Erysivena, Symonds & Cassis, 2018

Symonds, Celia L. & Cassis, Gerasimos, 2018, Systematics And Analysis Of The Radiation Of Orthotylini Plant Bugs Associated With Callitroid Conifers In Australia: Description Of Five New Genera And 32 New Species (Heteroptera: Miridae: Orthotylinae), Bulletin of the American Museum of Natural History 2018 (422), pp. 1-229 : 147-154

Erysivena , new genus

TYPE SPECIES: Erysivena kalbarri , n. sp., by original designation.

DIAGNOSIS: Defined by the following characters: small to medium size; subovate to elongate; (faded) yellowish-green coloration, cuneus with red tip in most species, forewing membrane with subcuneal clear spot, membrane veins red (fig. 14); pygophore with short dark spinelike setae proximate to genital opening on ventral surface (fig. 58A, B); pygophore dorsal margin moderately to strongly concave, with one, two, or three (elongate) sclerotized tergal processes (figs. 51H, 57H, 63H, 71H); left lateral tergal process always present, elongate and serrate; left paramere L-shaped, moderately (figs. 52B, 53B) to strongly expanded (figs. 72A, 73B) medially, apophysis short or elongate (e.g., figs. 73B, 69B), apex hooked or unhooked (e.g., figs. 50B, 49B); right paramere shape more variable, C-shaped (fig. 49C) or hammer shaped (fig. 50C), widest medially or subapically, mostly with some variation of an expanded medial flange on inner dorsal margin or broad lobe on dorsal margin; right paramere apex short or elongate, curved and mostly serrate; phallotheca dark and heavily sclerotized, often narrow or flattened distally, not always capped by proctiger (figs. 49D, 50D, 51H, 53E, F); in situ, phallotheca projected beyond parameres from genital opening, left paramere situated along majority of (fig. 58B) or over ventral margin of pygophore (fig. 52B), and right paramere situated mostly external and right lateral to phallotheca with curved apex meeting sensory lobe of left paramere (figs. 51H, 52A, B, 57H, 58A, B); aedeagus with three sclerotized, elongate endosomal spicules (fig. 11); proximal endosomal spicule (PES) positioned ventral (or left ventrolateral), second dorsal endosomal spicule (DES2) left dorsolateral to secondary gonopore, and first dorsal endosomal spicule (DES1) left lateral or dorsad to DES2; PES and DES2 associated with secondary gonopore, while DES1 is removed from secondary gonopore and adjacent to DES2; PES unbranched, with or without medial (figs. 56F, 64D, 54E) or subbasal (fig. 53H, 58C, 59E) process(es), constricted above base (figs. 52C, 53G) or more uniformly tapering to apex (fig. 54E, 72C), with smooth or serrate distal margins; DES2 unbranched or bifurcate in distal half, serrate distally and medially (figs. 49F, 50F, 52E); DES1simple and unbranched with smooth or serrate margins (figs. 58E, 59E, 64C, 65C), or complex with serrate bifurcate apex and processes (figs. 52E, 53G), with basal keel situated adjacent to DES2 (fig. 49F); female vestibulum heavily sclerotized (60A, 61A, B); ventral labiate plate with spiniferous lateral lobes (figs. 60C, 61A); large spiniferous mediolateral lobes on dorsal labiate plate (figs. 60B, 61B); interramal sclerites deeply divided with V-shaped junction, with two pairs of interramal lobes, separated at base (figs. 60D, 61C).

DESCRIPTION: Male: Macropterous, small to midsized, ovate-elongate, sides parallel or slightly round, body length 2.84–4.19 mm.

COLORATION: Dorsal aspect yellowish green, or bright green, often faded to brownish yellow as specimens age, sometimes with bright green patches of pigmentation on lateral margins of pronotum, on hemelytra above and below costal fracture, and on lateral margins and tip of cuneus; pronotum sometimes with anterior half yellow and posterior half bright green, anterior margin sometimes lighter yellow or pale dusty green; mesoscutum and scutellum often faded to yellow; cuneus tip bright red to dark red, sometimes just with slight hint of red; sometimes extending from tip to basal inner half; forewing membrane translucent silvery gray-brown, light with darker patches laterally or dark and more uniform all over, clear spot present below cuneus (fig. 12B), veins bright red; venter yellowish green, often faded to yellow; coxae and adjacent plate structures yellow; legs mostly faded to yellow with little green pigmentation, tarsi darkened; ethanol-preserved specimens faded to white (fig. 14). SUR- FACE AND VESTITURE: Dorsum smooth, weakly polished; sparse to moderately dense covering of simple setae, mostly pale or sometimes darker medium brown, semierect; setae sometimes more bristlelike on head and along lateral dorsal margins; antennae with dense distribution of short, pale simple setae, antennal segment I also with few elongate erect bristlelike setae (approx. 3 or 5); pygophore, ventral surface proximate to genital opening with short spinelike setae, medium brown or black, adpressed; femora with dark, simple setae and some longer, erect spinelike setae; tibiae with pale simple setae, hind tibiae with several rows of minute dark spinulae, tibial spines light brown. STRUCTURE: Head: Subovate in lateral view; weakly or strongly expanded anteriorly. When weakly expanded anteriorly, frons sloping downward, mandibular and maxillary plates and clypeus not enlarged (e.g., fig. 63A, B), but when strongly expanded anteriorly, frons strongly protruding beyond anterior margin of eyes, mandibular and maxillary plates and clypeus large (e.g., figs. 51A, B, 57A, B, 71A, B). Maxillary plate subovate, anterodorsal margin round and protruding beyond mandibular plate; clypeus prominent, round or slightly pointed in dorsal view, dorsal edge round in lateral view; vertex flat; eyes medium or large. When eyes medium, occupying less than threequarters of head height, weakly exerted beyond outline of head, not or very slightly extending past anterolateral angle of pronotum (e.g., figs. 51A–D, 57A–D), but when eyes large, occupying most of head height, exerted moderately or greatly beyond outline of head, extending well past anterolateral angle of pronotum (e.g., figs. 63A–D, 71A–D). Posterior margin of eyes con- tiguous with posterior margin of head; labium midlength and extending to meso- or metacoxae or elongate and extending beyond metacoxae, over abdomen. Antennae: Elongate, inserted just above ventral margin of eye, insertion contiguous with or slightly removed from anterior margin of eye (figs. 51B, 57B); AI short, between 0.9–1.3× vertex width, subcylindrical, slightly wider than remaining segments; AII moderately elongate, on average 1.25× pronotal width; AIII, average 1.5× AIV; AIV, subequal or less than AI; AIII and AIV slightly thinner than AII. Pronotum: Trapezoidal, transverse, moderately short, anterior margin weakly concave, collar reduced to thin lip; callosite region undefined; lateral margins straight, angled 30°–45° to midline; humeral angles round; posterior margin very weakly sinuous to straight (figs. 51C, D, 57C, D, 63C, D, 71C, D). Mesoscutum and scutellum: Slightly raised; mesoscutum exposed, one-third length of scutellum (fig. 14). Metathorax: Metathoracic spiracle moderately elongate, narrowly subcylindrical, with evaporative area narrowly spanning anterior margin and expanded slightly dorsally (figs. 51E, 57E, 63E, 71E); metathoracic scent gland with ostiole subovate to lanceolate, peritreme subovate to subtriangular, extending weakly or strongly onto posterior ostiole margin, evaporative area with small subovate to elongate evaporative bodies (figs. 51E, F; 57E, F; 63E, F, 71E, F). Hemelytra: Not greatly elongate, abdomen extending to middle of cuneus; lateral margins parallel to slightly round; cuneal fracture well defined; cuneus short to moderately elongate; forewing membrane major cell mostly elongate, major membrane vein round or straight, curved or parallel to inner margin of cuneus (fig. 14). Legs: Elongate, femora slightly flattened, hind femur moderately incrassate; pretarsus with moderately long claws, pulvilli moderately sized, fleshy parempodia broad, apically convergent (figs. 51G, 57G, 63G, 71G). GENITA- LIA: Pygophore: Short, transverse (wider than long), narrowing slightly posteriorly (figs. 51H, 53A); genital opening large, round (figs. 52A, 53A, 5bA); dorsal margin of genital opening strongly concave, sometimes slightly asymmetrically (figs. 51A, 53A); one, two, or three sclerotized, elongate, serrate tergal processes, usually lateral to midline (figs. 51H, 57H, 71H); left lateral tergal process always present, variously shaped (figs. 49A, 53A, 54A); second tergal process positioned right lateral when present, linear shape, sometimes with an expanded, lobed base (figs. 49A, 50A); third tergal process positioned medially, linear, paired with right lateral tergal process, when present (figs. 56A, 71H); far right lateral tergal lobe with smooth margins, sometimes present, either broad posterior facing (figs. 51H, 53A) or elongate and inward facing (figs. 71H, 72B, 73A); ventral margin straight to weakly concave (figs. 58B, 64B), sometimes asymmetrically (e.g., fig. 72A); narrow, round, phalloguide present, extending from ventral margin, modified ventrad to right paramere articulation as sclerotized ridge (figs. 52B, 53A) or an expanded elongate lobe (figs. 72A, 73A); left paramere wrapped along (just inside) ventral pygophore margin at rest (e.g., figs. 58B, 64B, 72A), rarely directed anteriorly across outer ventral surface of pygophore at rest (e.g., fig. 52B); right paramere mostly visible externally at rest, usually situated right lateral to phallotheca and aedeagus and curved round to meet left paramere medially under phallotheca (e.g., figs. 51H, 52B, 57H, 58B, 63H, 64B, 71H, 72B); phallotheca and aedeagus visible externally, extending beyond parameres posteriorly (e.g., figs. 63H, 64A, B); where left paramere is greatly expanded, phallotheca held cupped inside (e.g., figs. 71H, 72A, B); often without capped proctiger over phallotheca (e.g., figs. 51H, 52H, 63H, 64A); proctiger sometimes present covering phallotheca (e.g., figs, 57H, 58A) or present but removed from phallotheca (e.g., figs. 71H, 72B). Left paramere: L-shaped; moderately (figs. 52B, 53B) to strongly expanded and subtriangular medially (figs. 72A, 73B); sensory lobe broad and round (e.g., figs. 58B, 59B) or more prominently expanded (figs. 64B, 65B), outer margin sometimes with an angular edge; sensory lobe and outer surface with pale simple setae, sometimes also with short, light to dark brown, bristlelike setae on sensory lobe (figs. 64B, 65B, 66B); apophysis moderately to greatly elongate, narrowing to apex, inner margin sometimes with small spines or serrations (figs. 53B, 54B, 62B, 65B), ca. <90° to rest of paramere; apex, either round and curved inward but without hook (figs. 49B, 52B, 53B), or hooked (figs. 50B, 54B). Right paramere: Shape variable; C-shaped, curved from apex round to medial flange on dorsal inner margin (e.g., figs. 49C, 53C, 56C, 59C, 73C) or “hammer” shaped with expanded subapical lobe on dorsal margin and elongate apex (figs. 50C, 62C, 65C, 66C); medial flange present on dorsal inner margin expanded to differing degrees, from broad, weakly expanded lobe or serrate edge on dorsal margin (figs. 49C, 50C, 53C), or an expanded angular flange (fig. 54D, 68C), often constricted to point (fig. 56C, 59C); medial flange on dorsal margin projected inward or on inner lateral margin projected inward and downward, sometimes greatly reduced; apex generally on inner ventral margin, curved inward, weakly to strongly curved, sometimes greatly expanded/elongate; medial flange and apex mostly with serrate margins or small spines, rarely smooth; with or without simple setae. Phallotheca: Darkly sclerotized; protruding moderately from pygophore and visible externally in repose (figs. 52A, 64A), not always extending beyond parameres (figs. 58A, 72B); dorsal opening small (distal only) (e.g., fig. 50D) or large (distal to medial) (e.g., fig. 49D); distally round (fig. 49D) or slightly acuminate (fig. 50D); subapical ventral surface mostly with single tumescence (fig. 54D); distally narrow, without or with slight (e.g., fig. 59D) or full (e.g., fig. 53F) lateral compression (to form flat crest); small opening phallotheca, opening round, drop shaped with small point at apex or subovate sometimes with lobe on basal margin (figs. 50D, 56D); large opening phallotheca, right dorsal margin of genital opening with or without medial (or subbasal) lobe (figs. 49D, 69D). Aedeagus: Secondary gonopore elongate, cylindrical, membranous, and faint; three elongate, sclerotized endosomal spicules (fig. 11); proximal endosomal spicule (PES) ventral or left ventrolateral to secondary gonopore; second dorsal endosomal spicule (DES2) dorsad to left lateral of PES and secondary gonopore; PES and DES2 associated with secondary gonopore, but first dorsal endosomal spicule (DES1) is removed from secondary gonopore and adjacent (left lateral or dorsad) to DES2; all spicules originating in line with base of secondary gonopore or slightly distad to it; PES wrapped sheathlike around secondary gonopore, unbranched, with simple or complex forms. With simple PES (e.g., figs. 64C, D, 74C, D), tapering evenly from base to apex, with or without medial process, distally smooth or serrate, sometimes medially serrate (e.g., figs. 56F, 50F, 65E), or with complex PES (e.g., figs. 52C, 58C), base very broad, then weakly (e.g., fig. 49E) or acutely (e.g., fig. 53G) constricted and narrow and ribbonlike to apex, with or without subbasal straplike process attached just above constriction point (figs. 58C, 59E), expanded base with small, very fine process present on ventral margin proximate to apex of secondary gonopore (figs. 58C, 59E), distally smooth (fig. 67E) or weakly serrate (fig. 49E); complex PES sometimes joined to DES2 by membrane at base on right lateral side of secondary gonopore (figs. 49F, 52E, 53G, 58E, 59E, 67E). DES2 unbranched or bifurcate, sometimes expanded medially, distally serrate, serrations extend from apex to midsection on right side margin (figs. 52E, 64E), rarely with small row of serrations on right side margin more subbasally (fig. 62E), sometimes twisted or curved downward distally (figs. 65E, 68F), unbranched form sometimes with small medial process (figs. 69E, 70E, 72D); DES1 with simple or complex form: with simple DES1 (e.g., figs. 58E, 59E, 64C, 69C, 72C, 73G), unbranched, sometimes expanded medially, sometimes medially serrate, distally smooth or serrate; sometimes curved downward distally, apex bisected or whole; with complex DES1 (e.g., figs. 52E, 53G), bifurcate with two short serrate basal processes (below bifurcate portion), branches distally serrate; DES1 with elongate basal keel (DESk) present, membranous join at base to DES1, situated more adjacent and dorsal to DES2, with some anchoring membrane (figs. 49F, 50E).

Female: Almost identical to male, with very slight sexual dimorphism in slightly smaller eyes and hemelytral membrane length. On average, subequal size to males and sometimes slightly larger or smaller in body length, body length 2.84–4.08 mm. GENITALIA: Vestibulum with heavily sclerotized asymmetrical tubular processes, species specific (figs. 55A, 60A, 61B); ventral labiate plate sclerotized, mesial surface spiniferous with spiniferous clavate lateral lobes arising adjacent to gonapophyses 8 (figs. 60B, C, 61A); dorsal labiate plate with paired spiniferous mediolateral lobes on posterior lateral sides of genital chamber, lobes round, extending from margin (figs. 30A, 60B, 61B); sclerotized rings large, club shaped anteriorly, anterior surface spiniferous (figs. 55A, 61B); posterior wall heavily sclerotized; medial region of posterior wall more lightly sclerotized, margin straight or slightly convex, level with or slightly raised above anterior margin of interramal sclerites (figs. 55B, 60D, 74); interramal sclerites (IRS) deeply divided with V- or U-shaped junction, fused posteriorly below medial region of posterior wall forming posterior margin of posterior wall, posterior margin straight, broadly convex, broadly concave, or medially convex (figs. 30B, 60D, 61C, 74); IRS with two sets of paired interramal lobes, with species specific differences (fig. 74); lateral interramal lobes (laIRL) elongate and spiniferous with inner margin smooth and with spiniferous area or spiniferous lobe on inner margin at base (figs. 60D, 74); medial interramal lobes (mIRL) ventral to laIRL, with base embedded in socket and separated from laIRL (figs. 60D, 74), although proximate and with attaching membrane at base (e.g., fig. 74, anterolateral view); mIRL shorter and broader than laIRL (figs. 60D, 74); mIRL subquadrate or subrectangular, margins and sometimes also distal surface spiniferous (figs. 60D, 74).

ETYMOLOGY: Named for the distinctive red wing coloration on the hemelytral membrane veins of all species in this genus, from the Greek root erysi- (“red”) and the Latin word vena (“vein”). The gender is feminine.

REMARKS: Erysivena is best recognized externally by the red color of the hemelytral membrane veins and the red cuneal tips in the majority of species, yellow-green body, simple hairlike vestiture, small to medium size and elongate ovate body (fig. 14). The red forewing membrane veins are always present in Erysivena , but can be reduced within some species. Live specimens are bright green (fig. 4), with the first antennal segment also bright green and the remaining antennomeres lighter yellowish brown, and the lower third of the tibiae and tarsi are light orangebrown, with tarsal claw dark brown.

Erysivena and Callitricola are sister taxa and are similar in body size and shape; yellowgreen coloration; hairlike setae and lack of scalelike setae on dorsum; sclerotized tergal processes on the dorsal margin of the genital opening of the pygophore; dark bristles on the ventral surface of the pygophore; round genital opening of the pygophore; dark and strongly protruding phallotheca; simple L-shaped left paramere; PES positioned ventrad to the secondary gonopore; DES2 and DES1 left lateral to left dorsolateral of secondary gonopore; female interramal sclerite with two interramal lobes. In both genera, the shape of the left paramere is conserved and there is more interspecific variation in the right paramere.

Erysivena is less variable in size and shape than Callitricola . Erysivena can be distinguished from Callitricola by the red (rather than green) hemelytral coloration on the cuneus (cf. figs. 13, 14); presence of up to three elongate tergal processes on the pygophore, a moderately to greatly expanded left paramere with or without a hooked apex (cf. weakly expanded, strongly curved, and always hooked apex in Callitricola ); an externally projected and medially expanded right paramere mostly with a serrate apex (cf. club shaped, widest distally and often smooth at apex in Callitricola , with only the apex visible externally in repose); the phallotheca is tapered distally and not as strongly projected or as darkly sclerotized as in Callitricola ; the phallotheca is often not capped by the proctiger in Erysivena ; PES (cf. fig. 11) is not bifurcate and mostly narrow and smooth distally (cf. distally or medially bifurcate and distally serrate in Callitricola ); and DES2 is always distally serrate and medially serrate (cf. variable and never medially serrate in Callitricola ).

There are three main clades of Erysivena , with the three Western Australian species ( E. drepanomorpha , E. notodytika , E. kalbarri ) distinctly different from all other congeners, with complex PES and DES1 endosomal spicules, and PES and DES2 joined by a membrane basally on the right lateral side of the secondary gonopore.