 to 10 pairs of ventral setae plus 1 or 2 adjacent UR– setae (ZV4 and ZV5 sometimes absent), all similarly short (10–12), slender except for slightly larger JV5 (15) ( Fig. 17 ). Metapodal plates broadly oval, greatest dimension 15.

Tectum, chelicerae, ventral gnathosomatic structures and palpi similar to those in adult female, except capitular setae not swollen basally, and without pair of gland-like structures between setae hyp1.

Relative leg lengths, excluding pretarsi, similar to those of female, I (360–380), II (310–325), III (325– 350), IV (380–400). Form of legs and their setae as in adult female, except leg II no thicker than legs III–IV; tarsus I with pedestal bearing pretarsus less defined than on protonymph; tarsi II–IV in dorsoventral view tapered apically beyond insertions of al-1, pl-1. Coxa IV seta short (10), at most half as long as coxa III setae. Dorsal seta ad-1 increasingly longer on femora I to IV (10 to 20), thicker, poorly tapered but not rod- or oarlike on femur I and sometimes on femur II, but slightly oar-like on femur III and clearly so, in contrast with adjacent simple setae, on femur IV ( Fig. 44 ).

PROTONYMPH. Idiosoma 305–345 long, about 215–235 wide at level of setae r5 (n = 5), with lightly sclerotized, fully reticulated podonotal and pygidial shields ( Fig. 13 ); podonotal shield 207–220 long, 170–190 at widest level near posterior margin, with longitudinal distance from alveoli of j1 to j5 about 0.8 that from j5 to j6; pygidial shield 72–80 along midlength, 143–175 at widest level near seta S4, with anterior margin shallowly excavated at either side of midline, at level of setae J2, and usually with poroid-bearing protuberance laterad seta S5. Dorsal body setae mostly of similar short length (12–20) and simple form, opisthonotal setae Z5 slightly longer (22–25), S4, S5, Z5 slightly barbed, clunals J5 shortest (5). Peritremes extending anteriorly nearly to level of anterior margins of coxae III.

Tritosternum similar in form to that in deutonymph and adult female, laciniae free for about two-thirds of length (50). Sternal shield ( Fig. 14 ) smooth; setae st5 inserted on soft cuticle much shorter (6) than st1–st3 (15) and shorter than opisthogastric setae; paragenital poroids absent. Anal shield unornamented, similar in form and structure to those on deutonymph, length subequal to greatest width (60–65); postanal seta subequally as long as para-anal setae (12). Opisthogastric setae on soft cuticle all similarly short (9–10) except for slightly larger JV5 (12). Metapodal plates faint, elliptical (6 length, 10 width).

Tectum more broadly triangular and bluntly pointed than in deutonymph ( Fig. 15 ); other gnathosomatic structures similar to those in deutonymph, except palpi with normal protonymphal complement of setae.

Leg lengths, excluding pretarsi, I (280–305), II (232–268), III (240–275), IV (270–325); pretarsi II–IV with apical setalike processes nearly half as long as length of pretarsi to base of claws. Form of legs and their setae generally as in deutonymph, except tarsus I apically with more clearly defined pedestal (5) bearing pretarsus. Coxa IV seta shorter (11) than coxa III setae (15). Femora I–IV with dorsal seta ad-1 similar in length (about 10), slightly more thick and setulose than adjacent setae, but tapered, not rod- or oar-like ( Fig. 40 ).

LARVA. Idiosoma 290–360 long, 175–215 wide at level of setae s4 (n = 6), with lightly sclerotized, faintly and incompletely lineated podonotal and pygidial shields ( Fig. 10 ); podonotal shield 185–205 long, 150–155 at widest level near posterior margin, with longitudinal distance from alveoli of j1 to j5 0.7 that from j5 to j6, and with undulate lateral margins constricted behind levels of setae z2 and s4; pygidial shield sparsely punctate, 65–70 along midlength in dorsal view (but further length of 15–20 folded under ventrally), 145–150 at widest level near anterior margin, with anterior margin deeply invaginated on either side of midline, at level of setal alveoli J3, and posterior margin also deeply invaginated on either side of midline at level of setal alveoli J5 ( Fig. 11 ). Dorsal body setae similarly short (6–9), simple on podonotal shield, slightly shorter (4–5) on soft cuticle, but Z3–Z4 on pygidial shield much longer (37–40), thicker, capitate distally.

Tritosternum with base longer (17–25) than wide (9–11); laciniae free for about three-fourths of total length (40 excluding base). Sternal shield smooth, with setae st1–st3 similar in length (12–14) ( Fig. 11 ). Anal shield smooth, slightly wider (50–55) than long (45–50); para-anal setae (15–18) longer than postanal seta (12–13). Four pairs of opisthogastric setae (ZV2 present) plus S5 and Z5 on soft cuticle all similarly short (3– 4) except for slightly longer JV2 (7–14).

Tectum with anterior margin bluntly unevenly triangular ( Fig. 12 ). Fixed digit of chelicera with dentition and pilus dentilis lobe as in protonymph; movable chela 20–22 long, edentate or with a minute tooth, and with ventral mucro ( Fig. 6 ).

Leg lengths, excluding pretarsi: I (255–280), II (192–228), III (210–230); pretarsi II–III with apical setalike processes about half as long as length of pretarsi to base of claws. Form of legs and their setae generally as in protonymph. Tarsus I apically with well defined pedestal (7) bearing pretarsus. Femora I–III with dorsal seta ad-1 (8–9) resembling pd-1 of femur and ad–pd setae of genua, i.e., slightly more thick and setulose than other femoral setae, but not rod- or oar-like ( Fig. 38 ).

Type material. COSTA RICA, Heredia Province, La Selva Biological Station, (10° 26' 1" N, 84° 1' 2" W) elevation 50–150 m: HOLOTYPE: adult female, Research Swamp, ex inflorescences of Palmaceae, Calyptrogyne sarapiquensis , 10 February 1994, coll. S. Cunningham & E.E. Lindquist. PARATYPES: 4 adult females, 10 adult males, 6 deutonymphs, 14 protonymphs, 8 larvae, with same data as holotype; 2 adult females, 1 adult male, 6 protonymphs, 3 larvae, Research Swamp, ex inflorescences of C. sarapiquensis , 15 March 1991, coll. S. Cunningham; 1 adult female, 1 adult male, 2 protonymphs, ex scarab beetle on inflorescence of C.

sarapiquensis , 27 March 1991, coll. S. Cunningham; 1 adult female, 1 protonymph, Research Swamp, ex elytrum of scarab, Lagochile collaris , on inflorescences of C. sarapiquensis , 2 May 1994, coll. S. Cunningham; 4 adult females, ex scarab, Lagochile collaris , 22 February 1996, coll. M. Paniagua; 2 adult females, 1 protonymph, ex curculionid beetle, 24 February 1994, coll. S. Cunningham; 4 adult females, 9 adult males, ex bats, January & February 1993, coll. S. Cunningham; 1 adult female, 2 adult males, 1 protonymph, ex bats(?), 20 March 1995, coll. S. Cunningham.

Note: The above data are based on the labels affixed to slide preparations or associated with residual samples preserved in ethanol. Calyptrogyne sarapiquensis H. Wendl. ex Burret is presently considered to be a junior synonym of C. ghiesbreghtiana (Linden & H. Wendl.) (see Grayum 2003b).

Etymology. The specific epithet, calyptrogynae , is based on the name of the genus of palmaceous plants in whose inflorescences these mites undergo their life histories.

Remarks. Of the five recognized species of small palms of the Neotropical genus Calyptrogyne , C. ghiesbreghtiana is the most widespread and variable species, occuring in both lowland rainforests and mid-elevation cloudforests on both the Atlantic and Pacific coasts, from southern Mexico to Panama ( Grayum 2003b). Calyptrogyne ghiesbreghtiana is the only well documented case in the palm family Arecaceae of being pollinated, in part at least, by bats ( Cunningham 1995). Spadiseius calyptrogynae is the first flower-dwelling melicharine mite recorded to use pollinating bats for dispersal ( Tschapka & Cunningham 2004). Eight species of bats, including nectar-specialists and frugivores, are known to visit inflorescences of C. ghiesbreghtiana in a given study area of lowland rainforest in Costa Rica, and their visiting behaviors differ greatly. While the glossophagine nectar-specialists visit inflorescences in extremely short hovering flights of less than one second, the usually frugivorous stenodermine bats, including two species of Artibeus , land and perch on the inflorescences for minutes, occasionally up to 17 minutes. Although the glossophagine bats are much more frequent visitors than Artibeus to C. ghiesbreghtiana , their short hovering flights and lack of a smooth extended walkway like that provided by hummingbird bills evidently do not afford the mites sufficient time and means to mount or dismount ( Tschapka & Cunningham 2004). In contrast, perching by Artibeus bats provides close body contact between them and inflorescences along with longer visit durations. Mites were nearly always found on the surfaces of these bats’ wings, in the same places that most of the C. ghiesbreghtiana pollen grains were found.

Apart from the diversity of mites that are parasites of bats, few other mites have been noted to be phoretic on bats. Adult females of a tarsonemid, Alkithoenemus cynopterus Ochoa & OConnor, thought to undergo development within the fleshy syconial receptacle of figs, are phoretic on the frugivorous bat, Cynopterus nusatenggara Kitchener & Maharadatunkamsi. However , this bat is not a pollinator of figs, and species of Cynopterus feed on a wide variety of fruits, even though figs may constitute large parts of their diets ( Ochoa et al. 1996).

The above collection records support observations by Cunningham (1995) that a scarabeid beetle, Lagochile collaris Blanchard , which feeds destructively on C. ghiesbreghtiana flowers, provides alternative transportation for this mite. About two-thirds of the beetles observed on inflorescences carried mites ( Tschapka & Cunningham 2004), which locate on external surfaces, not underneath the elytra (Ellen Sperr, personal observation, July 2007). The single collection record of a few female and protonymphal mites phoretic on an unidentified curculionid beetle, without a more specific identification of the latter, needs confirmation as a phoretic association between these mites and a carrier of this family of beetles.

A sample of two fully developed inflorescences of Calyptrogyne ghiesbreghtiana taken at the La Selva Biological Station, 10 February 1994 by one of us (EEL), contained between 400 and 500 mites, not accounting for eggs, including an abundance of all developmental instars and adults.

The greatly enlarged peritrematal gland pore in adult male S. calyptrogynae is a form of sexual dimorphism not observed before among mesostigmatic mites. Adult males available of three other congeneric species ( S. spathiphyllae and two of the undescribed species) lack this attribute.

Figures

FIGURE 18. Spadiseius calyptrogynae n. sp., adult female, idiosoma, dorsal aspect. Supernumerary setae denoted by

FIGURES 1 – 9. Figs. 1 – 5, Spadiseius calyptrogynae n. sp., adult female: 1, subcapitulum; 2, labrum, ventral view; 3, salivary stylus; 4, tritosternum; 5, palpus, ventral view. Fig. 6, S. calyptrogynae n. sp., larva, chelicera, paraxial view. Figs. 7 – 9, Spadiseius spathiphyllae n. sp., adult female: 7, tectum; 8, chelicera, dorso-paraxial view; 9 a, chelicera, movable digit, dorsal view; 9 b, chelicera, movable digit, antiaxial-ventral view (Figs. 2 – 4, same scale).

FIGURES 10 – 12. Spadiseius calyptrogynae n. sp., larva: 10, idiosoma, dorsal aspect; 11, idiosoma, ventral aspect; 12, tectum, variations.

FIGURES 13 – 15. Spadiseius calyptrogynae n. sp., protonymph: 13, idiosoma, dorsal aspect; 14, idiosoma, ventral aspect; 15, tectum. Setae added to larval complement denoted without parentheses.

FIGURES 16, 17. Spadiseius calyptrogynae n. sp., deutonymph: 16, idiosoma, dorsal aspect; 17, idiosoma, ventral aspect. Setae added to protonymphal complement denoted without parentheses; supernumerary setae denoted by ‘ x’.

FIGURES 19, 20. Spadiseius calyptrogynae n. sp., adult female: 19, idiosoma, ventral aspect; 20, spermathecae, detail

FIGURES 21, 22. Spadiseius calyptrogynae n. sp., adult male, “ wartless ” morph: 21, idiosoma, dorsal aspect; 22, idiosoma,

FIGURES 23 – 25. Spadiseius calyptrogynae n. sp., adult male: 23, idiosoma, “ warted morph ”, dorsal aspect; 24, peritrematal shield and detail of enlarged gland pore; 25, greatly enlarged detail of peritrematal gland pore.

FIGURES 26 – 29. Spadiseius calyptrogynae n. sp., adult male: 26, subcapitulum; 27, chelicera and spermatodactyl, paraxial view; 28, tarsus II, anterolateral view; 29, tarsus IV, dorsal view, without pretarsus. Abbreviation: k, keel.

FIGURES 30 – 34. Spadiseius calyptrogynae n. sp., adult female. Figs. 30 – 33, dorsal views of legs excluding tarsi: 30, leg I; 31, leg II; 32, leg III; 33, leg IV; anterolateral (al) and posterolateral (pl) faces indicated by arrows. Fig. 34, tarsus IV, dorsal view.

FIGURES 38 – 41. Dorsal subapical femoral setae of legs. Figs. 38, 39, larval femora I – III: 38, Spadiseius calyptrogynae n. sp.; 39, Spadiseius spathiphyllae n. sp. Figs. 40, 41, protonymphal femora I – IV: 40, S. calyptrogynae n. sp.; 41, S.

FIGURES 42 – 45. Figs. 42, 43, Spadiseius spathiphyllae n. sp., deutonymph: 42, idiosoma, dorsal aspect; 43, idiosoma, ventral aspect; supernumerary setae denoted by ‘ x’. Figs. 44, 45, dorsal subapical femoral setae legs I – IV, deutonymph: 44, Spadiseius calyptrogynae n. sp.; 45, Spadiseius spathiphyllae n. sp.

FIGURES 65 – 69. Figs. 65 – 67: Spadiseius spathiphyllae n. sp., adult male: 65, tarsus IV, dorsolateral view, without pretarsus; 66, 67, femora IV, dorsal view: 66, “ short-haired ” morph; 67, “ long-haired ” morph. Figs. 68, 69, Spadiseius calyptrogynae n. sp., adult male, femora IV, dorsal view: 68, “ wartless ” morph; 69, “ warted ” morph.

FIGURES 46 – 50. Spadiseius spathiphyllae n. sp., adult female: 46, idiosoma, dorsal aspect, supernumerary setae denoted by ‘ x’; 47, idiosoma, ventral aspect; 48, spermathecae, detail of calyx; 49, position of spermatheca at base of leg IV;

Tables

TABLE 1. Polymorphic male measurements (µm) - Spadiseius calyptrogynae n. sp.

Morph	shield length	shield width / ratio	leg IV length	st5	JV5	s6	S1	S2	S5	IVFe -v IVFe- ad-1
wartless	440	320 0.72	430	45	20	30	20	30	57	37 30
wartless	467	312 0.67	450	87	30	62	40	80	80	52 42
warted	485	400 0.82	492	112	60	120	82	135	125	85 67
warted	460	375 0.82	465	125	50	100	70	97	107	87 55
warted	485	400 0.82	467	115	45	120	82	137	132	80 57
warted	487	395 0.81	500	100	38	100	65	100	107	80 55
warted	480	360 0.75	467	112	32	112	65	100	107	80 50
warted intermed	467	350 0.75	445	90	30	80	50	80	87	67 50