Leucochrysa (Leucochrysa) nigrilabris (Banks, 1915)

Tauber, Catherine A., Sosa, Francisco, Albuquerque, Gilberto S. & Tauber, Maurice J., 2013, Adults and larvae of two Leucochrysa (Leucochrysa) species (Neuroptera: Chrysopidae): descriptions, biological notes, and relationships, Zootaxa 3750 (2), pp. 101-129 : 103-117

Leucochrysa (Leucochrysa) nigrilabris (Banks, 1915)

Figs 1–12 , 19

Allochrysa nigrilabris Banks , [1915] (Banks 1914–1915: 623) original description: [ Colombia] “St. Antonio, Colombia, 1,800 m., December (Fassl)”.

Leucochrysa nigrilabris (Banks) . Banks (1944: 32) first reference to combination; Penny (1977: 23) species list. Leucochrysa (Leucochrysa) nigrilabris (Banks) . Brooks & Barnard (1990: 276) subgeneric designation, species list; Tauber (2004: 1140) possible synonymy with Leucochrysa (Leucochrysa) insularis (Walker) ; Oswald (2007) catalog listing.

Type specimen. One type, from San Antonio, Colombia, MCZ (male, examined). We assume that the locality “St. Antonio” refers to the city of San Antonio in the Department of Tolima. Banks (1914–1915) did not state how many specimens he used to write his description; it is likely that he had only one. There is only one type in the MCZ and none in the AMNH, USNM, or the Academy of Natural Sciences of Philadelphia, where Banks occasionally deposited types. Thus, we consider the specimen in the MCZ to be the holotype, by monotypy. Its labels read: (1) “St Antonio / Colombia S. Am” [Banks’ hand]; (2) “ 1800 m / Dec” [Banks’ hand]; (3) “Collection / N. Banks”; (4) “ Type ” [red, Banks’ hand]; (5) “ Type / 12004”; (6) “ Allochrysa / nigrilabris / type / Bks” [white, red border, Banks’ hand]; (7) “Genitalia / prep P. Adams / 1982”.

The specimen is in fairly good condition; the body (thorax and head) are somewhat flattened, the forewings and one hindwing are missing, and the terminalia are in a vial with glycerine. We took images that are now placed in the MCZ Type Database (http://insects.oeb.harvard.edu/MCZ/index.htm).

Known geographical distribution. Colombia: Department of Tolima. Venezuela: States of Aragua, Mérida, Portuguesa, Yaracuy. Brazil: State of Rio de Janeiro.

Diagnosis (adult). Externally, L. (L.) nigrilabris adults are distinguished from many Leucochrysa (Leucochrysa) species by a dark brown to black labrum, white frons, a single, small spot on the forewing (brown suffusion on the membrane around the distal Psm-Pcu crossvein), and all veins mostly green except inner and outer gradates dark brown to black. However, these features are shared with other Leucochrysa (Leucochrysa) species, including some L. (L.) “ varia -like species” and L. (L.) insularis . Thus, a reliable identification requires examination of the genitalia. The males of L. (L.) nigrilabris are distinguished by: (i) dense microtholi on sternites 3–8; (ii) small genital structures [gonarcal span of 0.40–0.65 mm; cf., 0.67–0.84 mm in L. (L.) varia ]; (ii) a narrow, elongate mediuncus that is well separated from the gonarcal bridge, and terminates in a long, slender hook and a pair of large, inflated flanges; and (iii) terminus of S8+9 with an attached pair of flat, sclerotized, plate-like projections with toothed mesal margins. Female L. (L.) nigrilabris have a doughnut-shaped spermatheca with a broad, bent velum, a deep, relatively narrow invagination, a relatively long, robust spermathecal duct, and apparently no bursal duct; the subgenitale has a well-sclerotized ventral process that extends perpendicularly from the distal margin of the base.

Redescription (adult). Head ( Fig. 1 ): 1.7–1.9 mm wide (including eyes). Frons, clypeus white to creamcolored, frons without red or pink suffusion, clypeus with or without tinge of pink; genae reddish brown; labrum black throughout; maxillary, labial palpi yellowish to cream-colored. Vertex with central area raised, green mesally, golden brown to reddish brown laterally; lateral, frontal areas cream-colored, with prominent, dark red to dark brown, scalloped to V-shaped mark along anterior margin, extending anteriorly to or between mesal margins of scapes, extending laterally to eyes; post-ocular area cream-colored, without spots. Antenna: scape cream-colored ventrally, with reddish brown mark of various sizes on distal, dorsolateral surface; pedicel cream-colored basally, with brown band distally; flagellum cream-colored, with brown bristles; mesal-ventral surface of basal one to seven flagellomeres lightly tinged with brown (variable); dorsal antennal fossae cream-colored.

Thorax ( Figs 1 D–H): Cervical sclerite with large, red mark laterally. Pronotum 0.92–0.98 mm long, 1.41–1.44 mm wide, light green, with elongate, dark red stripe on anterolateral margin, diffuse, light brown, sublateral spots anteriorly, posteriorly. Mesonotum, metanotum variable, with two distinct morphs [ Figs 1 D, E (brown), 1F (white)]. Brown morph: mesonotum, metanotum dark brown throughout, except anterior margin of prescutum cream-colored, with pair of sublateral, dark brown spots; mesoscutellum slightly lighter brown than remainder of notum; metascutellum black throughout. White morph: mesonotum mottled black to brownish black, anterior margin of prescutum cream-colored, with pair of sublateral, dark brown spots, posterior margin with cream-colored areas, mesoscutellum entirely white; metanotum largely dark brown to black, metascutum with large, white posteromesal patch; metascutellum black throughout.

Wings ( Fig. 2 ) Forewing 18.8–19.9 mm long, 6.5–6.9 mm wide (at widest point); ratio of length: maximum width = 2.6–2.7:1. Costal area moderately broad; tallest costal cell (#9–10) 1.6–1.7 mm tall, 3.0–3.1 times width, 0.2–0.3 times width of wing (midwing). First intramedian cell quadrangular, width (anterior margin) 1.4–1.7 times width (anterior margin) of third median cell, 0.9–1.1 times length of posterior margin of m3; length of basal vein (= ma, median arculus) 2.1–6.2 times greater than length of distal vein. First radial crossvein distal to origin of radial sector (Rs); radial area (between Radius and Rs) with single row of 17–18 closed cells; tallest cell (#8) 2.3–2.4 times taller than wide. No crassate veins; 5–6 b cells (= cells beneath Rs, not including an inner gradate vein). Two series of gradate veins; 10–12 inner gradates, 8–10 outer gradates. Height of fourth gradate cell 4.3–5.3 times width. Eight to nine b' cells (cells beneath pseudomedia after second intramedian cell). Three intracubital cells (two closed). Membrane clear except stigma opaque, with dark brown mark basally, distal two crossveins of last b’ cell (distal Psm-Pcu crossveins) surrounded by dark brown suffusion, and (some specimens) small amount of brown suffusion basally. Veins green, except distal two crossveins of last b’ cell, inner and outer gradates dark brown to black, sometimes anterior tips of basal R-Rs crossveins, base of Rs, basal Rs-m, basal medial crossveins, tips of posterobasal marginal veins lightly darkened.

Hindwing 16.7–17.7 mm long, 5.5–5.8 mm wide. Two series of gradate veins; 9–10 inner, 8–9 outer; 15–16 radial cells (counted from origin of Radius, not false origin). Five to six b cells (including small b1 cell); seven to nine b' cells beyond second intramedian cell; two intracubital cells (one closed). Membrane clear; stigma with pronounced brown mark basally. Veins light green.

Abdomen ( Figs 3–8 ): Tergites with mostly short, slender setae throughout, sternites with longer, slender setae; microsetae dense; pleural region with setae small, very sparse, microsetae very small. Tergites narrow, roughly rectangular, with rounded or irregular margins. Spiracles oval externally; atria not enlarged. Sternites S2–S5 longer than wide (lateral view). Coloration: mostly green, with diffuse yellow dorsal stripe; tergites T5, T6, base of T7 each with large dark brown to black spot, bordered with yellow and patches of red; callus cerci white; setae, trichobothria golden.

Male. S6, S7 height ca 1.1–1.2 times length (lateral view). Microtholi dense on S3–S8, absent from S1–S2, S9. Callus cerci oval (ca 1.3–1.5x taller than wide), greatest diameter 0.22–0.26 mm, with 26–35 trichobothria of variable length. T9+ectoproct soft, lightly sclerotized, rounded posterodorsally, truncate distally, broadly fused mesally, midline without distal cleft (mature specimen; cleft present in teneral specimen); ventral section tapering proximally, extending beyond midline of S9; dorsal apodeme lightly to moderately sclerotized, straight, extending ca half distance along dorsal margin of ectoproct; small, lightly sclerotized apodeme around proximal margin of callus cerci. S8+9 fused, without suture, with clear intersegmental demarcation throughout; dorsal margin with lightly sclerotized, straight apodeme; S8 1.7x taller than long, ca one-half (0.47–0.51x) length of S8+9; S8+9 (lateral view) with proximal margin relatively straight, dorsal surface straight, sloping ventrally to distal margin; terminus rounded; tip with pair of flat, sclerotized, plate-like projections with mesal edge toothed, lateral edge rounded. Gonarcus arcuate; bridge straight, narrow mesally (0.19 mm), curved abruptly at exterior margin of gonocornua; gonarcal bridge, gonocornua covered with robust membrane bearing dense gonocristae; gonocornua short (length, 0.07 mm), extending forward from edge of gonarcal bridge, stout basally, tapering to rounded apex; distance between inner bases of gonocornua 0.07 mm basally, distance between tips 0.12 mm; gonarcal lateral apodeme long, relatively narrow (0.60 mm long, 0.20 mm wide), not extending above bridge, curved distally (lateral view); distance between apodemes distally 0.72 mm. Gonarcus, between lateral apodeme and gonocornu, with moderately sclerotized, elongate posteroventral projection holding membrane that extends beneath mediuncus. Mediuncus well separated from gonarcus, with basal membranous connection stout; basal section of mediuncus consisting of heavily sclerotized platform with distal projections laterally, distal section consisting of well sclerotized, ventrally-projecting, smooth plate terminating in sharp, curved, elongate, mesal beak, pair of large lateral lobes; heavy membrane from gonarcus extending beneath mediuncus, recurving above beak to form gonosaccus with pair of elongate, lateral fields of robust gonosetae on bulbous chalazae; fields of gonosetae on surface of membrane facing mediuncus; each field with ca 15–20 gonosetae. Hypandrium internum broadly Vshaped, with comes not visible (mature specimen); arm 0.20–0.22 mm long, distal span between arms 0.19 mm.

Female. S6 height ca 0.67–0.92 times length, S7 height ca 0.55–0.65 times length. Callus cerci round, diameter ca 0.17–0.19 mm, with 29–34 trichobothria. T8 roughly elliptical (lateral view) with rounded corners, similar in depth to T6. T9+ectoproct elongate, vertically erect; ventral margin slightly rounded, extending below level of gonapophyses laterales. S7 with dorsal margin straight, distal end tapering abruptly; terminus truncate, unmodified, with terminal (posteroventral) setae slightly more numerous, longer than in other areas. Gonapophysis lateralis rounded throughout, ~0.48–0.52 height of T9+ectoproct; inner membranous surface slightly expandable, with vertical patch of small, delicate setae arising from slightly swollen membrane. Colleterial complex consisting of membranous gland, connected to colleterial reservoir via broad duct, and large accessory gland, both opening to exterior via narrow common duct above transverse sclerite; colleterial gland elongate, delicate; colleterial reservoir shorter, robust; accessory gland probably bulbous (torn in our specimens), elongate, granular, with multiple forks distally; transverse sclerite broad, flat, bent distally, with relatively dense, elongate, hair-like setae. Spermatheca simple, doughnut-shaped, with broad, flat, L-shaped velum, maximum diameter 0.26–0.31 mm, total length (spermatheca + velum) ca 0.35 mm; spermathecal invagination deep, narrow, with rounded terminus; velum with elongate slit opening to bursa copulatrix; tip of velum bending slightly into bursa copulatrix; bursal duct very small or absent; spermathecal duct relatively long, ca 1.13–1.51 mm, originating from side of spermatheca distal to slit in velum, basal half well sclerotized, tapering throughout, with lateral bend, then U-shaped loop; distal ca half narrow, brushy throughout, with curve, followed by distinct swelling mesally, with densely hirsute, swollen terminus. Bursa copulatrix saccular, with heavily textured surface distally (below subgenitale), becoming smoother anteriorly, with pair of elongate, tubular bursal glands attached to distal end, connections appearing unremarkable. Subgenitale with sclerotized surface smooth (unfolded), attached basally to heavy, transversely folded basal membrane (base of subgenitale then top of bursa copulatrix), with two projections, bilobed one on dorsodistal margin, unilobed one on anteroventral margin base withdrawn above tip of S7 [with pressure, the withdrawn lobe can be extruded].

Intraspecific variation (adult). Leucochrysa (L.) nigrilabris expresses considerable variation in head and mesothoracic coloration, as well as wing size, shape and degree of brown suffusion surrounding various veins. The most striking aspect of the variation is the polymorphism in mesothoracic coloration ( Figs 1 D–F). Among the specimens we examined, eight (4♀, 4♂) had brown mesoscutella, and five (2♀, 3♂) had white; both color forms occurred among the Venezuelan specimens.

The male genitalia show considerable developmental variation (Compare Fig. 5 with Figs 3 , 4 ). In the teneral male, the gonarcus is weakly sclerotized and narrow; the mediuncal bridge appears wide (in lateral view) and is tightly encased by the gonocristae-bearing membrane; the mediuncus is very narrow; and the hypandrium internum is unexpanded [length of arms, 0.20 mm—as in mature specimens; distal span between arms, 0.06 mm— ca 3x narrower than mature specimens]. Females, too, show considerable variation with age and maturation. For example, in the teneral female, the subgenitale is small, weakly sclerotized, and withdrawn, and the spermatheca is shifted anteriorly from the distal wall; whereas in mature specimens, the subgenitale is large, robust, heavily sclerotized and expanded, so that the spermatheca appears surrounded by the distal wall ( Figs 6 , 7 B).

Description (First Instar, Semaphoront A). Body ( Fig. 10 ). 2.8–3.0 mm long. All setae smooth.

Head ( Figs 10 A–C). 0.55–0.56 mm wide, 0.38–0.39 mm long; mandibles 0.59–0.61 mm long; ratio, mandible length: head width, ~1.09:1. Epicranial marking brown, paired, extending along entire basal margin of cranium, laterally extending to eye, mesally becoming confluent with base of postfrontal marking. Postfrontal marking slightly darker brown than epicranial marking, elongate, extending to base of antenna. Frontal marking V-shaped, arms extending forward through dark tentorial pits becoming confluent with intermandibular marking. Intermandibular marking broken mesally, extending laterally from tentorial pits to base of mandibles, anteriorly to tip of clypeus. Gena with single, elongate, brown stripe extending from base of eye to posterior edge of cranium. Labial palpus cream-colored basally, light brown distally. Mandibles amber to brown. Antenna light brown to cream-colored. Venter cream-colored to white mesally, light brown laterally, with brown transverse stripe distally; cardo, stipes brown.

All cephalic setae straight, with pointed terminus. Primary setae S1–S12 present; S1, S11 long; S2, S3, S5 short. Labial palpus: basal segment with two setae—one lateral, one ventral; midsegment with two distal setaeone lateral, one mesal. Mandible with one basolateral seta; maxillae with basal seta. Palpiger with two basal setae. Scape without setae; pedicel with one small distal seta; flagellar seta slightly longer than one-half length of segment.

Thorax ( Figs 10 D, G, H). Notum with diffuse, reddish brown, submesal markings; Setae (LS) on lateral tubercles (LTs) dark brown; dorsal setae light brown. Legs mostly light brown to brown; trochanters, tarsi white to cream-colored; claws, empodia brown. LS gently curved throughout, with tips pointed, fine; all other setae straight to gently curved, with terminus pointed.

T1: LT with two long LS, without microseta between. First primary sclerite (Sc1) light brown, large, with associated seta (S1Sc1) long, arising from small chalaza; S2Sc1 short, anterior to S1Sc1. S1 long, arising from small chalaza; S2 absent; S3, S4, S5 short.

T2: Spiracles prominent, tubercle-shaped, apex sclerotized, dome-shaped, amber-colored. LT with three long LS. Sc1, Sc2 small; associated setae (S1Sc1, S2Sc1 and S1Sc2) very small; Sc3 intermediate-sized, light brown, with associated seta (S1Sc3) small. Posterior subsegment with pair of short setae mesal to Sc3, row of four setae posterior to Sc3 (mesal pair short, lateral pair long, arising from robust chalazae).

T3: LT with three long LS. Sc1 small, with S1Sc1 very small; Sc2 light brown, with S1Sc2 small. Anterior region with pair of short setae; posterior region with four long setae arising from robust, fleshy chalazae.

Abdomen ( Figs 10 E–H). White to cream-colored; dorsum of A1–A3 with diffuse, submesal, reddish brown band; dorsum of A4, A6–A8 each with pair of reddish brown submesal spots. LTs each with two long LS (A2–A7), without microseta between; LS straight, with pointed tip, dark brown; submedian setae (SMS), setae (LDS) on laterodorsal tubercles (A1–A5) hooked, light brown. A1, A6–A8: Spiracle without associated seta (SSp); A2–A5: each spiracle with small to minute SSp. A1–A5: Laterodorsal tubercles (LDTs) each with two long, hooked LDS, one microseta between LDS. A1: Single row of four SMS between LDTs. A2–A4: Anterior row of four SMS, posterior row of two SMS, both between LDTs. A5: Anterior row of two to three SMS, posterior row of two SMS, both between LDTs. A6: Two short, straight setae anteriorly, two intermediate-length, hooked setae posteriorly; LDT with one long, hooked LDS, one short, straight LDS. A7: LDT with one intermediate-length, straight LDS, two very short LDS; dorsum with pair of minute setae anteriorly. A8: LT with one short, straight LS, two to three small setae, base of LT with spiracular tubercle projecting from anterior surface; spiracular base and sclerotized ring elongate, projecting posteriorly; dorsum with three to four short, straight setae between LTs. A9: Dorsum with three rows of short, straight setae (anterior two minute, middle four short, posterior four slightly longer). A10: Apparently without setae; terminus light brown, with small eversible pouch.

Description (Second & Third Instars, Semaphoront B). Body ( Figs 9 , 11–12 ). L2: 4.7–4.9 mm; L3: 8.6–9.0 mm long. Dorsum cream-colored, with dark brown markings.

Head ( Figs 11 A, B, 12A, B, G, H). L2: 0.81–0.83 mm wide, 0.53–0.58 mm long; mandibles 0.93–0.96 mm long; ratio mandible length: head width 1.14–1.17:1. L3: 1.3–1.4 mm wide, 0.86–0.91 mm long, mandibles 1.5– 1.7 mm long; ratio mandible length: head width, 1.19–1.23:1. Epicranial marking paired, brown to dark brown, covering entire basal portion of cranium, laterally becoming lighter brown, extending to eye, mesally becoming confluent with base of postfrontal marking. Postfrontal marking slightly darker brown than epicranial marking, elongate, extending to base of antenna. Frontal marking V-shaped, arms extending forward through dark tentorial pits, then curving to base of mandibles becoming confluent with intermandibular marking. Intermandibular marking brown, divided into three sections—two lateral sections at mesal base of mandibles, mesal section between, but separated from two lateral sections; clypeolabral region partially covered by intermandibular marking. Gena with single, elongate, brown stripe extending from base of eye to posterior edge of cranium; region around eyes cream-colored; stemmata, except posteroventral margin, surrounded by brown. Labial palpus: basal segment light brown; mesal segment light brown basally, darker distally, with well defined annulations; terminal segment dark brown. Mandibles brown basally to dark brown at tip. Antenna light brown to brown. Ventrolateral cranial surface light brown. Cardo with light brown mark basally, dark brown mark distally, narrow; stipes light brown to tan, narrow, elongate. Mentum white to cream-colored, with tinge of light brown anteriorly; palpiger light brown. Cervix cream-colored, with prominent lateral patches of brown.

Anterior margin of head with slightly rough, obtuse lateral edges; posterior section of cranium spiculate. All cephalic setae smooth, straight, with acute tip; S1–S12 present; S1, S11 long; S2, S3, S5 short. Labial palpus: basal segment with three setae, midsegment with distal annulation bearing four setae, remaining annulations with ca six (L2) or seven (L3). Mandible with one basolateral seta; maxillae with basal seta. Palpiger with two basal setae. Scape with one distolateral seta; pedicel with one small distal seta; flagellar seta ca one-half length of segment.

Thorax ( Figs 9 , 11 C, F, 12C, H, I). LTs white to cream-colored, with diffuse to very pale, reddish brown markings at dorsal base submesally; LS mostly dark brown; large sclerites Sc1 (prothorax), Sc3 (mesothorax), Sc2 (metathorax) light amber, shiny. Venter white to cream-colored, with small, diffuse, rosy, submesal patches on each segment. Spiracles amber to light amber; setae mostly brown or light brown. Legs: coxa with dark brown marking on dorsal connection to base, pair of brown marks, one anterior, one posterior; trochanter with pair of dark brown marks, one anterior, one posterior; femur brown dorsobasally, light brown distally; tibia brown throughout; tarsus light brown; claws curved, large, dark brown; empodia dark brown.

LTs smooth; LS smooth, except very fine tips sometimes slightly salebrose, seldom sparsely and finely barbed, slender, gently curved throughout, tips tapering to very fine terminus; tips of distal LS mostly pointed, tips of lateral LS mostly with small hook. All dorsal, ventral setae smooth.

T1: All dorsal, ventral setae straight; Sc1 large, extending laterally to dorsal surface of LT, with S1Sc1 long, slender, S2Sc1 very short, slender. Sc2 transparent, oval-shaped, ca one-third length of Sc1. LT (L2) with 19–21 LS (ca 10 apical, ca 9–11 lateral); LT (L3) with ca 27–30 LS (ca 10 apical, ca 17–20 lateral). Primary setae S1 medium-length, S2, S3, S4, S5 short, slender; no secondary setae.

T2: Anterior subsegment without secondary setae between spiracles; anterior sclerite (Sc1) with three associated setae (S1Sc1, S2Sc1, S3Sc1); spiracles prominent, with opening sclerotized, dome-shaped, ambercolored ring on small tubercle; walls of chamber parallel, not bulbous; SSp absent. Posterior subsegment with anterior sclerite (Sc2) bearing two very small associated setae (S1Sc2, S2Sc2); Sc3 with small associated seta (S1Sc3); anterior row with two intermediate-length, slender, straight setae; posterior row (L2) with ca 5–6 setae (3–4 long, robust, 1–2 slender, hooked or straight), arising from large, plump chalazae; posterior row (L3) with ca 11–14 setae (6–8 long, robust, 5–7 slender, hooked or straight), arising from large, plump chalazae. LT (L2) bearing ca 17–19 LS (ca 8–9 apical, ca 9–10 lateral, mostly straight); LT (L3) bearing ca 26 LS (ca 10 apical, ca 15–17 lateral).

T3: Sclerites (Sc1, Sc2) each with one associated setae (S1Sc1, S1Sc2); sometimes small seta on anterior fold, lateral to Sc1. Anterior row with one to two short (L2) or two to four intermediate-length (L3), slender, straight setae; posterior row (L2) with ca 12 long, robust, straight or hooked setae arising from large, plump chalazae; posterior row (L3) with ca 15–16 setae (ca ~12–14 long, robust, 2–4 intermediate-length, slender, straight or hooked), arising from large, plump chalazae. LT (L2) bearing ca 19–20 LS (ca 9 apical, 10–11 lateral, mostly straight). LT (L3) bearing ca 21–25 LS (ca 9 apical, 12–16 lateral).

Abdomen ( Figs 11 D–F, 12D–F, H, I). Dorsum, venter white to cream-colored; anterior segments without dorsal markings; A6–A8 with diffuse, reddish dorsal marks; A10 with small, brownish mesal spot distally. Venter of A2– A4 with diffuse, light reddish sublateral patches; A10 with pair of brown, submesal spots. LS on A2–A3 light brown, A4–A7 dark brown; SMS light brown to dark amber.

All setae smooth. SMS of A1–A6 hooked, slender, arising from plump chalazae; SMS longest, chalazae largest on A1, A2, becoming smaller posteriorly. A1–A5: LDT plump, with two long, slender, hooked LDS, one microseta between LDS. A1–A8: Spiracle small, slightly raised, dome-like, with minute to very small associated seta (SSp?) anteriorly or mesally.

A1: Single transverse row of eight (L2) or ca nine to ten (L3) SMS mesal to LDTs.

A2–A3: LT broad, plump, fleshy, with seven (L2) or eight (L3) long, robust LS arranged transversely along dorsal surface, ca two (L2) or four (L3) shorter, straight setae basally. L2: Dorsum with anterior row of four SMS, mesal row of four, posterior row of four (A2) or six (A3) between LDTs. L3: Dorsum with anterior row of ca ten (A2) or ~12 (A3) SMS, mesal row of four (A2, A3), posterior row of ca 12 (A2, A3) between LDTs.

A4–A5: LT tuberculiform, with ca five (L2) or 12 (A4, L3), 9–10 (A5, L3) robust, hooked or straight LS apically, dorsally, ca four (L2, L3) short, straight LS basally. LDS intermediate length, hooked. L2: Dorsum with anterior row of four (L2) intermediate-length SMS, mesal row of four, posterior row of four between LDTs. L3: Dorsum with anterior row of ca 12 (A4) or ca nine (A5) intermediate-length SMS, mesal row of four, posterior row of ca ten between LDTs.

A6: LT tuberculiform, with ca five (L2) or six (L3) robust, straight LS apically, ca three short, straight LS basally. LDT with one long, hooked LDS, two to three short, straight LDS. Anterior row of three (L2) or five (L3) intermediate-length SMS, no other SMS.

A7: LT tuberculiform with posterior projection, with ca five (L2) or six (L3) robust, straight LS apically, ca three short, straight LS basally. LDT with one intermediate-length, straight LDS, one to two short, straight LDS. Two pairs of very small setae between spiracles.

A8: LT with four to five short, straight LS (two short, two to three very short). Row of four to five very short setae between spiracles. Posterior margin with ca four (L2) or seven to eight (L3) very short setae.

A9: Two transverse rows of ca four to six short setae.

A10: Dorsum with ca four to five very short setae.

Intraspecific variation (larva). Three variable features warrant notation. (i) As with the larvae of all chrysopids we have studied, L. (L.) nigrilabris larvae exhibit considerable bilateral asymmetry in the numbers and size of setae across the dorsum of body. Odd numbers of setae are common. (ii) The left side of the first abdominal segment (A1) of one L 3 specimen has a fleshy, lateral protuberance (LT?) bearing an elongate, robust, hooked seta. This feature is unusual because LTs rarely occur on the A1 (e.g., in some Chrysopa spp., see Principi 1940, Tsukaguchi 1978), and they have not been reported for leucochrysine larvae. Moreover, LTs almost always bear more than one elongate seta. Thus, we do not know whether this structure is homologous to those on the larva’s subsequent segments (A2– A7) and/or to those on the Chrysopa A1. (iii) Among the first instars, the elongate setae (LS) on the apex of the lateral tubercles are mostly very long, straight, and tapering to pointed tips; occasionally some are hooked.

Biological notes. Adults were collected in disturbed habitats (e.g., coffee plantation, roadside, along trail) at the edges of mature forests. One female specimen, collected in the State of Rio de Janeiro, was maintained in the laboratory for ca two weeks. During that time she laid two batches of eggs. Each batch consisted of a hanging cluster of ca 15– 20 eggs attached to the ends of a series of long intertwined stalks.

The L. (L.) nigrilabris eggs were maintained at room temperature; hatching occurred within ca ten days. The larvae applied and carried dense packets of trash [in this case, frozen & thawed eggs of Ephestia kuehniella (Zeller) ] on their dorsa. The first moult occurred ca six days after hatching, and the second moult ca seven days after the first; cocoons were spun ca 10 days later. Four pharate adults emerged from their cocoons; only one completed ecdysis to the adult stage.

Specimens examined (adults and larvae, in addition to the type listed above). VENEZUELA. Aragua: Parque Nacional Henri Pittier (formerly Parque Nacional Rancho Grande), 1100 m, 11–13.ii.1969, J. Salcedo (1♂, MIZA); idem, 22.v.1967, L. Rodriguez V. (1♀, MIZA, determined by Adams 1979); idem, 9.xi.1990, V. Savini, trampa de luz (1♂, MIZA); idem, 5.vi.1958, F. Fernández Y. (MIZA, determined by Adams 1979]; idem, 16– 31.x.1966, S. S. & W. D. Duckworth (5♀, USNM); idem, 10–21.x.1969, Duckworth & Dietz (1♂, 1♀, USNM); idem, 8.vi.1967, R. W. Poole (2♀, CUIC); idem, 5.viii.1967, R. W. Poole (1♂, CUIC); idem, 20.ix.1973, B. Villegas (1♂, UCD); idem, 24–25.i.1996, white light, J. &. A. Skevington (2♂, 2♀, DEBU); idem, 1150 m, at lights on bldg, 24.ii–12.iii.1996, S. Marshall (1♂, 1♀, DEBU); El Limón, 450 m, 20.iv.1964, E. Osuna (MIZA, determined by Adams 1979); Mérida: Mérida, el. ca 4500 ft., 8.5942°N / 71.1429°W, 22–23.ii.1994, J. P. Donahue (1♀, LACM); Portuguesa: Córdoba 800 m, 24–29.xi.2008, R. Páz, Malaise Trap (1♀, UCOB); Yaracuy: La Cumbre, Chivacoa Council, 1075 m, 10º14’N / 68º58’W, 22.vii.2012, in Coffea arabica, F. Sosa , manual net (1♀, UCOB); idem, 1200 m, 10º16’N / 68º56’W, 11.ii.2013, in Coffea arabica, F. Sosa , manual net (1♂, UCOB); idem, 18.iv.2013, in Coffea arabica, F. Sosa, C. Martins and F. Díaz , manual net (2♂, 1♀, UCOB); El Candelo, Cocorote Council, 1309 m, 10º20’N / 68º49’W, 18.iv.2013, in forest, F. Sosa, C. Martins and F. Díaz, manual net (1♂, UCOB). BRAZIL. Rio de Janeiro: Sta. M. Madalena, Terras Frias, 28.x.2003, G. S. Albuquerque, M. J. Tauber, C. A. Tauber Expedition, Oct.–Nov. 2003 [1♀, 3?sex (lab-reared, pharate adults, incompletely emerged), 9 larvae (Tauber Lot 2003:067, UCB); 15 larvae (Albuquerque Lot 2003:021, UENF)].

Figures

FIGURE 1. Leucochrysa (L.) nigrilabris external features. A, B. Head, frontal; C. Head, dorsal; D. Body, dorsal E, F. Head and thorax, dorsal [note the variation in the coloration of the mesoprescutum and mesoscutellum]; G. Head, thorax, lateral; H. Head, prothorax, lateral [A, D, State of Yaracuy, Venezuela, female; B, C, F – H, State of Rio de Janeiro, Brazil, female; E, State of Aragua, Venezuela, female].

FIGURE 2. Leucochrysa (L.) nigrilabris wings. A, B. Forewing, two specimens to illustrate relatively minor variation; C. Hindwing [A, State of Yaracuy, Venezuela; B, C, State of Aragua, Venezuela, all female]. Scale applies to all three images.

FIGURE 3. Leucochrysa (L.) nigrilabris male abdomen, mature specimen. A. Second and third abdominal segments, lateral [note absence of microtholi from S 2, dense microtholi on S 3]; B. Terminal segments, lateral; C. Terminal segments, dorsal [note large brown spots on T 5, T 6, base of T 7 (arrow)]; D. Terminal segments, dorsal [note dorsally fused T 9 + ectoproct, without suture or distal cleft. The U-shaped darkening is the distal margin of S 8 + 9 below]; E. Callus cerci; F. Fused sternite 8 + 9, ventral, with sclerotized terminal plates extended distally [all, Aragua, Venezuela]. Abbreviations: c. c., callus cerci; d. a., dorsal apodeme of ninth tergite + ectoproct; s. p., sclerotized plate at tip of S 8 + 9; S 2, S 3, second and third sternites; S 8 + 9, fused eighth and ninth sternites (in parentheses, out of focus through dorsal surface); tr, trichobothria; T 5, T 7, T 8, fifth to eighth tergites; T 9 + ect, fused ninth tergite and ectoproct.

FIGURE 4. Leucochrysa (L.) nigrilabris male genitalia, mature specimen. A. Gonarcus, posterodorsal (mediuncus removed); B. Gonarcal complex, posterior (view into gonosaccus from behind, through membrane) [note the mediuncal beak and flanges extend forward (away) and downward]; C. Gonarcal complex, dorsal [view into gonosaccus from above, unobscured by membrane, base of mediuncus in focus, mediuncal beak present (extending forward), but out of focus]; D. Gonarcal complex, frontal [note the gonosaccus is below the mediuncal flanges; the gonosetae are within the gonosaccus]; E. Gonarcal complex, lateral; F. Hypandrium internum (comes very reduced or absent) [all, Aragua, Venezuela]. Abbreviations: bk, beak-like tip of mediuncus; fl, mediuncal flange; gc, gonocornu, encased in membrane; g. a., gonarcal apodeme; g. b., gonarcal bridge; g. p., gonarcal process (membrane attachment); gsac, gonosaccus; gse, gonosetae (within gonosaccus); mu, mediuncus (fused rods and beak); mu-b, sclerotized base of mediuncus.

FIGURE 5. Leucochrysa (L.) nigrilabris male abdomen, teneral specimen. A. Terminal segments, lateral (arrows mark the large brown spots on T 6, base of T 7); B. Gonarcal complex, frontal; C. Gonarcal complex, lateral [note thorny membrane over gonarcal bridge and gonocornua]; D. Gonarcal complex, lateral (thorny membrane above gonarcus partially removed); E. Tip of sternite 8 + 9, dorsal, with sclerotized terminal plates extended distally, hypandrium internum (attached to small membrane that had been everted beneath the sclerotized plate); F. Sclerotized plate at tip of S 8 + 9, lateral (all, Aragua, Venezuela). Abbreviations: bk, beak-like tip of mediuncus; c, comes; cleft, cleft separating distal margins of T 9 + ect; c. c., callus cerci; d. a., dorsal apodeme of T 9 + ect; fl, mediuncal flange; gc, gonocornu; gs, gonarcus; gsac, gonosaccus with robust gonosetae on bulbous chalazae; g. a., gonarcal apodeme; g. b., gonarcal bridge; h. i., hypandrium internum; mu-b, sclerotized base of mediuncus; mu-d, elongate distal section of mediuncus; s. p., sclerotized plate at tip of S 8 + 9; S 8 + 9, fused eighth and ninth sternites; S 9, ninth sternite (tip); T 7, seventh tergite; T 9 + ect, fused ninth tergite and ectoproct ..

FIGURE 6. Leucochrysa (L.) nigrilabris female abdomen. A. Fifth to terminal segments, lateral; B-D. Seventh to terminal segment, lateral [A, B, mature specimens, State of Rio de Janeiro, Brazil; C, D, teneral specimens, State of Aragua, Venezuela]. Abbreviations: g. l., gonapophysis lateralis; sg, subgenitale; sp, spermatheca; S 7, seventh sternite; T 5, T 8, fifth and eighth tergites; T 9 + ect, fused ninth tergite and ectoproct.

FIGURE 7. Leucochrysa (L.) nigrilabris female abdominal and genitalic structures. A. Callus cerci; B. Subgenitale, posterior; C. Spermatheca, lateral; D. Genitalia, ventral [A, B, D, State of Rio de Janeiro, Brazil, mature specimen; C, State of Aragua, Venezuela]. Abbreviations: base, sclerotized base of subgenitale; b. c., bursa copulatrix; b. gl., bursal gland; inv, spermathecal invagination; lobe, rounded lobe of subgenitale; sg / b. c., heavy folded membrane [distally (right): dorsum of subgenitale, proximally (left): venter of bursa copulatrix]; sp, spermatheca; sp. d., spermathecal duct; tr, trichobothria; vel, velum.

FIGURE 8. Leucochrysa (L.) nigrilabris colleterial complex. A. Colleterial complex, lateral; B. Base of colleterial complex, lateral (pulled posteriorly); C. Transverse sclerite, posterior. [all, State of Rio de Janeiro, Brazil, mature specimen]. Abbreviations: acc. gl., colleterial accessory gland; acc. gl. d., duct of colleterial accessory gland; c. d., common duct (colleterial reservoir and accessory gland to transverse sclerite); col. gl., colleterial gland; col. r., colleterial reservoir; tr. s., transverse sclerite.

FIGURE 9. Prothorax of third instar of four Leucochrysa (Leucochrysa) species. A. L. (L.) nigrilabris; B. L. (L.) insularis; C. L. (L.) varia; D. L. (L.) boxi [A, C, D, State of Rio de Janeiro, Brazil; B, North Carolina, United States].

FIGURE 10. Leucochrysa (L.) nigrilabris first instar. A. Head, dorsal; B. Head, ventral; C. Head, lateral; D. Thorax, dorsal; E. Abdominal segments 1 to 5, dorsal; F. Abdominal segments 4 - 10, dorsal; G. Body, lateral; H. Body, ventral [all, State of Rio de Janeiro, Brazil]. Abbreviations: Ax, abdominal segment number; AxRx, setal row number.

FIGURE 11. Leucochrysa (L.) nigrilabris second instar. A. Head, dorsal; B. Head, lateral; C. Thorax, dorsal; D. Abdominal segments 1 to 4, dorsal; E. Abdominal segments 4 – 10, dorsal; F. Body, lateral [all, State of Rio de Janeiro, Brazil]. Abbreviations: Ax, abdominal segment number; AxRx, setal row number.

FIGURE 12. Leucochrysa (L.) nigrilabris third instar. A. Head, dorsal; B. Head, ventral; C. Thorax, dorsal; D. Abdominal segments 1 to 3, dorsal; E. Abdominal segments 3 – 7, dorsal; F. Abdominal segments 6 - 10, dorsal; G. Head, lateral; H. Body, ventral; I. Body, lateral [all, State of Rio de Janeiro, Brazil]. Abbreviations: Ax, abdominal segment number; AxRx, setal row number.

FIGURE 19. Species-specific genital characteristics of Leucochrysa (L.) insularis (left column) and L. (L.) nigrilabris (right column). A, B. Gonarcal complex; C, D. Subgenitale [A, C. Dominican Republic; B, D. Aragua, Venezuela]. Abbreviations: bk, beak-like tip of mediuncus; gc, gonocornu; g. a., gonarcal apodeme; g. b., gonarcal bridge; mu, mediuncus (fused rods and beak); sg-bl, bilobed, dorsal process of subgenitale; sg-v, protruding ventral process of subgenitale; sp, spermatheca.