Hispiniphis Moraza & Lindquist, 2015, new genus

Hispiniphis new genus (Figs 41–92, 101– 105) Type species. Hispiniphis parva new species. Genus based on adult female, adult male, nymphal and larval material representing six species, of which one is newly described here, the others to be described subsequently. Diagnosis. Adult Hispiniphis are distinguished from those of Makarovaia in having the dorsal shield less extensive anterolaterally, leaving at least setae z 2, all s, and all r setae on adjacent soft cuticle, which has a thicker, micropunctate appearance distinct from the normally striate ventral soft cuticle. Adults have less strongly sclerotised and ornamented shields on the venter, including a smaller, inversely subtriangular anal shield, slightly longer than wide. Male stenitigenital shield incompletely formed, deeply excavated posteriorly, leaving setae st 5 on soft cuticle or on a separated platelet medially. Hypostome of nymphs and adults short, corniculi inserted at level only slightly behind insertions of hypostomatic setae hp 1. Movable cheliceral digit of female and nymphs tridentate, and with pointed process discernible on midventral face. Male chelicerae lacking a digitiform process at level where movable digit hinged to fixed digit, and with spermatodactyl projecting freely anteriorly. In addition to leg chaetotactic reductions noted for genus-group description, femur I and tibia I lacking dorsal seta ad - 3; genu II lacking lateral seta al - 2; genua II and III retaining ventral seta av. Tarsus I with ventrodistal acuminate process extending under pretarsus; tarsi II–IV with ventral projection under pretarsus seeming articulated, acuminate in immature instars and adults. Protonymph and to lesser extent deutonymph with opisthosoma reduced in form, size, sometimes flanked closely by posteriorly-directed legs IV. Description. Idiosomatic dorsum . Adult female (Fig. 42). Dorsal shield moderately well sclerotised, with reticulate ornamentation; shield relatively smaller in size, compared to overall dorsal surface area, not extensive enough to bear setae s 1, z 2, r 2, r 3 which remain with s 4, s 5, s 6, r 5 on surrounding soft cuticle; soft cuticle thickly striate, micropunctate, giving ornamented effect in contrast with opisthogastric soft cuticle (Fig. 103, 104). Dorsal shield with a maximum complement of 13-20 pairs of setae, including seven to 12 podonotal (j 1, j 3 –j 6, z 4, z 5, z 6, sometimes j 1 off shield, sometimes j 4 duplicate) and six to 11 opisthonotal pairs (J 1 +, J 3, J 4 +, J 5, Z 2 –Z 5, sometimes Z 2, Z 3, and Z 5 duplicate) (Fig. 88); dorsal setae usually similar in form, somewhat variable in length, except for J 5 always minute. Dorsal shield with complement of 13–15 pairs of discernible pore-like structures (three to five podonotal, 10 opisthonotal), of which five to seven pairs (two to four podonotal, three or four opisthonotal) superficially appear secretory (gland pores) and seven to nine pairs (one or two podonotal, six or seven opisthonotal) non-secretory (poroids). Soft lateral cuticle with 20–45 pairs of lateral, marginal and submarginal setae, including z 2, all s, all r, sometimes Z 2, and all S, R, and UR, and at least six pairs of marginal poroids, including idRp; few to many of setae on soft cuticle may be duplicate. Adult male ( Fig. 56). Dorsal shield usually similar in size, ornamentation and setation to that of female; sometimes capturing one to three more pairs of setae than in female; form and relative lengths of shield setae as in female. Idiosomatic venter. Adult female (Fig. 45). Tritosternum with laciniae fused for about half of their length, without basal elaborations; laciniae nearly devoid of pilosity (Fig. 47). Ventral shields variably poorly sclerotised. Presternal region smooth or faintly lineated, without platelets (Fig. 45). Sternal shield fragmented, its form and setation as noted for genus-group description; sternal area with varying faint ornamentation; shield fragment sometimes with bases of setae st 1, st 2 circumscribed by sclerotised thickenings, while unsclerotised medial region sometimes faintly ornamented but not striated with soft cuticle. Epigynal shield lightly sclerotised, unornamented, narrow, with hyaline anterior margin hardly widened between legs III, and with posterior margin hardly widened but strongly rounded or bluntly pointed (Fig. 48). Opisthosomatic venter lacking metapodal platelets, with soft striated cuticle bearing four to six pairs of opisthogastric setae (JV 1, JV 2, ZV 2, JV 5, sometimes JV 3, JV 4) and four pairs of poroids, flanked laterally and posteriorly by further thickened, ornate cuticle variably bearing several pairs of R - or UR - setae. Anal shield small, well sclerotised, smooth or reticulated, pyriform or inversely subtriangular, slightly longer than wide; para-anal setae inserted at mid-level of anus, shorter than postanal seta; shield with pair of gland pores gv 3 on lateral edges at level of insertions of para-anal setae, and with cribrum along tapered posterior margin (Fig. 49). Peritrematal shield weakly sclerotised, extending without interruption to about midlevel of coxae I; shield with gland pore gp 1 located at anterior extension at level of coxa I (Fig. 43); exopodal strip continuous alongside peritrematal shield only to level of coxae III, extensions between bases of coxae II–III rudimentary or absent. Peritremes thick, reaching at most to level of anterior margin of coxae II (Figs 43, 45). Spermathecal apparatus sometimes with small section of tubular piece discernibly sclerotised, slightly widened (Fig. 55). Adult male (Fig. 60). Form of tritosternum and presternal area as in female. Sternitigenital shield anterior margin well defined at level of genital opening; shield lightly sclerotised, united with endopodal strips between coxae I–II, II–III, III–IV; shield with four pairs of setae, st 1 -st 4, and two pairs of poroids; shield incompletely formed, deeply excavated posteriorly, leaving setae st 5 on soft cuticle or on a separated platelet medially (Figs 60, 62). Form and extent of exopodal and peritrematal structures much as in female (Fig. 60). Opisthogastric region much as in female, with same complement of poroids but only four or five pairs of opisthogastric setae (JV 1, JV 2, ZV 2, JV 5, sometimes JV 3). All ventral idiosomatic setae smooth, similar in shape and length as female. Anal shield similar in form and structure as in female. Gnathosoma. Female and male. Gnathotectum with anterior margin convex, denticulate (Figs 52, 57). Chelicerae generally as described for the genus-group description. Fixed cheliceral digit sparsely multidentate (offset subapical tooth and six to nine teeth), with pilus dentilis reduced to alveolar remnant or indiscernible; movable digit of female tridentate, with a small pointed process on mid-ventral face (Fig. 44). Movable cheliceral digit of male with one small tooth, and lacking small process on ventral face; spermatodactyl digit-like, variously sinuous, directed anteriorly, not recurved basally (Figs 58–59); male chela lacking any arthrodial process on paraxial surface at base of movable digit. Corniculi of female and male similar in form, short, stout, inserted at level of insertions of setae hp 1 (Figs 51, 53, 61); paraxial process at base of corniculus slightly shorter than corniculus (Fig. 51); bifid internal malae elongate, extending nearly to or beyond apex of palpgenu (Fig. 53); apex of labrum extending to palptibia. Subcapitulum with a pair of pore-like structures at level of insertion of corniculi (Fig. 53). Subcapitular setae smooth, with hp 1 clearly longer and thicker than other pairs. Deutosternum with seven moderately narrow, multidenticulate rows of denticles, connected by lateral margins; basal two rows not narrowed, sometimes more finely multidenticulate. Palpal setation and form of setae as described for genus-group description (Fig. 54 a–c); palptrochanter with inner seta somewhat longer than external setae (54 a). Legs. Female and male. Legs moderately short, similar in length to dorsal shield but clearly shorter than idiosoma. Legs I and IV similar in length and thickness to legs II, III (Figs 63–70). Tarsus I with sensilla s inconspicuous, short and acute (Fig. 72 a), and with ventrodistal acuminate process extending under pretarsus (Fig. 72 b). Pretarsus of legs I to IV with slender claws and large, rounded pulvillus; those of legs II–IV with short, inconspicuous paradactyli formed as narrow sclerotised shafts flanking claws (Fig. 74). Legs II to IV with tarsus (excluding pretarsus) about twice as long as tibia. Tarsi II–IV with apical setal processes ad - 1, pd - 1 inconspicuous, shorter than short pretarsi, and with apical ventral process short, triangular, more retrorse in male than in female (Figs 73, 74). Setation and its ontogeny on segments of legs I to IV deficient from full complement of Melicharidae (as presented by Lindquist & Evans 1965 for Melicharini), coxae, 2 - 2 - 2 - 1; trochanters, 5 - 5 - 5 - 5; femora, 11 (2 2 / 1 2 / 2 2) – 9 or 10 (1 2 or 3 / 1 2 / 2 1) – 6 (1 2 / 1 2 /0 0) – 6 (1 2 / 1 2 /0 0); genua, 11 (2 3 / 1 2 / 1 2) – 8 (1 3 / 1 2 /0 1) – 7 (1 2 / 1 2 /0 1) – 7 (1 2 /0 3 /0 1); tibiae, 10 (2 2 / 1 2 / 1 2) – 7 (1 1 / 1 2 / 1 1) – 7 (1 1 / 1 2 / 1 1) – 7 (1 1 / 1 2 / 1 1); trochanter I seta ad vestigial, only alveolus discernible. In addition to setal absences noted for family: femur I, tibia I, and sometimes femur II lacking seta ad - 3; genu II lacking seta al - 2; however, genua II–III retaining av, and genu IV retaining pd - 3. Several setae moderately thickened, spinelike: ad - 1, pd - 1, pd - 2 on femora I–II and ad 1, pd - 1 on femora III–IV; ad - 2 on femur IV. Seta av on femur III and pv - 2 on trochanters I–IV (on female and male) attenuated. Other leg setae simple, not strongly differentiated on either sex (male leg setal dimorphism not developed). Deutonymph. Idiosomatic dorsum. Discoidal form. Dorsal shield poorly sclerotised, without lateral incisions, lightly lineated, with ca. 35 pairs of similarly short setae except more posterior ones smaller: 17 pairs (j 1, j 3 –j 6, z 2, z 4 –z 6, s 1, s 2, s 4 –s 6, r 2, r 3, r 5) on podonotal region, and a maximum 18 pairs (J 1, J 3, J 4, J 5, Z 2 –Z 5, S 1 –S 5, R 1 -R 6) on opisthonotal region (Figs 75); many of these setae displaced more marginally on dorsal shield than in adult. Ventrolateral soft cuticle nearly devoid of setae. Dorsal idiosomatic complement of poroids and gland-pores similar to that in female. Idiosomatic venter. Sternal region lightly sclerotised, unornamented, without endopodal extensions between coxae, with three pairs of platelets bearing sternal setae s 1 -st 3 and poroids iv 1, iv 2; setae st 4 -st 5 inserted on soft cuticle between coxae III and IV (Fig. 76); poroids iv 5 absent. Anal shield prominent, lightly sclerotised, unornamented, roughly rounded, with circumanal setae similar in length; adanal gland pores (gv 3) at level of paraanal setae; cribrum as in adult female. Opisthogaster with four to six pairs of ventral setae (the four larval JV 1, JV 2, JV 5, ZV 2, sometimes plus two minute JV 3, JV 4), sometimes not flanked by further setae on soft cuticle (Fig. 76). Peritrematal shields and peritremes shortened, paedomorphic, as in protonymphal instar, developed only alongside level between coxae III–IV (Fig. 76). Peritrematal gland pore gp 2 and poroid ip 2 on soft cuticle between levels of coxae II and III; poststigmatic poroid ips adherent to stigma on posterior edge of peritrematal shield, and poststigmatic gland pore gp 3 and poroid ip 3 on soft cuticle. Rim of exopodal plate behind coxa IV inconspicuous or indiscernible, with gland pore gv 2 not discernible. Gnathosoma. Gnathotectum, chelicerae and other mouthpart structures, corniculi and adjacent structures as in adult female; subcapitular and palpal structures similar to those in adult female. Legs. Pretarsal structures, and chaetotaxy of legs I–IV as in adult, including these additions to protonymphal setal complement: av - 2 on trochanters I–IV; al - 2 and pl - 2 on genu I, ad - 3 on genua I–II, pd - 3 on genu IV, pl - 1 on genu IV, av - 1 on genua II–IV, pv - 1 on genu II, al - 2 and pl - 2 on tibia I. Form and shape of leg setae similar to those on adult female, except seta pl on femur, genu, tibia, basitarsus of leg IV strongly developed and recurved/bent. Legs IV directed posteriorly, as in protonymph. Tarsi II–IV with form of setae and acuminate form of apical ventral process as in protonymph. Protonymph. Idiosomatic dorsum . Discoidal to ovoidal form. Dorsal shielding weakly sclerotised, with or without delineation between podonotal and pygidial shields, and with faint lateral, mostly podonotal ornamentation; mesonotal scutellae not evident (Figs 77, 89); dorsum maximally with 24 to 33 pairs of similarly short setae except more posterior ones smaller, J 5 minute (protonymphal setae s 5, r 2, r 3, r 5, Z 2, S 2, R 1, sometimes J 1 added to larval complement, j 2 absent); setae J 3, J 5, Z 4, Z 5 sometimes on pygidial shield if delineated (Fig. 89). Sometimes as many as eight pairs of dorsal deutonymphal setae (s 1, z 6, S 1, R 2 -R 6) may accelerate their ontogeny to this instar (Figs 77, 78). Body dorsum with complement of 11 pairs of discernible pore-like structures, of which six non-secretory (poroids) and five superficially appear to be secretory (gland pores). Idiosomatic venter. Sternal region with two or three pairs of weakly sclerotised platelets (without endopodal elements) bearing setae st 1, st 2 and poroids iv 1, iv 2 (iv 2 added to larval complement); setae st 3 on either a third pair of weak platelets or on soft cuticle; setae st 5 rudimentary, paragenital poroids indiscernible. Opisthogastric region reduced to small area behind coxae IV. Anal shield faintly sclerotised, unornamented, yet prominent, about as wide as long, with postanal seta longer than para-anal setae; para-anal setae inserted at mid-level of anal opening, roughly in transverse line with adanal gland pores; cribrum similar in development as in deutonymph. Opisthogaster with four or six pairs of ventral setae; JV 1, JV 2, sometimes ZV 2 anterior to anal shield; larval complement sometimes augmented by normally deutonymphal setae JV 3, JV 4 on posterior margin, along with larval JV 5 (Figs 78, 90). Peritrematal region on each side with poroid and gland pore (ip 2, gp 2) on soft cuticle at mid-level of coxae III, anterior to apex of weakly formed peritreme between coxae III and IV, and with poststigmatic poroid and gland pore (ip 3, gp 3) on soft cuticle behind peritreme. Exopodal rim and gland pore gv 2 behind coxa IV not discernible. Gnathosoma. Form of gnathotectum, corniculus and adjacent paraxial process, elongated internal malae (Fig. 85), and other gnathosomatic structures similar to those in deutonymph, except palpi with normal protonymphal complement of setae (see Evans 1964), including only one trochanter seta. Cheliceral movable digit bi- or tridentate, with mid-ventral projection rudimentary or indiscernible; fixed digit weakly multidenticulate, with offset subapical tooth and sparse (five or six) teeth along masticatory surface (Fig. 87). Legs. Legs I to IV with pretarsi, well-developed claws, and normal protonymphal complement of setae as described for Ascidae by Lindquist & Evans (1965) except for absence of second ventral seta on femur II: coxae, 2 - 2 - 2 - 1; trochanters 4 - 4 - 4 - 4 (1 0/ 2 1 on I, II, 1 1 / 2 0 on III, IV); femora 10 (2 2 / 1 2 / 1 2) – 7 (1 2 / 1 2 /0 1) – 5 (1 2 / 1 1 / 0 0) – 4 or 5 (1 2 /0 1 /0 0 or 1); genua 8 or 9 (1 2 or 3 / 1 2 / 1 1) – 6 or 7 (1 2 or 3 /0 2 /0 1) – 6 (1 2 /0 2 /0 1) – 5 or 6 (1 2 /0 2 /0 0 or 1); tibiae 8 (1 2 / 1 2 / 1 1) – 7 (1 1 / 1 2 / 1 1) – 7 (1 1 / 1 2 / 1 1) – 7 (1 1 / 1 2 / 1 1); normally deutonymphal seta pl - 1 sometimes accelerated in appearance on femur and genu IV, and ad - 3 similarly on genua I and II. Coxae I–II distal rims with serrated posteroventral strips. Leg setae generally simple. Legs IV directed posteriorly, with seta pl on femur, genu, tibia, and to less degree on tarsus IV slightly to clearly thicker and longer than other setae of segments (Fig. 79). Tarsi II–IV with setae ad - 1, pd - 1 small but as long as short pretarsus; md, al - 2, av - 1, pl - 1 and pl - 2 longer, thinner than other tarsal setae, except av - 2 similarly attenuate on tarsus IV; apical ventral tarsal process acutely triangular. Larva. Idiosomatic dorsum . Body with dorsal shielding weakly sclerotised, not clearly delimited, less discernibly ornamented than on nymphal instars; mesonotal scutellae indiscernible (Figs 80, 91). Body dorsum with 17 pairs of setae: nine podonotal pairs, and eight opisthonotal pairs (J 2); discernible pore-like structures include eight non-secretory poroids (three podonotal, five opisthonotal) and three apparently secretory gland pores (two podonotal, one opisthonotal). Peritrematal region on each side with gland pore gp 1 at level of setae j 3, gland pore gp 2 and poroid ip 2 at level above coxae II–III, and poststigmatic gland pore gp 3 and poroid ip 3 at level of anal shield anterior margin. Idiosomatic venter. Tritosternum normally developed, similar in form as in subsequent instars. Sternal region with two or three pairs of weakly (hardly discernible) sclerotised platelets bearing setae st 1, st 2 and poroids iv 1; setae st 3 on soft cuticle or on a third pair of platelets; sometimes protonymphal setae st 5 accelerately present between coxae III (Figs 81, 92); pair of subcutaneous structures sometimes discernible at level of posterior margins of coxae III. Anal shield faintly sclerotised, elongate or as wide as long; para-anal setae inserted at level slightly posterior to anal opening, and slightly longer than postanal seta; adanal gland pores (gv 3) at level of postanal seta; cribrum rudimentary or absent. Venter with six pairs of opisthogastric setae on soft cuticle around anal shield, including JV 1, JV 2, sometimes rudimentary ZV 2, JV 5 anterolaterally and ventrally displaced S 4, S 5 posterolaterally (sometimes JV 5 and S 4 or S 5 absent). Gnathosoma. Gnathotectum with anterior margin less denticulate than in subsequent instars (Fig. 83). Cheliceral movable digit with two teeth, and a rudimentary mid-ventral projection (Fig. 84); dentition of fixed digit weakly developed, with offset subapical tooth and masticatory ridge with ca. two to four weak teeth. Form of elongated internal malae, corniculus and adjacent acuminate process much as in subsequent instars (Fig. 82); deutosternum with rows of denticles similar to those in nymphs. Palpus with normal larval complement of setae (see Evans, 1964); palptrochanter nude. Legs. Legs I to III with pretarsi, well-developed claws, and with normal larval complement of setae as described for Ascidae by Lindquist & Evans (1965) except for retardation of appearance of seta ad - 2 on femur II until protonymph: coxae, 2 - 2 - 2; trochanters 4 - 4 - 4; femora 10 (2 2 / 1 2 / 1 2) – 7 (1 2 / 1 2 /0 1) – 5 (1 2 / 1 1 /0 0); genua 8 (1 2 / 1 2 / 1 1) – 6 (1 2 /0 2 /0 1) – 6 (1 2 /0 2 /0 1); tibiae 8 (1 2 / 1 2 / 1 1) – 7 (1 1 / 1 2 / 1 1) – 7 (1 1 / 1 2 / 1 1). Coxae I- III distal rims smooth. Leg setae generally simple, not markedly differentiated. Tarsi II and III with setae ad - 1, pd - 1 as long as short pretarsus; apical ventral tarsal process acutely triangular. Etymology. The name of the genus, feminine in gender, is a Latinised combination of the term hispidus, meaning bristly, upon which the family-group name of the beetles is based, and iphis, meaning stalwart, and a suffix commonly used among names for gamasine mites. Distribution and habitats. The genus is currently based on one newly described species and on additional material representing five undescribed species. All instars of mites of the genus Hispiniphis are known only as subelytral symbionts of adult hispine beetles of the genus Cephaloleia, tribe Immatidiini (or Cephaloleiini). Over 200 species of the neotropical genus Cephaloleia are recognised, including at least 40 sympatric species from lowland tropical rain forest in the La Selva Biological Station, Costa Rica, alone; larvae of the majority of species feed on unfurled leaves of various families of the Order Zingiberales (Staines 1996, 2008, 2009, Garcia-Robledo et al. 2010), Staines & Garcia-Robledo 2014). Remarks. Reduction in size of opisthosoma. Among species of Hispiniphis for which specimens of the nymphs are available, a smallness of the opisthosomatic region is notable (Figs 78, 90). There is a compactness of body behind the legs IV in both nymphal instars, leaving the anal shield and adjacent structures squeezed into a relatively small space. Perhaps this may be correlated with evolution towards a rapid life cycle, and lessened need for enlargement of the body region not involved in mobility. This reduction is not evident in the nymphs of the one known species of the sister-genus, Makarovaia. Interspecific variation in adult dorsal shield attributes. The shape and ornamentation of the adult dorsal shield, as well as the complement and size of its setae, vary markedly among the five species of Hispiniphis for which we have material at hand (Fig. 88). In general, the description of the genus accounts for these differences, until such time as other of these species are described subsequently. Such readily discernible morphological differences may be quite useful in determining whatever level of host specificity there may be between species of this genus of mites and their beetle hosts. Symmetry of dorsal idiosomatic hypertrichy. Adults (to some extent preceded by nymphs) of