Leucochrysa (Nodita) azevedoi
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- Leucochrysa azevedoi Navás, 1913: 97; Type: missing; original description: “Brasil: Rio de Janeiro, Agosto de 1911. R.P. Joaquín da Silva Tavares S.J. (Col. m.)”. Navás (1912–1913: 303) [species list].
- Nodita azevedoi (Navás). First combination in Nodita by Navás (1928: 111) [collection record: Guayaquil, Ecuador, May 1926, probably misidentified; see below]. Navás (1929: 859) [collection record: Prov. de Rio de Jan., Coll. v. Bönninghausen, 20-X-1906, M. H., specimen probably destroyed]; Penny (1977: 25) [species list].
- Leucochrysa (Nodita) azevedoi Navás (Brooks and Barnard 1990: 277) [species list]; Oswald (2007) [catalog listing, nomenclature]; Legrand et al. (2008: 117) [probably Nomen Dubium]; Mantoanelli et al. (in press) [larval descriptions]. Here: confirmed as a valid species.
The 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 find it in the Muséum national d’Histoire naturelle, Paris (MNHN). It is reasonable 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 identified 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 Nodita camposi showed significant differences.
Earlier, we had noted that Navás had confused Leucochrysa (Nodita) camposi and azevedoi (see Legrand et al. 2008: 116). That is, he had labeled a specimen of Leucochrysa (Nodita) camposi in the MNHN as Nodita azevedoi; this specimen had been collected from the Nodita camposi type locality in 1930. [Note: Navás may also have similarly misidentified 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 Leucochrysa (Nodita) azevedoi. Indeed our specimens were collected in the state of Rio de Janeiro, where Leucochrysa (Nodita) azevedoi was originally collected; moreover our specimens fit Navás’s original description of Leucochrysa azevedoi well. Although Navás’s type of Leucochrysa azevedoi is missing, the similarities in external features, coupled with his confusion of the two species, give us confidence 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 misidentifications, we now consider that Leucochrysa (Nodita) azevedoi ranges throughout coastal Brazil [and probably also into the state of Pará] and that Leucochrysa (Nodita) 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 Leucochrysa azevedoi neotype, a specimen (male) from the Brazilian state where the original type specimen was collected. Its labels read: (1) “Brazil: R[io de] J[aneiro], Campos dos Goytacazes, Est. Exp. PESAGRO (Estação Experimental da Empresa de Pesquisa Agropecuária do Estado do Rio de Janeiro) [21°44'55"S, 41°18'30"W], 06/IV/2002”; (2) “G. S. Albuquerque, Collector”; (3) “NEOTYPE Leucochrysa azevedoi Navás 1913, desig. C.A. Tauber, G.S. Albuquerque 2010.” In the Coleção Entomológica Pe. Jesus Santiago Moure (Universidade Federal do Paraná, Curitiba, PR, Brazil).
Externally, Leucochrysa (Nodita) 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 flagellomeres, 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 (Nodita) azevedoi is slightly smaller than many of these species, including Leucochrysa (Nodita) camposi and Leucochrysa (Nodita) morenoi (wing length 17.7–19.5 mm versus ~21.3–21.6 for Leucochrysa (Nodita) camposi and morenoi), but it can only be identified with accuracy by its genitalic characters (male and female) (Figs 4–6). In the male Leucochrysa (Nodita) azevedoi, the gonarcal arms are broadly spread; they extend almost the full width of the segment. The 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 Leucochrysa (Nodita) camposi gonocornua. The 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. The hypandrium internum has a broad V-shape. Females can be recognized by their tubular spermatheca, small bursa, relatively small, fluted bursal duct. The 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.
[Measurements: head, thorax, abdomen, wings (n=4), genitalia (n=2 mature males, 2 mature females)].
Head: 1.7–1.9 mm wide (including eyes); ratio of head width to eye width = 2.3–2.5:1. Vertex raised, with smooth surface, prominent, rounded posterior fold, without setae. Antenna 24.7 mm (~1.3 times length of forewing); scape longer than broad, (0.41–0.44 mm long, 0.32–0.34 mm wide), width = 3.2–4.4× distance between scapes, with three to four long setae distally on dorsal surface, shorter setae laterally; lateral margin fairly straight, mesal margin straight basally, curved outward distally; pedicel ~0.10 mm long, ~0.15 mm wide (at widest point); proximal flagellomeres short (segments 1, 2, 3: length = 0.85–1.0 times width), with three to four concentric rings of setae; middle and distal segments becoming longer (segments 6–8: length = 1.7–2.1 times width; distal segments: length = 3.0–3.2 times width), with four concentric rings of setae. Distance between scapes 0.08–0.10 mm; distance between tentorial pits 0.57–0.61 mm; length of frons (midway between scapes – midway between tentorial pits) 0.44–0.55 mm. Frons relatively flat mesally, with scalloped fold below toruli; surface smooth to slightly textured, with short setae. Clypeal margins straight; surface slightly textured, not horizontally striated. Labrum with distal margin indented mesally; dorsal surface smooth, rounded, setose distally. Ratio of genal length to distance between tentorial pits = 0.26–0.33:1.
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; flagellum 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.
Thorax: 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, diffuse, reddish brown spots mesally; metascutellum unmarked or with pair of small, diffuse 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 gradate vein). Two series of gradate veins; 9 inner gradates, 8–10 outer gradates; Eight b’ cells (cells beneath pseudomedia after second intramedian cell). Three 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); five 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.
Abdomen: Distal segments (beyond A4) moderately expanded; pleural region ca. twice height of sternites. Sternites, tergites with microsetae moderately dense; pleural region with very dense microsetae. Male: S6 approx. 1.2–1.4 times longer than tall, S7 approx. 1.0–1.1 times longer than tall (lateral view); female: S6 approx 1.4–1.7 times longer than tall, S7 approx. 1.7–2.0 times longer than tall. Tergites narrow, roughly rectangular, with lighter setae and longer microsetae than on sternites. Spiracles oval externally; atria not enlarged. Coloration: Dorsum with medial yellow stripe, deep red sublateral markings; sides green with deep red marks posterodorsally; venter white; callus cerci white; trichobothria pale.
Male: Callus cerci oval to round, 0.20–0.27 mm diameter (range), with 29–37 relatively thin trichobothria of various lengths. Sternites 3–8 (not S1 or S2) with microtholi. T9+ectoproct truncate distally, fused mesally, midline without deep cleft, setae robust throughout, slightly smaller proximally than distally; ventral section of T9+ectoproct with broad, elongate proximal extension reaching full length of A8; proximal section well sclerotized, with apodeme heavy, unbranched, extending around proximal margin of callus cerci. S8+9 fused, without suture in mature specimens; S9 without microtholi; S8 much shorter, slightly taller than S9; S8+9 (lateral view) with proximal margin straight, rounded dorsally, acute ventrally; distal margin straight, with rounded apices, approximately 1/3 height of proximal margin. Setae on S9 slightly heavier than those on S5-S8; terminus of S9 without gonocristae. Subanal region membranous, with small striations around anus, no setae. Gonarcal complex connected to terminus of ectoproct by long, clear, smooth membrane that attaches to gonarcal bridge around base of gonocornua; gonocornua protruding through membrane; section of membrane below mediuncus extending interiorly and holding hypandrium internum. Gonarcus robust, very broadly arcuate, with lateral apodemes extending laterally almost entire width of T9+ect; gonarcal apodemes robust, stiff, slightly concave dorsally, rounded distally. Gonocornua broadly attached to gonarcal bridge, mesal to mediuncus, extending upward, away from mediuncus, short, stout, with inner margin curving outward, outer margin perpendicular to bridge, with lateral knob distally. Mediuncus largely membranous, dorsally with pair of longitudinal ridges, trough between, distally with pair of rounded lobes, recurved arcessus with heavy, rounded, beak-like tip; sides of mediuncus membranous, expanded, forming hollow depression; membrane inside depression bearing ~3 pairs of robust, medium-length setae on small chalazae; distal margins of mediuncal membrane with dense patches of relatively long, fine setae. Entoprocessus, tignum, gonapsis, pseudopenis, spinellae, gonocristae, gonosaccus absent. Hypandrium internum with broad V-shape, apex rounded, large, delicate, lightly sclerotized; comes elongate, thin, sometimes not visible.
Female: Callus cerci round, 0.16–0.22 mm maximum diameter, with 25–33 trichobothria of mixed length. Tergite 8 roughly quadrate (lateral view), similar in depth to T6. Tergite 9+ectoproct elongate; posterior margin: dorsal half straight, almost perpendicular to dorsal margin of tergite, ventral half indented, angled inward, straight; ventral margin rounded, extending slightly below gonapophyses laterales. Sternite 7 with transverse weakness midlength, dorsal margin straight, not tapering distally; terminus unmodified, with terminal (posteroventral) setae slightly more dense, robust, and longer than other setae. Gonapophysis lateralis angled dorsally, rounded distally, ventrally, ~0.65–0.68 length of T9+ectoproct; inner membranous surface not expandable, with ~two vertical rows of short setae. Colleterial gland smooth-walled, delicate, with fluted, globate reservoir or secondary gland attached via a narrow duct immediately before transverse sclerification. Transverse sclerification robust, flat, platform-like with lateral margins upturned, with coarse longitudinal striations. Bursa copulatrix saccular, consisting of robust, densely folded membrane, extending anteriorly over spermatheca, into distal section of S7; pair of robust, branching, elongate, tubular bursal glands, connected to dorsolateral surface of bulbous end of bursa via short, very narrow duct. Spermatheca tubular, bent, thick at opening, tapering slightly (0.17–0.20 mm diameter at mouth, 0.12 mm diameter at midsection), 0.8–0.9 mm long, with broad, shallow (0.06–0.09 mm) invagination; slit along entire side of spermatheca opening to bursa. Spermathecal duct attached to anterior margin of spermatheca; basal section thick, straight, extending directly into base of subgenitale, then gradually tapering at first U-shaped bend, recurving posteriorly with two curves; basal ~one-half to two-thirds well sclerotized; distal section clear, brushy. Subgenitale broad, rounded, with smooth, rigid surface distally, with folded, membranous surface, with terminus broad, bilobed, with shallow, flat depression between lobes (heart shaped in posterior view); small ventral fold on S7 without setae.
Described elsewhere (Mantoanelli et al. in press).
This 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).
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 Leucochrysa (Nodita) 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 Leucochrysa (Nodita) amazonica was based on a single teneral male from the lower Amazon. Our study shows that Adams’ figure of the specimen from Pará is typical of Leucochrysa (Nodita) azevedoi.]
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.
In addition to the specimens listed below, we examined one specimen (a male) from the State of Rio Grande do Sul. This specimen’s background color and markings are lighter than those of the other specimens; in addition, the beak on its mediuncus is sharp and straight, not blunt and decurved as in Leucochrysa (Nodita) azevedoi. A determination of whether this specimen represents a separate species or a variant population of Leucochrysa (Nodita) azevedoi awaits the availability of additional specimens.
Adult specimens examined
In addition to the neotype listed above: Brazil. Rio de Janeiro: Campos dos Goytacazes, VI/22/1999, R. R. Moraes (1♂, 1♀, Tauber Research Collection, TRC); Campos dos Goytacazes, Est. Exp. PESAGRO (Estação Experimental da Empresa de Pesquisa Agropecuária do Estado do Rio de Janeiro, 21°44'55"S, 41°18'30"W, 14m, VII-2005 – III-2007, G. S. Albuquerque, J. S. Multani, Collectors (2♂♂, 2♀♀, alcohol, UENF; 3♂♂, 3♀♀, alcohol, 1♀ pinned, TRC). São Paulo: Jaboticabal, FCAV, 10 July 1998, Freitas, S., 10 (1♂, California Academy of Sciences, CAS). Mato Grosso: Itiquira, P. E. Michelin, 18 July 1998, Freitas, S., M10 (1♂, CAS).
Latinized proper name, genitive. Navás named the species in honor of Ignacio de Azevedo (1528–1570), a Portuguese Jesuit who labored in Brazil and was killed along with 39 Jesuit companions, while sailing near the Canary Islands on a return voyage from Rome to Brazil. They are known as the “Fourty Martyrs of Brazil”.
- Catherine A., T; Gilberto S., A; Maurice J., T; 2011: Nomenclatorial changes and redescriptions of three of Navás’ Leucochrysa (Nodita) species (Neuroptera, Chrysopidae) ZooKeys, 92: 9-33. doi: 10.3897/zookeys.92.828
- Navás L (1912–1913) Les Chrysopides (Ins. Névr.) du Musée de Londres [Ia]. Annales de la Société Scientifique de Bruxelles 37(pt. 2): 292–330.
- Navás L (1928). Insectos neotropicos. 4.a serie. Revista Chilena de Historia Natural 32:106-128.
- Navás L (1929). Insectos del Brasil. 3.a serie [IIIa]. Revista do Museu Paulista, Sao Paulo 16:857-864.
- Penny N (1977) Lista de Megaloptera, Neuroptera e Raphidioptera do México, América Central, ilhas Caraíbas e América do Sul. Acta Amazonica 7(4) (Suplemento): 1–61.
- Brooks S, Barnard, P (1990) The green lacewings of the world: a generic review (Neuroptera: Chrysopidae). Bulletin of the British Museum of Natural History, Entomology 59:117-286.
- Oswald J (2007) Neuropterida Species of the World. Version 2.0. http://lacewing.tamu.edu/Species-Catalogue/. [accessed on Nov. 17, 2010]
- Legrand J, Tauber C, Albuquerque G, Tauber M (2008) Navás’ type and non-type specimens of Chrysopidae in the MNHN, Paris [Neuroptera]. Revue française d’Entomologie 30:103-183.
- Mantoanelli E, Tauber C, Albuquerque G, Tauber M (in press) Larvae of four species of Leucochrysa (Nodita) from Brazil’s Atlantic coast (Neuroptera: Chrysopidae: Leucochrysini). Annals of the Entomological Society of America.
- Navás L (1913) Crisópidos sudamericanos. Brotéria (Zoológica) 11: 73–104, 149–168.
- Monserrat V (1985) Lista de los tipos de Mecoptera y Neuroptera (Insecta) de la colección L. Navás, depositados en el Museo de Zoología de Barcelona. Miscellània Zoològica 9:233-243.
- Freitas S, Penny, N (2001) The green lacewings (Neuroptera: Chrysopidae) of Brazilian agro-ecosystems. Proceedings of the California Academy of Sciences 52:245-395.
- Multani J (2008) Diversidade e abundância de crisopídeos (Neuroptera, Chrysopidae) e interações com presas, parasitóides e fatores abióticos em pomares de goiaba em Campos dos Goytacazes, RJ. PhD Dissertation, Universidade Estadual do Norte Fluminense, Campos dos Goytacazes, RJ, Brazil.
- Adams P (1987) Studies in Neotropical Chrysopidae (Neuroptera) III. Notes on Nodita amazonica Navás and N. oenops, n. sp. Neuroptera International 4:287-294.