Scaralina Yanega

Yanega, Douglas, Goemans, Geert, Dam, Matthew Van, Gómez-Marco, Francesc & Hoddle, Mark, 2024, Description of a new genus of North and Central American planthoppers (Hemiptera: Fulgoridae) with fourteen new species, Zootaxa 5443 (1), pp. 1-53 : 5-8

Scaralina Yanega , gen. nov.

Type species. Calyptoproctus marmoratus Spinola, 1839 , by present designation.

Etymology. As these species are intermediate in many features between the related genera Scaralis and Alphina , the name is a portmanteau of these two generic names. For purposes of gender agreement, the name should be considered feminine.

Diagnosis. Species of this taxon are most similar to those of Scaralis Stål , especially in the reduced femoral ridges and mesocoxal spines, as well as numerous minor details of the head and thoracic structure and sculpture, mostly not shared by other genera (other than Jamaicastes ), such as the pattern of notal carinae, the long rostrum, and the broad lateral lobes of the lower frons. There are, however, several characters that together distinguish all members of Scaralina from at least the type species of Scaralis ( S. picta (Germar, 1830)) and many of the other included species we have been able to examine ( S. corallina (Gerstaecker, 1860) , S. neotropicalis (Distant) , and S. obscura (Distant) ; also see below). Most reliable among these features in Scaralina are the following: the second antennomere (small and globose in Scaralina versus large and asymmetrically ovoid, larger than the antennal socket in Scaralis ( Fig. 7 )); the more elongate clypeus (usually roughly twice as long as wide, and reaching the apex of the forecoxae); the forewings without an arcuate impressed nodal line at the base of the apical hyaline portion (very well-defined in Scaralis s.s.); the distal forewing membrane patterned with irregular maculations and variable venule coloration (membrane hyaline, evenly-tinted, or gradually shading, with venules uniform or very gradually shading in almost all Scaralis ); a projecting, abbreviated, and concave trapezoidal versteifung with a strongly angulate proximal margin (in Scaralis the “versteifung” is lower and sometimes elongated, weakly concave, and more rounded at the proximal end; compare Figs 4–5 ); the prominent leg markings (contrasting transverse bands on fore- and mid-femora and tibiae); the numerous black granular sublateral pits on the abdominal terga (these pits concolorous with the surrounding cuticle in Scaralis , and fewer in number); the male gonostyli more visibly enclosing the aedeagal apex, with a setose bulge at the base of an incurved dorsal surface (in Scaralis only the extreme apex of the aedeagal complex is sometimes enclosed dorsally, and the setose bulge is very small and approximates the inner margin of the gonostyle ( Figs 8, 9 )); the fine but very readily visible pubescence on the dorsal thorax, in particular (the dorsal thorax is usually bare in Scaralis , as are the wing veins, or at most with barely visible short, fine setae). Several more variable or occasionally unreliable features can be added to this list, for distinguishing Scaralina from Scaralis (Scaralis) : the sub-ocular lobes (nearly absent in most Scaralina , versus distinct and rounded or subacute in most Scaralis ); the well-defined and contiguous carinae of the mesonotum in Scaralis (no species of Scaralina has all of these carinae well-defined for their entire length; one or more are reduced to low ridges or entirely obsolescent, at least in part); the reduced female supra-anal plate in Scaralina (absent or only partially concealing the anal tube versus completely covering it; in genera such as Scaralis this plate is typically at least three times the length of any of the preceding tergites); the deep punctures and/or wrinkles in the dorsal and lateral faces of the pronotum in Scaralis s.s.; the red or orange coloration of the hindwing bases (only two Scaralina have blue coloration basally, while Scaralis typically do); the greatly reduced wax production in Scaralina (in Scaralis , the face and pleura often have large areas bearing wax pollinosity ( Fig. 103 ), and the spiracles are often entirely occluded by wax, plus a distinct mass of wax on either side of the terminalia in females); the hyaline or weakly infumate anal region of the hind wing (strongly infumated with pale venation in Scaralis s.s.); the broadly rounded apical concavity of the male anal tube (generally deeply notched in Scaralis s.s.; compare Figs 10–11 , 23 ). Despite some variability, the combined list of such features that differ between the two groups is enough, we believe, to justify a generic-level separation, and we further believe this group as defined here is likely to be monophyletic; nonetheless, it would not be surprising to discover (e.g., if and when a thorough molecular phylogenetic analysis with all of the South American species is performed) that Scaralina as here defined render Scaralis paraphyletic, or the converse, in which case the two groups may need to be relegated to subgenera within a more inclusive genus Scaralis . Scaralina chapina and S. monzoni , in particular, both described herein, show more intermediate features than any of the other taxa (e.g., these species have the clypeus only slightly longer than broad, have blue hindwing bases, with the forewing basal markings extending to the point where RP deviates from ScP+RA, and lacking translucent basal areas in the forewings; chapina also has the male gonostylar hooks short and thickened as in Scaralis ; compare Fig. 9 and Fig. 83 ), and were it not for these two species, the separation of the two genera would be far more definitive.

Composition. Included species (15): Scaralina aethrinsula Yanega & Van Dam, sp. nov.; S. chapina Goemans & Yanega, sp. nov.; S. cristata Yanega & Van Dam, sp. nov; S. durango Yanega. sp. nov.; S. gigantea Yanega, sp. nov.; S. hawksi Yanega, sp. nov.; S. marmorata ( Spinola, 1839) ; S. metcalfi Yanega & Van Dam, sp. nov; S. monzoni Goemans & Yanega, sp. nov.; S. obfusca Yanega, sp. nov.; S. obrienae Yanega & Van Dam, sp. nov; S. orientalis Yanega, sp. nov.; S. rileyi Yanega, sp. nov.; S. sullivani Yanega, sp. nov.; S. veracruzensis Yanega & Van Dam, sp. nov.

Description. Head: Head broader than long, not projecting. Vertex roughly 4 times as wide as long, demarcated on all sides by lamellate rim, produced ventrolaterally into rounded supra-ocular lobes, which are appressed against dorsoposterior eye margin; posterior rim reflexed anteriorly. Sub-ocular lobes absent (or very weak and rounded). Vertex with two pairs of posteromedial spots (e.g, Fig. 47 ), which may or may not contrast with the surrounding color (so they may not be evident), but of a distinctly differing texture (dull and granular) from the surrounding cuticle to be detectable; the pair nearer the midline are transversely ovoid, and well-defined, outer spots larger, more variable in shape, and irregular, with less definite edges. Frons roughly rectangular (wider than long), broadly flat or very gently convex for most of its surface; typically smoother and sometimes shallowly impressed below upper portion, often very finely rugose below and without well-defined ridges, though a few species (e.g., Figs 31 , 39 ) show a medial ridge and/or converging lateral ridges; the lower margin just above the clypeal suture is sometimes thickened into a low transverse ridge; lateral margins of frons vary from nearly straight and subparallel to slightly concave and divergent below, with lower lobes bearing a depression (sometimes distinctly expanded at corners), which is impressed and often bearing a small translucent window in the depression (e.g., Fig. 34 ); frons reflexed along upper portion to form an appressed flap, typically at an obtuse angle relative to the plane of the lower frons, delimited above by strong transverse crease, reflexed portion defined laterally by secondary oblique crease a short distance below upper corner of flap, and with the dorsal reflexed portion commonly having distinct, closely-spaced vertical wrinkles; individual specimens sometimes possess small incomplete interstitial ridges (partial flaps) between the upper lateral edge of the transverse frontal crease and the anterior rim of the vertex (these may be remnants of an embryonic frontal sac). Clypeus elongate triangular (distinctly longer than broad, typically at least 1.5 times as long, reaching to apex of forecoxae), nearly straight along sides, surface fairly smoothly convex but often very shallowly concave near midline, with distinct, parallel-sided lateral areas; upper margin varies from almost straight to distinctly dorsally arcuate, sometimes very slightly indented medially. Rostrum with 3 visible segments, middle segment longest, typically extending well beyond metacoxae, the tip of the rostrum reaching almost to the hind femoro-tibial joint or beyond ( Fig. 2 ); in most species, mostly pale except apical segment, and anterior (ventral) ridges also somewhat darkened. Antennae short, base concealed in protruding, pale socket; basal antennal segment often darkened dorsally, otherwise pale; second segment small (diameter approximately equal to antennal socket diameter or very slightly larger), nearly spherical to slightly pyriform, symmetrical ( Fig. 32 ), typically dark, with pale circular to ovoid sensillae; stylus apical, elongate, dark, with a small basal bulbosity, arising from slight indentation. Eyes prominent, roughly circular in outline; lateral ocelli in angle between frons and lower eye margin, translucent yellowish.

Thorax. Dorsal and lateral surfaces with very fine and slightly scattered pale erect pubescence, generally longest on pleura and ventral sclerites. On the notum of nearly all species these setae are longer than the interspaces between setae, and easily seen (especially in S. rileyi ( Fig. 3 )), but the type species, S. marmorata , has diagnostically shorter setae; the density is similar to other species, but shorter and slightly recumbent, making them difficult to observe unless viewed from an oblique angle. Pronotum elongate (more than 2x length of vertex along midline), anterior margin concave, following contour of head, disc with a distinct medial carina, sometimes projecting and sub-lamellate; with a sharply carinate lateral anterior rim that starts near the inner eye margin, and continues to near the middle of the tegula, below the eye; there is an oblique carina just ventral to this, continuing onto the ventral posterior lobe of the pronotum, which approximates the forecoxal bases and partly overlaps the anterior face of the mesepisternum; posterior margin of pronotum usually with angulate medial notch, and commonly a low transverse ridge immediately preceding posterior margin. Pronotum with paired, dark granular spots similar to those on vertex; a somewhat pit-like anteromedial pair, and a lateral post-ocular pair (at the indentation immediately behind the outer posterior edge of the eye), and often with fine transverse wrinkles in the posterior half. Mesonotum at midline about equal in length to the vertex and pronotum combined, bearing low longitudinal medial and lateral carinae, the latter sinuate to varying degrees, ending posteriorly at the outer edges of a pair of small dark posterior granular spots (the inner edges of which may sometimes be bordered by a weak inner posterior carina that starts at the terminus of the lateral longitudinal carina), and the inner edges of much larger granular spots (dark at least in part) which are variously elongated anteriorly; the lateral carinae at the extreme anterior edge are strongly angled inward and converge at the midline, but these anterior “arms” are hidden underneath the posterior pronotal margin, and thus seldom visible (they may be exposed if the pronotum is somewhat depressed); there are short transverseoblique ridges located laterad of the lateral carinae, but these become obscure before attaining the lateral carinae; the medial carina becomes obscure posteriorly, ending in an upward-angled semi-acute lobe that is typically creased or wrinkled transversely at its base. Mesopleuron delimited from mesosternum by a strongly and fairly evenly arcuate ridge running from the upper posterior corner of the procoxal membrane to the upper anterior corner of the mesocoxa; the degree of curvature of the ridge reduces gradually from anterior to posterior. Metacoxa with a prominent dorsoposterior spine, this spine absent or reduced to a minute pointed tubercle on the mesocoxa. Femora weakly convex on anterior face, more strongly so on posterior face, with only two well-defined longitudinal setose ridges, along the anterior dorsal and ventral edges. Fore- and mid-legs virtually identical in color pattern: anterior coxal face typically with two dark spots which may coalesce; femora with at least two black bands, the basal one largest, and sometimes with a very small third apical or subapical band or mark; tibiae with three black bands, the apical one generally largest, often but not always reaching the apex; pro- and mesotarsi black except for pale dorsal mark on third tarsomere; metafemur typically with one black band near apex, metatibia with three dark bands and four or five lateral spines, metatarsi with first two tarsomeres mostly pale except dorsally at base, third tarsomere generally matches those of other legs.

Wings. Forewings without evident nodal line; “versteifung” posteriorly concave, abbreviated and strongly protruding, angulate at least on proximal margin, trapezoidal to almost triangular ( Fig. 5 ); pigmented portion of forewing (excluding postcostal cell) extending over less than half the length (ending near the claval juncture). Distal portion of forewing with membrane faintly to strongly maculated, never entirely hyaline, and venules varying from light to dark, never uniform in color. Nearly all species with the basal forewing markings divided into a paler basal portion and a darker distal portion that forms a band from subcosta to clavus, near the first branch of MP (about 1/3rd of the wing); the gap between MP and CuA is almost always less than the gap between it and ScP+R. Venation relatively coarse, with relatively few branches and relatively few crossveins compared to most poiocerine genera. In most species, the crossveins are simple and define square to rectangular cells, though there are typically some reticulated venules in the radial cell (between the base of MP and ScP+R) and in the anal region (especially between Pcu and A 1); a few species show reticulated venules elsewhere, such as the postcostal cell, which in most species has simple crossveins or may be nearly devoid of crossveins. Hindwings with colored bases, typically red or orange (rarely pale blue), membrane of anal region entirely clear to slightly infumate.

Abdomen. Most species with tergites and pleurites predominantly dark, and sternites predominantly pale; dorsal pale markings, when present, most commonly yellow to orange, though sometimes red, and sometimes grayish-buff. Spiracles large, one pair dorsal at base of abdomen ( Fig. 6 ), others in the dorsal pleurites. Terga 3–6 typically with small, black, pit-like lateral depressions with granular texture; typically arranged in two sets on each side, an anterior set of 3–4 pits and a posterior set of one large pit and one very small pit ( Fig. 6 ). Female tergum 6 often produced into a supra-anal plate, but generally not entirely concealing anal tube; at most twice the length of any of the preceding terga, but unmodified in two species ( aethrinsula and durango ), where terga 5 and 6 are nearly identical in length. Wax production evidently limited, typically a faint residue at the base of the genitalia, and traces in spiracular apertures.

Male terminalia. Apico-dorsal portion of the gonostyli very strongly incurved and largely enclosing the apex of the aedeagal apparatus, in most species nearly parallel at the midline for most of the apical half. The outer inflection of the incurved portion marked by a swollen region with numerous sharp, short, bristle-like setae (the “setose bulge”); basal hook of the gonostyle located anterior to this swelling, the hook variably developed but generally small, recurved, and sharply acute, with a well-developed flap-like fold at the base. Inflatable aedeagal elements (in inflated preparations) with sac-like dorsal and ventral lobes on both sides, and medial outcurving horn-like lobes with slightly sclerotized inner basal edges and faintly sclerotized apices. Both sexes with anal tube somewhat heartshaped in dorsal view, widest near middle (in female) or past middle (in male) and narrowest at base; in female strongly compressed and only very slightly convex on upper and lower surfaces, in male more convex dorsally; margin of apical aperture broadly concave, semicircular, exposing pygofer ( Fig. 23 , resembling also Fig. 11 ).

Figures

FIGURE 2. Ventrolateral view of rostral apex, Scaralina cristata; note that tip of last rostral segment extends beyond the femorotibial joint of the hind legs when the hind legs are pointed straight back.

FIGURE 3. Mesonotum of Scaralina rileyi showing long dorsal pubescence.

FIGURE 6. Dorsum of abdomen (terga 2–7) of female Scaralina aethrinsula, showing dark granular sublateral pits and unmodified tergum 6. Anal tube in this specimen is angled upwards and foreshortened.

FIGURE 7. Antenna of Scaralis (Alphinoides) nr. semilimpida; arista not visible.

FIGURES 4–5. Versteifung, underside of left forewing, viewed from posteroventral angle in both photos, so wing base is to the left; (4) Scaralis (Alphinoides) semilimpida; (5) Scaralina obrienae.

FIGURES 8–9. Male genitalia of Scaralis (Scaralis) sp.; (8) dorsal view, anal tube removed; (Fig. 9) lateral view.

FIGURES 99–104. Faces of different genera of Poiocerini. (99) Calyptoproctus sp.; (100) Hypaepa nr. zapotensis; (101) Jamaicastes basistella; (102) Matacosa miscella; (103) Scaralis sp.; (104) Scaralina obfusca (female).

FIGURES 10–11. Male anal tubes, dorsal view; (10) Scaralis (Scaralis) sp.; (11) Scaralis (Alphinoides) inbio.

FIGURES 21–26. Habitus photos of Scaralina species. (21) S. monzoni (female); (22) S. obfusca (type male); (23) S. obrienae (male); (24) S. orientalis (male); (25) S. rileyi (male); (26) S. sullivani (female).

FIGURES 82–90. Male genitalia of Scaralina species, lateral view. (82) S. aethrinsula; (83) S. chapina; (84) S. cristata; (85) S. marmorata; (86) S. metcalfi; (87) S. monzoni; (88) S. obrienae; (89) S. orientalis; (90) S. rileyi.

FIGURES 42–50. Heads and nota of Scaralina species. (42) S. aethrinsula; (43) S. chapina; (44) S. cristata; (45) S. durango; (46) S. gigantea; (47) S. hawksi; (48) S. marmorata; (49) S. metcalfi; (50) S. monzoni.

FIGURES 27–35. Faces of Scaralina species. (27) S. aethrinsula; (28) S. chapina; (29) S. cristata; (30) S. durango; (31) S. gigantea; (32) S. hawksi; (33) S. marmorata; (34) S. metcalfi; (35) S. monzoni.

FIGURES 36–41. Faces of Scaralina species. (36) S. obfusca (male); (37) S. obrienae; (38) S. orientalis; (39) S. rileyi; (40) S. sullivani; (41) S. veracruzensis.