Vieira flinti, Sosa, Francisco & Tauber, Catherine A., 2017

Sosa, Francisco & Tauber, Catherine A., 2017, The genus Vieira Navás (Neuroptera: Chrysopidae): a new species, a key to the species, and new geographic records, Zootaxa 4258 (1), pp. 43-59 : 44-58

Vieira flinti , new species

Type material. Holotype, Male, CAS; Fig. 6 . Venezuela, Yaracuy, Aroa , 1920. iii.14, J. & E. B. Williamson (deeply stained, labeled as a n. sp. by N. Penny). There are three paratypes (see below).

Etymology. The specific epithet was chosen to honor Dr. Oliver S. Flint, Curator Emeritus of Neuropteroid Orders, Entomology, Smithsonian Institution, United States National Museum of Natural History, Washington, D. C. Although a specialist on Trichoptera, Ollie is also a long-time student of the Neuroptera , the person who brought this species to our attention, and a treasured friend.

Diagnosis ( Figs 1–7 , 10 ): Externally, adults of V. flinti are slightly larger, but similar in gestalt to those of V. elegans and V. brooksi . Of the three species, this one appears to be the most cryptic; its largely clear wings with white veins on a relatively unreflective membrane, and diffuse patches of light brown coloration with brown venation, make the insect almost invisible against a variety of backgrounds. It differs from the other two species in that the large marks at the tips of its fore and hind wings are diffuse, and those on the hindwing do not reach the wing margin. Also, the basal forewing marks are large—surrounding several crossveins in the basal costal area, the two intramedian cells, the three median cells, the intracubital cells, and the anal veins. The basal forewing markings on the other two small species of Vieira are darker and less extensive.

Description. Body robust; dorsum largely black or dark brown, with anterior region yellowish or greenish mesally, dark brown to black laterally.

Head: Vertex green to yellowish, with light brown longitudinal streak along eyes, anterior margin of vertex; surface smooth; width 1.5–1.6 mm (including eyes); ratio head width to eye width 1.6: 1. Distance between eyes (frontal) 0.66–0.67 mm; distance between tentorial pits 0.45–0.51 mm; length midantennae to midway between tentorial pits 0.37 mm. Antenna length 8.8–9.0 mm; proximal segments short (segments 2, 3: length 1.0–1.2 times width), with three concentric rings of setae; middle and distal segments longer (segments 7–8: length 2.2–3 times width), four concentric rings of setae. Scape slightly elongate, black marked dorsally; pedicel black throughout; flagellum dark brown to black basally, cream to yellowish distally; antennal fossae marked with black.

Frons, clypeus yellowish with conspicuous brown shaded (male) or, brown (female) markings below scapes, light brown shading on frons and clypeus (male), or with elongate brown spot mesally (female); gena with dark brown stripe from base of eye to mid-clypeal region. Labrum pale, with distal margin indented mesally, bearing numerous long, thick, golden setae. Labial palpus, maxillary palpus black, both with last segment stout, tapering apically, covered by long, golden to amber setae.

Thorax: Prothorax wider than long (length 0.6–0.8 mm, width 1.2 mm), yellowish green mesally, with broad, brown, longitudinal bands covered with elongate, black, stout setae mesolaterally. Mesothorax broadly marked by brown spots with tufts of black setae (more dense in male than female specimens). Metathorax mostly brown, covered by long, black setae. Mesothorax, metathorax with black line across middle of pleural region. Legs cream colored, without markings, covered by numerous brown to amber setae; tarsal claws with broad, dilated base, deep narrow cleft.

Wings: Membrane transparent, not especially reflective, with light and dark brown markings and streaks, most notably at base of wings. Stigma semi-transparent, with light brown spot at each end (forewing), at basal end (hindwing).

Forewing with three prominent brown marks: first one in basal area—involving first ca. twelve costal crossveins, subcostal cell, basal part of radial sector, area around anal veins, cells b1, r1, m2-m3, im1, ic1-ic4; second one at tip of wing—surrounding inner, outer gradates, apical marginal forked veins; third one smaller, surrounding penultimate crossvein between Psm and Psc. Setae enclosed within black spots long, black; other setae gold colored.

Forewing 14.6 (male) to 16.6 (female) mm long, 5.41 (male) to 6.7 (female) mm wide (at widest point); ratio of length: maximum width 2.4–2.5: 1. Costal area broad; tallest costal cell (#9–11) 1.8–2.1 mm tall, 1.4–1.8 times width, height 0.17–0.15 times width of wing (midwing). First intramedian cell ovate, height (along median arculus) 0.2–0.5 mm, width 2.5–4.5 times height, 0.62–0.66 times width of third median cell. First radial crossvein black and distal to origin of radial sector (Rs); radial area (between radius and Rs) with single row of 11–12 closed cells; tallest cell (#6–8) 1.3–1.5 times taller than wide; two to three rows of gradate veins: three to six inner gradates in regular pattern (basal two white); inner gradate series closer to outer gradates than to Radial sector; six to seven outer gradates (all black), first five aligned in regular pattern, distal ones irregularly aligned; sometimes a third series of two gradate veins present. Height of fourth gradate cell 2.9–3.9 times width. Five b cells, four b’ cells (cells beneath Psm after second intramedian cell). Four intracubital cells [two or three closed], Cu2 bifurcating so that icu3 and icu4 lie next to each other. Subcosta, radial sector and apical veins forked apically.

Hindwing with most basal veins surrounded by dark brown, with large brown spot distally, encompassing distal area below radial sector, distal inner, outer gradate veins, and distal gradate cells. Veins pale except near darkened areas.

Hindwing 13.5 (male) to 14.7–15.0 (female) mm long, 4.41 (male) to 4.8–5.0 (female) mm wide; 12 radial cells (counted from origin of radius, not false origin), first one black bordered; two to three series of gradates; two to five inner gradates; five to six outer gradates; ~19 costal cells before pterostigma, first eight black bordered; four large b cells (no small “t” cell); three to four b’ cells beyond second intramedian cell; two intracubital cells (one closed).

Abdomen: Robust, uniformly expanded throughout. Segments of medium length and height, not particularly narrow or elongated; spiracles not especially large or small, simple, with unenlarged, bilobed atrium.

Female: Abdomen yellowish green; T1–T8 heavily marked by irregularly shaped black marks; ninth tergite, ectoproct (T9+ect) unmarked; callus cerci located dorsally on T9+ect, round, with ca. 31 trichobothria; S1, S2 with small black marks; S1–S6 densely covered with long, amber to golden setae; S7 almost entirely black, with heavy, black setae; tergites, sternites with elongate setae, slender, delicate on basal segments, becoming thicker, more robust distally; pleural regions with microthrichiae, short setae; pleuron of A6 with scattered, long, thin setae, pleuron of A7, A8 with elongate, thick, more dense setae; apical region of ectoproct with heavy, dense, up-curved setae; left, right T9+ect fused dorsally ( Brazil), separated by deep invagination ( Venezuela). Gonapophyses laterales rounded ventrally, with dorsal margins angular proximally, curved distally, entirely covered with long setae, especially heavy dorsally.

Praegenitale absent. Colleterial complex with elongate, delicate gland, apparently with scattered particles; reservoir large, spherical, with colliculate, membranous surface, extending apically only into ectoproct; transverse sclerite delicate, rectangular, with slender, elongate striations. Spermathecal complex ( Fig. 5 ) simple; spermatheca doughnut-shaped, with large, U-shaped invagination, dorsally with heavy, cylindrical, slightly bilobed velum connecting directly to bursa copulatrix via dorsal slit, velum merging distally into bursal duct, basal section of bursa copulatrix; spermathecal duct elongate (~3.0–3.5× width of spermatheca), arising from dorsal margin of spermatheca with distal one third, curvy, hair-like; bursal duct leathery basally, elongate, extending posteriorly to bursa copulatrix; bursa copulatrix saccular, membranous; bursal glands, flat, filamentous, with rough surface, extending posteriorly from lateral margins of bursa. Subgenitale broader than long (internal, proximal section extending transversely ca. 3/4th width of segment), distal section with cordiform knob extending from thick, short, folded neck; knob with U-shaped arms, small, rounded, lower lip extending ventrally.

Male: S7 and S8 densely covered by microthrichiae and microtholi; A8 spiracle simple; T9+ect divided dorsally by groove; apical margins rounded dorsally; basal margin acute, with elongate, dorsal apodeme along dorsal margin, forked distally around callus cerci; callus cerci large, located very low on T9+ect, with ~33 trichobothria; S8+9 triangular, tapering apically; margin of apex heavily sclerotized, contiguous with ventral apodeme along dorsal margin of S8+9; apex, with dense, long setae.

Gonarcal complex robust; gonarcal bridge straight; gonarcal apodemes broad, flared distally; gonocornua strongly fused with the gonarcus, tapering, then widening, then tapering again apically to acute tip; mediuncus, with a pair of dorsal rods, pair of lateral lobes, mesal beak. Hood quadrate, heavily sclerotized, attached at base of mediucus, extending above dorsum of mediuncus, with shallow teeth on distal margin. Hypandrium internum broad, V-shaped, with slender comes.

Material examined (in addition to holotype). BRASIL: BA [Bahia], Camacan, Res. [Reserva Particular do Patrimônio Natural] Serra Bonita , 15°23'S –39°33'W, 800m, iv.2010, 1 ♀. L. Santos col.; Vieira sp., Det. O. S. Flint 2013 (paratype). VENEZUELA: Aragua, El Limón , 450 m, 25. iv.1982, 1 ♀, F. Fernández Y col. ( MIZA); Idem, 8 . iv.1987, 1 ♂, L. J. Joly col (MJMO) (paratypes).

Variation. The most notable variation we found among the specimens is in the dorsal fusion of the left and right sides of the ninth abdominal tergite+ectoproct (T9+ect). In the females ( Brazil, Venezuela) the two sides are fully and clearly fused; whereas in the Venezuelan males there is a distinct groove between the two sides.

Known geographic distribution. Venezuela (States of Aragua and Yaracuy) , Brazil (Bahia State).

Generic characters. With the description of the new species, it is worthwhile to review the validity of the diagnostic generic features that were proposed based on previously known species (see Tauber et al. 2006, Tauber 2006, 2007). Below, we discuss V. flinti in relation to nine morphological character states that have been used to distinguish Vieira from other genera.

Basal radial crossvein (compare Figs 2 A, 2C, 10A with Figs 8 A, 9A, 11A). Vieira elegans originally was placed in the genus Berchmansus Navás because it expresses a characteristic that was considered distinctive for that genusi.e., the basal radial crossvein leaves the Radial vein (R) before the origin of the Radial sector (Rs). Such is the condition in V. brooksi as well. However, in both V. flinti and V. lescehnaulti , the first radial crossvein originates from the Rs after its separation from the R. Thus, it appears that this feature is not a diagnostic character for the genus.

Splayed costal crossveins on forewing (compare Figs 2 A, 2C, 10A with Figs 8 A, 9A, 11A). A conspicuous and defining feature of the genus is the forewing with the costal area enlarged and with a pronounced brown patch surrounding a series of splayed costal crossveins. This character state continues to hold as a diagnostic feature for Vieira , and it expresses some valuable variation for distinguishing among Vieira species. In V. elegans and V. brooksi , the enlargement of the costal area is moderate; in V. leschenaulti and V. flinti it is more abrupt basally and more pronounced across the margin of the wing. The pattern and number of splayed crossveins also varies: V. elegans has a small, slightly splayed patch of ~five unbranched crossveins; V. brooksi has a similarly small, but slightly more splayed patch of five to six crossveins, one of which is usually branched near its base; V. flinti has a large, well splayed patch of about ten to 13 crossveins—all unbranched in the Venezuelan specimens and with one branched crossvein (#7, #8) in the Brazilian specimen; V. leschenaulti has fewer (~5) splayed crossveins—all unbranched, but they are relatively widely splayed.

Bifurcation of forewing Cubital (Cu) vein (compare Figs 2 A, 2C, 10A with Figs 8 A, 9A, 11A). In Vieira there is a distal bifurcation of the forewing Cu, and among the four species the placement of the bifurcation appears to exhibit some clinal variation: In two species, V. elegans and V. brooksi , the bifurcation occurs after the Cu has left the Psc (sometimes very near the furcation of Cu from Psc); in these species both icu3 and icu4 are present and open on the exterior margin of the wing. In V. flinti , the bifurcation occurs at or very near the point where the Cu leaves the Psc; in this species, the icu3 is closed, and icu4 is present and open on the wing margin. In V. leschnaulti the bifurcation occurs still earlier—before the Cu leaves the Psc; icu3 and icu4 are both present and open, side by side on the wing margin.

Slanted distal vein of the third medial cell (m3) (compare Figs 2 A, 2C, 10A with Figs 8 A, 9A, 11A). In V. leschenaulti , the distal vein of m3 is strongly slanted distally and is followed by six closed cells beneath the Psm. In V. flinti , the distal vein of m3 is slanted somewhat, and in V. brooksi and V. elegans it is vertical; in all three of the above species, the m3 is followed by four closed cells beneath the Psm.

Wing markings (compare Figs 2 A, 2B, 10A, 10B with Figs 8 A, 8B, 9A, 9B, 11A, 11B). In all four species, the following membranous areas of the wings are characteristically and prominently marked with brown to dark brown: around the base of the fore and hind wings, around the splayed costal crossveins (forewing), around a series of basal or midbasal crossveins (hindwing); around the distal inner and/or outer gradates (fore and hind wings); around one or more Psm or Psc crossveins (fore and hind wings), around the intracubital cells (fore and hind wings). Although these areas are consistently marked in the four Vieira species, the size and depth of the markings vary among them. For example, the markings around the gradate veins are relatively small and more widely separated on V. flinti , compared with those on the other Vieira species.

Head and body markings (compare Figs 10 C, 10D with Figs 8 C, 8D, 9C, 9D, 11C, 11D). All four Vieira species express the following distinctive markings: elongate, longitudinal markings on sides of vertex; scapes marked with dark brown or black at least dorsally; pedicel and one or more basal flagellomeres dark; distal flagellomeres yellow.

Setation (compare Figs 1 A, 10C with Figs 8 C, 9C, 11C). All four Vieira species have patches of long, heavy, black setae on the mesothorax and metathorax. In addition, the terminal segments of the labial palpus and the terminal segments (tergites and sternites) of the abdomen have dense, relatively heavy setae. Again, V. flinti shares these features with its congeners, but in V. flinti the labial setae are golden and quite striking ( Figs 1 C, 1D, 10D).

Gonarcal hood. Male specimens of three Vieira species have been diagnosed as having a gonarcal complex bearing a “hood”-like structure either above or below the mediuncus; the structure is sclerotized and prominent in V. elegans , V. brooksi ( Tauber et al. 2006, Tauber 2006), as well as V. flinti ( Figs 6 C, 6D, 6E); it was membranous to lightly sclerotized in the V. leschenaulti specimens studied to date; perhaps most were teneral ( Tauber 2007). Thus, this character appears to be diagnostic of the genus.

Spermathecal shape and size. Females of Vieira species were proposed to have a round spermatheca bearing an elongate, tubular, and at least somewhat bilobed velum that opens to the bursa copulatrix dorsally via an elongate dorsal slit, and also distally via the bursal duct. The spermatheca has an elongate, slender spermathecal duct ( Brooks & Barnard 1990; Tauber 2007). The reproductive tract of V. flinti females also expresses all of these features.

From the above, it is clear that the four Vieira species share a number of unifying characteristics, but they also express a wide range of interspecific variation. Most notably, the four species vary markedly in body size. Indeed, the variation in many of the above features may be correlated with body size. Such a broad range of variation in size and correlated features may not be unexpected if, as is the case for all known belonopterygines, the larvae of Vieira species feed on ant brood and have relatively narrow ranges of acceptable prey.

Key to species of Vieira (See Figs 8–11 ; wing veins are identified on Figs 2 C, 2D)

1. Body large, forewing 22–32 mm long; inner series of gradate veins aligned end to end in a row extending away from the Pseudomedius and the outer gradate series and arching toward the Radial Sector ( Fig. 11 A).............. V. leschenaulti

1’ Body smaller, forewing length <17 mm; inner and outer series of gradates aligned in parallel, step-like rows........... 2

2. Dark markings at base of forewing large, including ~ ten basal costal crossveins, the intramedian and third median cells, and the three cubital cells; large marks at tips of fore and hind wings diffuse, not reaching wing margin ( Fig. 10 A)..... V. flinti

2’ Dark marking at base of forewing small, including only the bases of the major veins (M, M1, M2, Cu, A1, A2) and a few basal crossveins (m-cu1, cu-a); large marks on tips of fore and hind wings dark, those on hindwing reaching margin of wing.....3

3. Forewing relatively small (11.4–11.7 mm long); large midcostal marking including six crossveins, ususally one of which is bifurcated; large mark at tip of forewing not reaching wing margin ( Fig. 8 A).............................. V. brooksi

3’. Forewing relatively large (13.6–15.2 mm long); large midcostal marking including four to six crossveins, all of which are simple; large mark at tip of forewing reaching wing margin ( Fig. 9 A)....................................... V. elegans

Figures

FIGURE 1. Ƒieira flinti, sp. nov., Paratype (female, Brazil). External features. A. Head, prothorax (dorsal); B. Head (frontal); C. Head, prothorax (lateral); D. Head (lateral); E. Head, thorax, legs (lateral); F. Tarsal claw; G. Abdomen (ventrolateral). S 7, seventh sternite.

FIGURE 2. Ƒieira flinti, sp. nov., Paratype (female, Brazil). Wings. A. Forewing; B. Hindwing; C. Base of forewing, with features labeled; D. Tip of forewing, with features labeled. Veins: C, costa; Cu, cubitus; M, media; Psc, pseudocubitus; Psm, pseudomedia; R, radius; Rs, radial sector; Sc, subcosta. Crossveins: c-sc, costa-subcosta crossvein; i. g., m. g., o. g., inner, middle and outer gradate veins; m-cu 3, third media-cubitus crossvein; r-m 1, first radial crossvein. Cells: b 3, b 5, third and fifth upper Banksian cells; b’ 1, b’ 4, first and fourth lower Banksian cells; icu 1, icu 2, icu 3, icu 4, first to fourth intracubital cells; im 1, im 2, first and second intramedian cells; m 1, m 2, m 3, first to third median cells. Vein splits: cuf 1, cuf 2, first and second furcations of cubitus; mf, first furcation of media. st, stigma.

FIGURE 3. Ƒieira flinti, sp. nov., Paratype (female, Brazil). Abdomen. A. Segments A 4 - terminus (lateral); B. Terminal segments (lateral); C. Genitalia; D. Tergite 8, tergite 9 + ectoproct (dorsal). b. c. bursa copulatrix; col. res., colleterial reservoir; g. l., gonapophysis lateralis; sg, subgenitale; sp, spermatheca; sp. d., spermathecal duct; S 7, seventh sternite; tr. pl., transverse plate; T 8, eighth tergite; T 9 + e, fused ninth tergite and ectoproct.

FIGURE 4. Ƒieira flinti, sp. nov., Paratype (female, Venezuela). Abdomen. A. Segments A 2 - terminus (lateral); B. Segments A 2 - terminus (dorsal); C. Callus cerci; D. Spiracle; E. Terminal segments and genitalia (extruded, lateral). g. l., gonapophysis lateralis, sg, subgenitale; S 7, seventh sternite; T 7, T 8, seventh and eighth tergites; T 9 + e, fused ninth tergite and ectoproct.

FIGURE 5. Ƒieira flinti, sp. nov., Paratype (female, Venezuela). Genitalia. A, B, E. Spermathecal complex (lateral); C, D. Subgenitale, ventral. b. c., bursa copulatrix; b. d., bursal duct; inv, invagination; sp, spermatheca; sp. d., spermathecal duct; vel, velum.

FIGURE 6. Ƒieira flinti, sp. n., Holotype (male, Venezuela). A. Terminus (lateral); B. Tergite 9 + ectoproct (lateral); C. Gonarcus (dorsal); D. Gonarcus (posterodorsal); E. Gonarcus (lateral). c, comes; c. c., callus cerci; d. a., dorsal apodeme; gc, gonocornu; g. a., gonarcal apodeme; g. br. gonarcal bridge; hd, gonarcal hood; h. i., hypandrium internum; mu, mediuncus; S 8 + 9, fused sternites S 8 + S 9; T 9 + e, fused tergite T 9 + ectoproct; v. a., ventral apodeme.

FIGURE 7. Ƒieira flinti, sp. nov., Paratype (teneral male, Venezuela). Abdomen and genitalia. A. Terminal segments (ventral); B. Terminal segments (dorsal); C. Terminal sternites (lateral); D. Terminal sternites (ventral); E. Gonarcus (lateral); F. Gonarcus (dorsal). c. c., callus cerci; gc, gonocornu; g. a., gonarcal apodeme; S 7, seventh sternite; S 8 + 9, fused eighth and ninth sternites; T 8, eighth tergite; T 9 + e, fused ninth tergite and ectoproct.

FIGURE 8. Ƒieira brooksi Tauber. (Male, Venezuela). External features. A. Forewing; B. Hindwing; C. Head, thorax (dorsal); D. Head (frontal). Scale applies to both A and B.

FIGURE 9. Ƒieira elegans (Guérin Méneville). (Male, Brazil). External features. A. Forewing; B. Hindwing; C. Head, thorax (dorsal); D. Head (frontal). Scale applies to both A and B.

FIGURE 10. Ƒieira flinti, sp. nov. (Female, Venezuela). External features. A. Forewing; B. Hindwing; C. Head, thorax (dorsal); D. Head (frontal). Scale applies to both A and B.

FIGURE 11. Ƒieira leschenaulti (Navás). External features. A. Forewing (female, Bolivia); B. Hindwing (female, Bolivia); C. Head, prothorax, mesothorax (female, Costa Rica; dorsal); D. Head (male, Costa Rica; frontal). Scale applies to both A and B.