Nomenclatorial changes and redescriptions of three of Navás’ Leucochrysa (Nodita) species (Neuroptera, Chrysopidae)

Abstract Three species that Navás described – Leucochrysa (Nodita) azevedoi Navás, 1913, Leucochrysa (Nodita) camposi (Navás, 1933) and Leucochrysa (Nodita) morenoi (Navás, 1934) – have received recent taxonomic attention. All three have many similar external features; indeed Navás himself, as well as subsequent authors, have confused the species with each other. Here, (a) misidentifications are corrected; (b) a neotype of Leucochrysa azevedoi is designated; (c) Leucochrysa (Nodita) morenoi, previously synonymized with Leucochrysa (Nodita) camposi, is recognized as a valid species [Reinstated status] All three species are redescribed and illustrated, with special emphasis on the types. Leucochrysa (Nodita) azevedoi was found to be relatively common in agricultural areas along Brazil’s Atlantic coast. The two other species are known only from their type localities: Leucochrysa (Nodita) camposi – coastal Ecuador, and Leucochrysa (Nodita) morenoi – Quito, Ecuador.


Introduction
Recent fi eld surveys of natural enemies associated with insect pests in neotropical agroecosystems show that the chrysopid fauna is very rich and taxonomically complex (e.g., in Brazil: Freitas and Penny 2001, Multani 2008. Th e Leucochrysines are especially diverse and diffi cult to identify; as a consequence, the literature on the group is rife with synonymies, errors, and uncertainty. To help clarify the systematics of the group, we have begun examining the leucochrysine types and comparing them with recently collected or reared specimens. Th is is a slow process. Here we report the results of our studies with three Leucochrysa (Nodita) species that Navás described. All three resemble each other closely, and have been misidentifi ed and confused with each other in both old and recent literature (Navás 1928, Freitas and Penny 2001, Legrand et al. 2008). One of these species occurs in agricultural settings throughout Brazil; thus timely clarifi cation of its taxonomy is important.
Type Material. Th e original type remained in Navás collection (Navás 1913: 97); however, it does not exist in the Navás collection at the Natural History Museum of Barcelona now (Monserrat 1985: 240). Also, Legrand et al. (2008: 117) were unable to fi nd it in the Muséum national d'Histoire naturelle, Paris (MNHN). It is reason-   able to assume that the specimen has been destroyed; therefore, Legrand et al. (2008) considered the species a Nomen Dubium.
Recently we attempted to identify a Leucochrysa (Nodita) species from orchards in northern Rio de Janeiro State. Using the key by Freitas and Penny (2001), we identifi ed the specimens as Leucochrysa (Nodita) camposi, a species that was described from coastal Ecuador; our specimens matched the drawings and description included with the key. However, comparison of our specimens with the type of N. camposi showed signifi cant diff erences.
Earlier, we had noted that Navás had confused L. (N.) camposi and azevedoi (see Legrand et al. 2008: 116). Th at is, he had labeled a specimen of L. (N.) camposi in the MNHN as Nodita azevedoi; this specimen had been collected from the N. camposi type locality in 1930. [Note: Navás may also have similarly misidentifi ed another specimen collected from the same locality in 1926 (Navás 1928: 111), but we have not seen that specimen. ] Navás's errors then led us to consider whether our specimens could be L. (N.) azevedoi. Indeed our specimens were collected in the state of Rio de Janeiro, where L. (N.) azevedoi was originally collected; moreover our specimens fi t Navás's original description of L. azevedoi well. Although Navás's type of L. azevedoi is missing, the similarities in external features, coupled with his confusion of the two species, give us confi dence that Navás's original description applies to the specimens we have at hand from northern Rio de Janeiro State.
As a result of our studies and the correction of Navás misidentifi cations, we now consider that L. (N.) azevedoi ranges throughout coastal Brazil [and probably also into the state of Pará] and that L. (N.) camposi occurs on the west coast of northern South America (currently known only from Ecuador). Now, to stabilize the nomenclature of these species, we designate as the L. azevedoi neotype, a specimen (male) from the Brazilian state where the original type specimen was collected. Its labels read: (1)  Diagnosis. Externally, L. (N.) azevedoi adults closely resemble several Leucochrysa (Nodita) species that have a red to reddish brown dorsolateral stripe on the scape, dark brown to black ventrolateral marks on the basal fl agellomeres, red to reddish brown marks on the sides of the raised portion of the vertex, a darkened section in the middle of the Radial sector (forewings and hindwings), and darkened terminal veinlets along the posteroapical margin of the hindwings (Figs 1A, 2-3). Leucochrysa (N.) azevedoi is slightly smaller than many of these species, including L. (N.) camposi and L. (N.) morenoi (wing length 17.7-19.5 mm versus ~21.3-21.6 for L. (N.) camposi and morenoi), but it can only be identifi ed with accuracy by its genitalic characters (male and female) . In the male L. (N.) azevedoi, the gonarcal arms are broadly spread; they extend almost the full width of the segment. Th e gonocornua are located mesally on the gonarcal bridge, well within the span of the mediuncus, and they are directed upward from the bridge, not laterally from the bridge as are the L. (N.) camposi gonocornua. Th e mediuncus is heavy; the beak is borne on a posteriorly projecting ventral plate (arcessus) that angles back towards the gonarcal arch; the beak is broad and blunt, and the membranous arms extending laterally from below the beak are rounded and each bears a lateral patch of setae. Th e hypandrium internum has a broad V-shape. Females can be recognized by their tubular spermatheca, small bursa, relatively small, fl uted bursal duct. Th e bursa has two elongate, branched, tubular glands attached to its dorsal surface, and the subgenitale is heavy-based; the terminus has a thick neck, paired short, dorsal lobes and a short protrusion. . Leucochrysa (Nodita) azevedoi, Male (Rio de Janeiro, TRC). A Abdomen, lateral B Gonarcus, lateral C Hypandrium internum, dorsal. Abbreviations: c.c., callus cerci; d.a. dorsal apodeme gc gonocornu g.a. gonarcal apodeme g.b gonarcal bridge mu mediuncus s.memb setose membrane lateral to mediuncus beak tip tip of mediuncus beak S8+9 fused eighth and ninth sternites T8 eighth tergite T9+ect fused ninth tergite and ectoproct.
Head coloration: Antennae: scape cream to golden with light greenish tinge, broad red to reddish brown dorsolateral stripe extending full length of scape, onto anterolateral corner of dorsal torulus; pedicel cream with dark brown band distolaterally; fl agellum cream with black setae, basal ~15 antennomeres with large dark brown to black mark ventrolaterally, fading on antennomeres ~16-20; marks forming prominent dark ventrolateral stripe. Vertex with central area raised, yellowish to greenish posteriorly, with prominent, triangular red to reddish brown mark laterally; area around raised area light green, unmarked. Frons, clypeus white, unmarked; labrum cream to amber; gena white, with narrow longitudinal red to reddish brown stripe immediately below tentorial pit. Torulus cream to amber. Maxilla, maxillary palpi, labium, labial palpi white to cream.
Th orax: Cervix small, largely withdrawn below prothorax, light green, with small reddish brown lateral marks. Prothorax (sclerotized region) 0.86-0.93 mm long; 1.1-1.2 mm wide; ratio of length : width = ~0.69-0.77:1; setae thin, long, light golden; pronotum light green, with broad, golden yellow mesal stripe, with pair of lateral to sublateral elongate, red to reddish brown marks with irregular margins, sometimes with pair of small anterolateral reddish marks, pair of small, mesal reddish brown marks. Mesothorax, metathorax light green with golden yellow mesal stripe, sometimes with white; mesoprescutum with pair of crescent-shaped reddish brown marks; mesoscutum with pair of bold, submesal dark red to reddish brown marks; mesoscutellum unmarked; metascutum unmarked or with pair of small, diff use, reddish brown spots mesally; metascutellum unmarked or with pair of small, diff use reddish brown spots laterally. Legs unmarked, with golden setae; coxae, femora cream to white; tibiae light green, tarsi golden. Tarsal claws with deep cleft, elongate, with quadrate base.
Wings: Forewing 17.7-19.5 mm long, 6.4-6.9 mm wide (at widest point); ratio of length : maximum width = 2.7-2.9:1. Costal area moderately broad; tallest costal cell (#7-9) 1.4-1.5 mm tall, 2.0-2.2 times width, 0.23-0.24 times width of wing (midwing). First intramedian cell triangular, 0.4-0.7 width of third median cell. First radial crossvein distal to origin of radial sector (Rs); radial area (between Radius and Rs) with single row of 16-18 closed cells; tallest cell (#7-8) 2.2-2.8 times taller than wide. No crassate veins; 5-6 b cells (= cells beneath Rs, not including an inner gra- A View from above, with gonocornua projecting from the gonarcal bridge to the right, mediuncus extending downward, away from camera [Note heavy membrane above gonarcus extending over gonocornua] B Gonarcus with gonocornua projecting directly upward towards camera, mediuncus fully extended to right, covered with heavy membrane extending from gonarcal bridge C Gonarcus, view from beneath, with gonocornua extending to left D Mediuncus, view from below the tip of the beak: D1 View at level of the beak; note the setose membranous areas lateral to the tip of the beak D2 View at level above tip of beak; note membranous gonosaccus and gonosetae well above beak. Abbreviations: gc gonocornu gse gonoseta g.a. gonarcal apodeme g.b. gonarcal bridge mu mediuncus s.memb, setose membrane lateral to mediuncus beak tip tip of mediuncus beak. date vein). Two series of gradate veins; 9 inner gradates, 8-10 outer gradates; Eight b' cells (cells beneath pseudomedia after second intramedian cell). Th ree intracubital cells (two closed). Membrane clear; stigma opaque, marked with brown. Veins green, with black on tips of costal veinlets, basal segment and midsection of Rs, two basal radial crossveins, distal arm of im2, bases of marginal forks.
Hindwing 15.3-16.9 mm long, 4.7-5.4 mm wide. Two series of gradate veins; 7-8 inner, 7-8 outer; 14-16 radial cells (counted from origin of Radius, not false origin). Five to six b cells (including small b1 cell); fi ve to six b' cells beyond second intramedian cell; three intracubital cells (two closed). Membrane clear, with posteroapical margin streaked with light brown; stigma pronounced, marked with brown basally. Veins mostly light green, but dark brown on tips of costal veinlets, midsection of Rs, tips of marginal forks, vein along distal half of posterior margin.
Larvae. Described elsewhere (Mantoanelli et al. in press). Biology. Th is species has been collected in orchards in the states of Rio de Janeiro, São Paulo, Mato Grosso and Rio Grande do Sul, in some cases in relatively large numbers. Its seasonal occurrence in orchards of coastal Brazil was assessed (Multani 2008).
Distribution. Currently known only from Brazil. We have seen specimens from the States of Rio de Janeiro, Matto Grosso, and São Paulo [also see Freitas and Penny 2001, as L. (N.) camposi]. Probably, the species also occurs in the State of Pará. [Note: Adams (1987) considered a specimen that he studied from Pará (illustrated in Fig. 18) to be an abnormal variant of Leucochrysa (Nodita) amazonica (Navás). It is not clear why he did so; Navás's description of L. (N.) amazonica was based on a single teneral male from the lower Amazon. Our study shows that Adams' fi gure of the specimen from Pará is typical of L. (N.) azevedoi.] Variation. See images on our project site at www.morphobank.org. In teneral males, S8 and S9 are only partially fused; the segments are fully fused, but well demarcated in mature males. When teneral, the gonarcus is thin and delicate; the gonocornua are small and closely aligned, adjacent to the mediuncus; and the setose lateral extensions below the beak of the mediuncus are not visible, but the beak is distinct.
Type material. Th e N. camposi lectotype, a male, is badly discolored, but its wings are in good condition, its body is mature, and the genitalia are well sclerotized. A female specimen collected one year later, at the same locality as the type, is also mature. Diagnosis. L. (N.) camposi can be distinguished from L. (N.) azevedoi by its somewhat larger size and more robust appearance (see above) and genitalic characters (male and female) (Figs 8-10). In the male L. (N.) camposi, the gonarcal arms are narrowly arched; the gonarcal arms extend perpendicularly from the gonarcal bridge, not laterally as in L. (N.) azevedoi. Th e gonocornua are located distally on the gonarcal bridge, well outside the span of the mediuncus, and they extend laterally away from the gonarcal bridge. Like L. (N.) azevedoi, the mediuncus is heavy apically and its beak is borne on a plate (arcessus) that bends back toward the gonarcal arch; however, the membrane extending from the beak does not have distinct arms or patches of setae, as are found on L. (N.) azevedoi. Th e hypandrium internum is delicate and has a very narrow U-shape. Th e L. (N.) camposi female can be recognized by its bulbous spermatheca, asymmetrical bursa that extends from its left side over the spermatheca and part of the bursal duct, and its highly coiled bursal duct that extends well into the sixth abdominal segment. Th e bursal glands are elongate, branching and ribbon-like.
Head coloration. Specimen largely discolored with age. Antennae: scape amber colored mesally, noticeably darker laterally, probably a broad reddish brown dorsolateral stripe extending full length of scape; pedicel probably with brownish band distolaterally; fl agellum cream with golden brown setae, basal ~20 antennomeres with large dark brown to black mark ventrally, fading on antennomeres ~20-28; marks forming prominent dark stripe ventrally. Vertex with raised central area discolored, with thin reddish brown streak across frontal margin, trace of large triangular red mark laterally; area between raised area and eyes maybe marked with red. Frons, clypeus white, unmarked; labrum cream to amber; gena discolored, possibly red throughout. Maxilla, maxillary palpi, labium, labial palpi white to cream.
Hindwing 18.5-18.8 mm long, 5.5-6.1 mm wide. Two series of gradate veins; 8 inner, 9 outer; 15 radial cells (counted from origin of Radius, not false origin). Four to fi ve b cells (including small b1 cell); fi ve b' cells beyond second intramedian cell; two intracubital cells (one closed). Stigma golden, marked with brown distally; membrane clear with posteroapical margin streaked with faintly brown tinge. Veins green, except midsection of Rs, tips of marginal forks (posteroapical margin of wing) dark brown to black.
Female: Callus cerci round, 0.26 mm maximum diameter, with 39 slender trichobothria. Tergite 8 roughly quadrate (lateral view), similar in depth to T6. Tergite 9+ectoproct elongate; posterior margin: dorsal one-third straight, perpendicular to dorsal margin of tergite, then indented, angled inward, straight; ventral margin rounded, extending to depth of gonapophyses laterales. Sternite 7 largely damaged. Gonapophysis lateralis angled dorsally, rounded distally, ventrally, ~0.60 length of T9+ect; inner membranous surface not expandable, with sparse, small setae on inner surface. Colleterial gland missing but short, thick duct attached immediately before transverse sclerifi cation. Bursa copulatrix concave, shield-like, lightly folded, thick membranous sac, extending anteriorly over spermatheca, into mid-section of S7, also extending, asymetrically on left side of S7 above ca. basal one-half of coiled bursal duct; with pair of slender, elongate, branching, ribbon-like bursal glands, one connected on either side of bursa, near interior tip of bursa. Bursal duct fl at to thickened, coiled, robust, tapering anteriorly to A6, then coiling back on itself to bursa; membranous throughout. Spermatheca broad, thick, U-shaped tube, thick walled with wide circular opening facing subgenitale, enlarged tire-shaped bulge at open end, prominent slit along entire dorsum, opening to bursal duct (0.16 mm diameter at mouth, narrower at midsection), ~0.5 mm long, with narrow (0.04 mm), moderate length (0.12 mm) invagination. Spermathecal duct uncolored, originating on dorsal surface of spermatheca at terminus of dorsal slit, making two sharp bends, a U-shaped bend in subgenitale, then glandulose thicked portion making broad bend back toward spermatheca. Subgenitale large, rounded, with rigid, folded, heavy membraneous texture, terminus broad, fl at-surfaced, bilobed dorsally (heart shaped in posterior view), with large, fl at transverse depression mesally below lobes, with ventrally protruding tip; small ventral fold above S7 without setae.

Leucochrysa (Nodita) morenoi
Type material. Th e N. morenoi lectotype, a male, is discolored, but otherwise in good condition; the abdomen is cleared and in a vial with glycerin. For this study, we re-examined the types of N. morenoi and N. camposi side-by-side; here, we provide images and a re-description to support our recognition of L. (N.) moreni as a valid species. Diagnosis. L. (N.) morenoi is known only from the type specimen, which is badly discolored with age. At this time, the only reliable way to identify specimens is with (N.) camposi, in the following traits: dark brown to black, lateral marks on the basal ~15 fl agellomeres that give impression of a streak on the exterior edge of the antenna, a darkened section in the middle of the Radial sector (forewings and hindwings), and darkened terminal veinlets along the posteroapical margin of the hindwings. Th e head and thorax are too discolored to determine if there are reddish markings on the vertex, gena, scape, or prothorax as there are on many of the other species with these traits. We suspect that they are not present on this species because the only marks that Navás (1934)  Adult description. Head: 2.12 mm wide (including eyes); ratio of head width to eye width = 1.85:1. Vertex approximately oval, raised slightly, with smooth to slightly textured surface, small posterior fold. Antenna 38 mm long (~1.8 times length of forewing); scape longer than broad, (~0.4 mm wide), width = 2× distance between scapes, with long setae distally on dorsal surface, shorter setae laterally; lateral margin fairly straight, mesal margin straight basally, curved outward distally; pedicel ~0.25 mm long, ~0.18 mm wide (at widest point); proximal fl agellomeres short (segments 1, 2, 3: length = 1.2-1.5 times width), with three to four concentric rings of setae; middle and distal segments becoming longer (segments 9-11: length = 1.6-1.7 times width; distal segments: length = 2.2-2.6 times width), with four concentric rings of setae. Distance between scapes 1.04 mm; distance between tentorial pits 0.65 mm; length of frons (midway between scapes -midway between tentorial pits) 0.77 mm. Frons relatively fl at, with slightly scalloped fold below toruli; surface smooth to slightly textured. Clypeus straight distally; surface slightly textured, not horizontally striated. Labrum with distal margin very slightly concave mesally; dorsal surface smooth, rounded, setose distally. Ratio of genal length to distance between tentorial pits = 0.41:1.
Th orax: Cervix small, largely withdrawn below prothorax, discolored. Prothorax (sclerotized region) 0.83 mm long; 1.03 mm wide; ratio of length : width = 0.56:1; prothorax (extended) 1.5 mm long; setae thin, long, golden; surface discolored, scattered patches of red tinge visible, but not mentioned in original description, small grey area mentioned in original description not apparent now. Mesothorax, metathorax discolored; mesoprescutum with pair of sublateral reddish brown marks (mentioned in original description); mesoscutum with pair of brown streaks on anterior and posterion margin; mesoscutelllum unmarked; metascutum with pair of brownish spots on posterior margin. Legs unmarked, with golden setae, originally described as greenish.
Hindwing 18.7 mm long, 6.2 mm wide. Two series of gradate veins; 9 inner, 9 outer; 17 radial cells (counted from origin of Radius, not false origin). Six b cells (including small b1 cell); seven b' cells beyond second intramedian cell; two intracubital cells (two closed). Membrane clear, probably streaked with light brown tinge along posterodistal margin; stigma pronounced, marked with brown. Veins mostly green (probably), but dark brown on midsection of Rs, tips of crossveins extending from darkened midsection, and posteroapical margin of wing. Abdomen: Distal segments (beyond A4) expanded; pleural region ca. two times height of sternites. Sternites, tergites with microsetae relatively sparse; male: S6 ca. same height as length, S7 approx. 1.1 times taller than long (lateral view). Tergites roughly rectangular, with rounded posterior and lateral margins, shorter setae than on sternites. Spiracles oval externally; atria not enlarged. Coloration: information not available.
Female: Unknown. Larvae and biology. Unknown. Distribution. Currently known only from Quito, Ecuador. Adult specimens examined. Known only from the type. Etymology. Navás named this species in honor of President Gabriel García Moreno of Ecuador (1821-1875). Moreno was a noted statesman, who strongly supported education, science, and the Roman Catholic Church.

Keys to species
Th e latest key to South American Leucochrysa species relies on external features of adults to identify species that occur in Brazilian agroecosystems (Freitas and Penny 2001: 285). As explained above, this key identifi es specimens of L. ) morenoi -are known from only a single specimen each (the types), and these specimens are too badly discolored to be useful in incorporating the species into a key using external characters. Reliable identifi cations require dissection of male and/or female genitalia, and at this time the genitalia of too few species have been studied to develop a useful key.
All three species treated here appear to be related to L. (N.) amazonica and its relatives (see Adams 1987, Freitas andPenny 2001); they are large bodied, apparently have similar body markings, and share a truncated plate at the tip of the mediuncus (arcessus of Freitas and Penny 2001) and elongate, coiled bursal ducts. Future studies should include comparisons among the species that share these traits.