Leucochrysa (Leucochrysa) varia (Schneider, 1851)

Tauber, Catherine A., Sosa, Francisco & Albuquerque, Gilberto S., 2013, Two common and problematic leucochrysine species - Leucochrysa (Leucochrysa) varia (Schneider) and L. (L.) pretiosa (Banks) (Neuroptera, Chrysopidae): redescriptions and synonymies, ZooKeys 310, pp. 57-101 : 60-75

Leucochrysa (Leucochrysa) varia (Schneider, 1851) Figs 114

Chrysopa varia Schneider [1851] ( 1851: 154, Plate 58) original description: "Brasilia, ubi a Dr. Clausen inventa; (collect. E. de Selys-Longchamps!)". Walker (1853: 268) brief redescription, collection records; McLachlan (1868a: 270) taxonomic note; Brooks and Barnard (1990: 247) confirmed type species of genus Leucochrysa .

Leucochrysa varia (Schneider). McLachlan (1868b: 208) first reference to combination, Leucochrysa varia designated as type species of genus; Navás (1917: 279) species list; Navás (1922a: 392) collection record; Navás (1922b: 89) collection record; Navás (1924: 28) collection record; Navás (1926: 12) collection record; Navás (1928a: 131) collection record (probably in error); Navás (1928b: 111) collection record; Navás (1928c: 35) collection record; Navás (1929: 862) collection records; Navás (1932: 57) collection record; Banks (1944: 31) note on type, species distribution records (probably in error); Banks (1945: 168) note on geographic distribution, comparison with Leucochrysa pretiosa ; Penny (1977: 23) species list; Adams (1979: 97) probable occurrence in Mexico.

Allochrysa varia (Schneider). Banks (1910: 150) first reference to combination; Navás (1912-1913: 313) species list, collection record; Navás (1913: 156) brief redescription; Navás (1917: 279) genus synonymized with Leucochrysa .

Leucochrysa (Leucochrysa) varia (Schneider). Brooks and Barnard (1990: 248, 277, figs 498-507) subgeneric determination, species list, figures; Freitas and Penny (2001: 282, 354, fig. 43) brief redescription, collection records, figures -- based on misidentified specimens in the FCAV-UNESP, not Leucochrysa (Leucochrysa) varia ; Mantoanelli et al. (2006: 7) description of larvae, developmental data, analysis of color polymorphism; Mantoanelli and Albuquerque (2007: 302) behavioral and developmental data; Oswald (2007) catalog listing.

Chrysopa internata Walker [1853] ( 1853: 252) original description: "a. Brazil. b. ----------? From Mr. Children’s collection". McLachlan (1868a: 269) synonymy with Chrysopa ampla Walker.

Leucochrysa internata (Walker). Navás (1912-1913: 303) first reference to combination, species list; Navás (1913: 102) brief redescription, reversal of McLachlan’s 1868a synonymy with Leucochrysa ampla ; Kimmins (1940: 444) identification of Walker’s specimen b, without locality data, as primary type; Penny (1977: 23) species list; Brooks and Barnard (1990: 277) apparently listed in error as a synonym of Leucochrysa (Leucochrysa) walkerina Navás; Oswald (2007) catalog listing as a synonym of Leucochrysa (Leucochrysa) walkerina as per Brooks and Barnard (1990). Syn. n.

Allochrysa internata (Walker). Banks (1914-1915: 625) first reference to combination; Navás (1917: 279, internata not specifically mentioned) genus synonymized with Leucochrysa ; Oswald (2007) catalog listing as synonym of Leucochrysa (Leucochrysa) walkerina , apparently in error.

Nodita internata (Walker). Navás (1917: 280) first reference to combination, species list (probably an error); Oswald (2007) catalog listing as synonym of Leucochrysa (Leucochrysa) walkerina .

Chrysopa ampla Walker [1853] ( 1853: 268) original description: "a. Georgia. From Mr. Abbot’s collection. b. -------? From Mr. Children’s collection". McLachlan (1868a: 270) taxonomic note and mention of close relationship with Chrysopa varia . Kimmins (1940: 444) identification of Walker’s specimen b, without locality data, as primary type. Penny (1977: 23) listing as a synonym of Leucochrysa internata (Walker), without comment.

Leucochrysa (Leucochrysa) ampla (Walker). Brooks and Barnard (1990: 276) first reference to generic, subgeneric combinations, species list; Freitas and Penny (2001: 280, 351, fig. 40) brief redescription, collection records, figures; Oswald (2007) catalog listing. Syn. n.

Leucochrysa vegana Navás [1917] ( 1917: 278) (not Nodita vegana Navás, 1925) original description: "Colombia: La Vega, Juni 1915 (Coll. Br. Apolinar Maria)". Banks (1944: 30) synonymy with Leucochrysa varia . Penny (1977: 23) listing as a synonym; Brooks and Barnard (1990: 277) listing as a synonym; Oswald (2007) catalog listing as a synonym. Nom. d.

Leucochrysa phaeocephala Navás [1929] ( 1929b: 21) original description: "America: "Niederl. Guayan. Obere Commaryne, 28. 11. 1908". Mus. Hamburg". Banks (1944: 30) synonymy with Leucochrysa varia . Penny (1977: 23) listing as a synonym; Brooks and Barnard (1990: 277) listing as a synonym; Oswald (2007) catalog listing as a synonym.

Leucochrysa (Leucochrysa) phaeocephala Navás. Valid status reinstated. See below.

Leucochrysa walkerina Navás [1913] ( 1913: 102) original description: “Brazil”. Penny (1977: 23), as " Leucochrysa walkerina Navás (1917) ", listing as a synonym of Leucochrysa internata , without comment.

Leucochrysa (Leucochrysa) walkerina Navás. Brooks and Barnard (1990: 277) subgeneric determination, species list; Freitas and Penny (2001: 283, 355, fig. 44) brief redescription, collection records, figures -- based on misidentified specimens in the FCAV-UNESP; Oswald (2007) catalog listing. Syn. n.

Known geographical distribution.

Our findings [based on confirmed published records and specimens examined] indicate that Leucochrysa (Leucochrysa) varia occurs from the western, lowland regions of the Amazonian drainage basin of Brazil, Ecuador and Peru, throughout much of forested Brazil (coastal and inland), and in northeastern, mid-elevation regions of Argentina. The specific areas that we have confirmed include: Argentina: Province of Salta. Brazil: States of Bahia, Distrito Federal, Mato Grosso, Minas Gerais, Pará, Rio de Janeiro, Rondônia,São Paulo. Ecuador: Provinces of Napo, Orellana. Peru: District ofMadre de Dios.Confirmed published records include: Freitas and Penny (2001: 280, as Leucochrysa (Leucochrysa) ampla ), and Mantoanelli et al. (2006: 8).

Unconfirmed, published records from South America include --- Brazil: EspíritoSanto ( Navás 1922b: 89), Mato Grosso ( Navás 1932: 57); Pará ( Navás 1912-1913: 314), Rio de Janeiro ( Navás 1922a: 392; Navás 1926: 12; Navás 1929a: 862), Paraguay: San Bernardino [Cordillera] ( Navás 1913: 157), and Bolivia: Buenavista [Santa Cruz] ( Navás 1928b: 111). The Brazilian records of Leucochrysa (Leucochrysa) varia and Leucochrysa (Leucochrysa) walkerina by Freitas and Penny (2001: 282) were based on misidentified specimens; thus, only their records for Leucochrysa (Leucochrysa) ampla [= Leucochrysa (Leucochrysa) varia ] are included here.

Leucochrysa (Leucochrysa) varia was reported from British Guiana, Suriname, Central America, and Mexico; however we have not confirmed any of these records. Banks (1945: 168) questioned the Navás (1928a: 131) record for Central America (Guatemala). Based on our study here, we also question this record. We confirmed that the Banks (1944: 31) record for British Guiana was in error; a specimen in the AMNH with the data he reported is probably an undescribed species; its abdomen is missing. The specimen(s) that he identified as Leucochrysa (Leucochrysa) varia from Suriname ( Banks 1944: 31) were not found in the MCZ (P. Perkins, personal communication), nor in the AMNH. This record probably applies to Leucochrysa (Leucochrysa) pretiosa or another, undescribed species. In summary, we have not seen specimens of Leucochrysa (Leucochrysa) varia from northern South America, the Caribbean region, Central America, or Mexico.

Type specimens and rationale for taxonomic changes.

Chrysopa varia . Holotype, by original designation, MCZ, male (examined).

The type is in fairly good condition. Its primary label is hand-written, apparently in Schneider’s hand; it reads: "varia nov. sp. / Schneider / Brasilia". The exact collection site is unknown. In addition to Fig. 1 here, images of the external and male features are in the MCZ Type Database (website: http://insects.oeb.harvard.edu/MCZ/index.htm).

Chrysopa internata . Holotype by monotypy, BMNH, male (examined).

Walker originally referenced two specimens under the name Chrysopa internata ( “a” from Brazil and “b” without locality data); he also stated that Chrysopa internata had two varieties. Because one of the specimens must represent Walker’s lettered variety ("var. β”), that specimen must be excluded from the type series of Chrysopa internata under Art. 72.4.1 (see Oswald 2007). Kimmins (1940: 444) identified the specimen without locality data (specimen “b”) as the primary Chrysopa internata type (a lectotype). Because the actual type series of Chrysopa internata consists of only a single specimen, his lectotype designation was unnecessary; we recognize Kimmins’ action as identification of the holotype.

The excluded "var. β” specimen (specimen “a” from Brazil), was subsequently designated the type of Leucochrysa walkerina Navás (see below).

The Chrysopa internata holotype is in fairly good condition; the head is separated from the body and the dissected abdomen is in glycerin, in a vial attached to the pinned specimen. The labels are hand-written and printed (Fig. 2B). The specimen carries no locality data; its collection site is unknown. All of its features, including the male terminalia, correspond to those of the Leucochrysa (Leucochrysa) varia type (see Figs 2-3).

Chrysopa ampla . Holotype by monotypy, BMNH, female (examined).

Walker originally referenced two specimens ( “a” and “b”) under the name Chrysopa ampla ; he also stated that Chrysopa ampla had two varieties. Because one of the specimens must represent Walker’s lettered variety ("var. β”), that specimen must be excluded from the type series of Chrysopa ampla under Art. 72.4.1 (see Oswald 2007). Thus, the actual type series of Chrysopa ampla consists of only a single specimen, the one that was listed by Walker as "var. α” and that carries no locality data. Therefore, this specimen constitutes the holotype by monotypy. This designation is consistent, nomenclaturally, with the action by Kimmins (1940: 444), who considered the specimen as the name-bearing type when he designated it as the “lectotype”.

The excluded "var. β” specimen was reported from “Georgia”. Tauber (2004: 1132) identified it as Leucochrysa (Leucochrysa) insularis Walker, and it bears her label with that name.

The Chrysopa ampla holotype is in fairly good condition; the dissected abdomen is in glycerin, in a vial attached to the pinned specimen. The labels are hand-written (not by Navás) and printed (Fig. 4A). The specimen carries no locality data; its collection site is unknown. All of the features (external and genitalic) of this type correspond to those of female Leucochrysa (Leucochrysa) varia (see Figs 4-5).

Leucochrysa vegana . Type(s) by original designation, probably missing.

Navás (1917: 278), in his original description, did not indicate the depository or sex of the type; he reported that it was from the "Coll. Br. [Brother] Apolinar Maria [ María]” (not examined). It was not found in the Navás collection ( Monserrat 1985: 240), the BMNH, the MNHN, or the MCZ. And despite searches by colleagues (see acknowledgements) in Colombia where Br. Apolinar María lived, it was not found. Apparently, the Br. Apolinar María collection was housed in the Museo de la Universidad de la Salle and was destroyed during a political upheaval in 1948.

Navás mentioned the similarity between Leucochrysa vegana and Leucochrysa varia ("variae Schn."), but he did not point out why he considered that Leucochrysa vegana was different. Most of the features that he described for Leucochrysa vegana are found on Leucochrysa (Leucochrysa) varia specimens, including the markings at the base of the Leucochrysa vegana forewing that he illustrated. Thus, although it is unlikely that Banks actually saw the Leucochrysa vegana type, his synonymy (based on Navás’ description) made sense at the time.

The type locality of Leucochrysa vegana (La Vega, which is in the Cordillera Oriental of Colombia), is considerably north of the currently known northern limit of Leucochrysa (Leucochrysa) varia [the western regions of the Amazon drainage basin of Ecuador and Peru (e.g., the Yasuní Reserve in the Province of Napo, Ecuador and the Tambopata district of Peru)], but well within the ranges of other, undescribed Leucochrysa (Leucochrysa) varia -like species. At this time, we suspect that Leucochrysa vegana is not synonymous with Leucochrysa (Leucochrysa) varia . However, the region remains very poorly collected, so it is possible that our suspicion is in error. Thus, while we await the collection of Leucochrysa (Leucochrysa) varia from La Vega or nearby, to confirm Banks’ synonymy, we consider the species name to be a nomen dubium.

Leucochrysa phaeocephala . Type(s), by original designation, Hamburg, probably destroyed during W.W. II, sex unknown (not examined).

Banks’ (1944: 30) synonymy was made on the basis of the description; it does not appear that he saw the type. Navás (1929b: 21) reported that the specimen was collected in Dutch Guiana in 1908 (now, Suriname). Because we have found no records of Leucochrysa (Leucochrysa) varia from the northern regions of South America, we reverse Bank’s synonymy; we will deal with the species in a later publication.

Leucochrysa walkerina . Holotype by monotypy, BMNH, female (examined).

Navás (1913: 102) considered that Walker’s internata variety β represented a distinct species; he explicitly applied the species name “walkerina” in honor of Walker. Kimmins (1940: 444) designated Walker’s specimen “a” of internata (labeled from Brazil) as the type of Chrysopa internata variety β; thus it became the holotype of Leucochrysa walkerina (see Oswald 2007).

The walkerina type is slightly teneral, but in good condition; the cleared abdomen is in glycerine, in a vial attached to the specimen. The pin carries four labels below the specimen (including a locality label, “Brazil”) (Fig. 6A); it also has a “Paralectotype” label above the specimen (Fig. 6B). The external characteristics (Fig. 6) are consistent with those of Leucochrysa (Leucochrysa) varia , and the elongate, coiled, spermathecal duct is that of Leucochrysa (Leucochrysa) varia (Fig. 7).

Diagnosis.

As the name varia implies, adults of Leucochrysa (Leucochrysa) varia exhibit a wide range of color variation; there are black and red morphs, with and without coloration on the mesoscutellum (See Fig. 8 here, Fig. 7 in Mantoanelli et al. 2006). In addition, preserved specimens of varia-like species tend to loose their natural coloration very quickly; old specimens are especially discolored. Thus, it is important to emphasize that accurate identification of the varia-like species can only be made by careful examination of the male or female genital characters.

Externally, Leucochrysa (Leucochrysa) varia and Leucochrysa (Leucochrysa) pretiosa adults usually (but not always) can be separated by differences in the darkness of certain veins and the degree of suffusion around the veins. Both species have black to dark brown suffusion surrounding the second m-cu crossvein, but the marking generally is less prominent in Leucochrysa (Leucochrysa) varia than in Leucochrysa (Leucochrysa) pretiosa . Moreover, in both species the distal two to four Psm–Psc crossveins and at least the three basal outer gradates are darkened and the membranes surrounding these veins are shaded to some degree. In Leucochrysa (Leucochrysa) varia , the degree of darkening and suffusion around these veins is generally uniform, whereas in Leucochrysa (Leucochrysa) pretiosa , the distal Psm-Psc crossvein is usually darker and more prominent than the other Psm-Psc crossveins or the outer gradates.

In Leucochrysa (Leucochrysa) varia males, the sclerotized mediuncal plate is elongate; its rods are narrow and parallel; its membranous connection to the gonarcal bridge does not extend laterally beyond the gonocornua and is soft [not broad, leathery and stiff, as in Leucochrysa (Leucochrysa) pretiosa ]. The Leucochrysa (Leucochrysa) varia female is distinguished by a very long, strongly coiled spermathecal duct and a spermatheca that is scoop-shaped distally and has a convoluted, tubular, basal section leading to a broad, fluted bursal duct.

Redescription.

Head (Fig. 8): 1.8-1.9 mm wide (including eyes). Frons, clypeus cream to red, with anterior clypeal margin dark red; gena red to reddish brown; maxillary, labial palp yellowish to cream-colored. Vertex with central area raised, yellowish to cream-colored, with prominent, dark red-wine-colored, V-shaped mark along anterior margin; lateral margin, midline sometimes also with red-wine-colored marks. Antenna: dorsum of scape lightly to darkly tinged with red-wine color; pedicel dark red to ivory-colored, with inner margin darkened; flagellum cream-colored, with amber bristles; inner margin of basal ca 3 flagellomeres tinged with red to black; antennal fossa marked with red laterally.

Thorax (Fig. 8): Cervix lightly tinged with red laterally. Pronotum 1.2-1.5 mm long, 1.2-1.5 mm wide, yellowish to greenish, unmarked except for small posteromesal red to black mark. Mesonotum, metanotum mostly dark, variable, with four types of color morphs (black entire, black open, red entire, red open) [see Fig. 8 and Mantoanelli et al. 2006].

Wings (Fig. 9 A–B): Forewing 17.3-20.5 mm long, 6.5-7.8 mm wide (at widest point); ratio of length: maximum width = 2.6-2.7:1. Costal area moderately broad; tallest costal cell (#9-10) 1.5-1.9 mm tall, 2.6-3.3 times width, 0.3 times width of wing (midwing). First intramedian (im1) cell quadrangular, width (anterior margin) 1.4-1.6 times width (anterior margin) of third median cell (m3), 2.3-3.2 times length of posterior margin of m3, length of basal vein (= ma, median arculus) 1.0-1.1 times greater than length of distal vein. First radial crossvein distal to origin of radial sector (Rs); radial area (between R and Rs) with single row of 16-18 closed cells; tallest cell (#6-8) 2.16-2.58 times taller than wide. No crassate veins; 5-7 b cells (= cells beneath Rs, not including an inner gradate vein). Two series of gradate veins; 6-11 inner gradates, 7-9 outer gradates. Height of fourth gradate cell 3.6-5.4 times width. Eight to nine b’ cells (cells beneath Pseudomedia after im2). Three intracubital cells (two closed). Membrane mostly clear except basal area with small, reddish brown patch, stigma opaque to light brown, with large very dark brown mark basally, small, light brown clouding around second m-cu crossvein and around distal Psm–Psc crossveins, sometimes with clouding around distal leg of im1 and around crossveins between distal b’ cells. Veins mostly green, except anterior tips of most costal crossveins, base of Radius, bases of ca three to five radial crossveins, distal three to four Psm–Psc crossveins, outer gradates, and forks of posterior marginal veins darkened to black; inner gradates, posterior veins of distal three to four b’ cells slightly darkened. Hindwing 16.2-18.3 mm long, 5.3-6.0 mm wide. Two series of gradate veins; 6-10 inner, 6-8 outer; 14-17 radial cells (counted from origin of R, not false origin). Four to six b cells (including small b1 cell); six to eight b’ cells beyond im2; two intracubital cells (one closed). Membrane clear; stigma with pronounced dark brown mark basally. Veins mostly light green; middle costal crossveins, outer gradates, bases of marginal forks, black.

Abdomen (Figs 1, 10-14): Sternites, tergites with long, slender setae throughout, microsetae moderately dense; pleural region with setae small, very sparse, microsetae very small. Rim around each sternite heavily sclerotized, especially anteriorly, fading posteriorly. Tergites narrow, roughly rectangular, with rounded margins. Spiracles oval externally; atria not enlarged. Sternites S2-3 longer than tall; S5-7 more square-shaped; distal segments (beyond A4) expanded, height of pleural region greater than height of sternites. Coloration: mostly green, with yellow mesally. Tergites T5, T6 with large black spots, bordered by red; callus cerci white; setae, trichobothria golden.

Male: S6 height and length approximately equal, S7 height ca 1.1-1.2 times length (lateral view); S4-S8 with dense microtholi, S3 with microtholi laterally, sometimes also across entire posterior region, absent or sparse anteriorly, mesally; S1-2, S9 without microtholi. Callus cerci round to slightly oval (ca 1.1 –1.2× taller than wide), diameter ca 0.16-0.28 mm, with ca 30-35 trichobothria of variable length. T9+ectoproct rounded posterodorsally, truncate to rounded distally, broadly fused mesally, midline with small distal cleft, with long setae; ventral section tapering, rounded proximally, extending above S8+9 only to suture between S8 and S9; dorsal apodeme substantial, but not thick, straight basally, forking midway on anterior margin of callus cerci; dorsal branch extending around dorsal margin of callus cerci to midway on posterior margin, ventral branch curving ventrally well below callus cerci, then bending posteriorly, extending along ventral margin of ectoproct. S8+9 fused, with trace of suture dorsally, with clear intersegmental demarcation throughout; S8 tall (1.5 –2.0× taller than long), ca one-half (0.44 –0.47×) length of S8+9 along ventral margin; S8+9 (lateral view) with proximal margin slightly convex, dorsal surface of S8 rounded, of S9 curved distally, steeply sloped at terminus; terminus without knob or gonocristae; membranous region above terminus of S9 with pair of large, eversible, lateral pouches. Gonarcus well sclerotized, widely arcuate (total span, 0.67-0.84 mm); bridge broad (0.38-0.62 mm long), dorsoventrally flattened, gently curved throughout; lateral apodeme rounded dorsally, more acute ventrally (0.21-0.24 mm wide; 0.29-0.33 mm tall); gonocornua extending forward from anterior edge of gonarcal bridge, basally stout, with ventral, straight projection, tapering to narrow, rounded apex (length, 0.09-0.15 mm); distance between inner bases of gonocornua 0.15-0.25 mm, distance between tips 0.20-0.26 mm. Mediuncus located beneath, and well separated from gonarcus, with mesal, recurved beak, lateral, well-sclerotized, stiff membranous, curved arms; mediuncus attached to gonarcal bridge via robust membrane that extends along width of gonarcal bridge between outside margins of gonocornua, folds beneath gonarcal bridge and beneath mediuncal beak to form deep gonosaccus; gonosaccus with two fields of five to seven short gonosetae on chalazae. Hypandrium internum: arm 0.25-0.37 mm long, distal span between arms 0.24-0.32 mm.

Female. Height of S6 ca 0.75 times length, S7 height ca 0.60 times length. Callus cerci round, diameter 0.17-18 mm, with 30-35 trichobothria. T8 roughly quadrate (lateral view) with rounded corners, similar in depth to T6. T9+ectoproct elongate, slanting anteriorly; ventral margin slightly convex, extending slightly below level of gonapophyses laterales. Dorsal margin of S7 with slight taper basally, becoming more pronounced distally; terminus unmodified, with terminal (posteroventral) setae slightly more dense than in other areas. Gonapophysis lateralis rounded to slightly acute dorsally, rounded distally, ventrally, ca 0.53-0.60 height of T9+ectoproct; inner membranous surface not expandable, with ca two vertical rows of short setae. Colleterial complex consisting of membranous gland connected to colleterial reservoir via broad duct, and elongate ribbon-like accessory gland; colleterial gland elongate, delicate, transparent; colleterial reservoir smaller, delicate, transparent, tapering to narrow, granulose, spiny duct; accessory gland narrow, elongate, forked distally, with spiny surface; accessory gland and colleterial duct connected to lightly sclerotized, widened, flattened platform extending from below transverse sclerite; transverse sclerite curved, lightly sclerotized, slender throughout, with long teeth (setae?). Spermatheca with initial (posterior) section scoop-shaped, broad, thick, tapering slightly at base (ca 0.25 mm wide along distal margin x ca 0.20 mm height from tip of distal margin to base of scoop), with elongate, broad, smooth, convoluted tube extending down one side, looping in U-shaped turn, then twisting through several loops before joining bursal duct [tube ca 1.1 mm in total length, ca 0.05 mm in width throughout]; spermathecal invagination not specifically identified. Spermathecal duct extremely long, well sclerotized throughout, densely, tightly coiled, arising from side of scoop-shaped section of spermatheca; coiled length ca 3-5 mm, including membranous, brushy, less coiled distal section, uncoiled length much greater. Bursal duct extending from tip of tubular spermathecal velum, basal section membranous, broad, curved, fluted; surface with striated folds, lateral margins of major folds heavily granulose. Bursa copulatrix small, saccular, extended over spermatheca, slightly into section of S7; ventral surface with small striated folds; dorsal surface smooth; pair of clear, elongate, tubular bursal glands attached dorsally to base of bursa via clear, pipe-fitting-like bases; bursal glands very long, unbranched; surface lightly granulose. Subgenitale with smooth (unfolded), rounded surface, broad, rectangular, robust, bilobed projection extending distally at ca 90° angle to subgenitale surface, lobes large, with minute setae on surface, region between lobes extending distally as smaller, acute lobe; base of bilobed projection with dense transverse folding, with sclerotized, knob-like mesal lobe projecting from pair of scalloped, sclerotized arms.

Larva.

All instars described ( Mantoanelli et al. 2006).

Biology.

Developmental and survival rates of immature stages under five constant temperatures and larval trash-carrying behavior studied by Mantoanelli et al. (2006) and Mantoanelli and Albuquerque (2007).

Variation.

The coloration of the head, mesonotum and metanotum is the most obvious variation expressed by Leucochrysa (Leucochrysa) varia [see above]. However, color is not the only feature that varies among Leucochrysa (Leucochrysa) varia specimens. For example, the expansion of abdominal segments 4-9 and the degree of sclerotization of the integument varies greatly among both male and female specimens (see Figs 10, 12). Differences in sclerotization are particularly noticeable in the ventral apodemes of the male T9+ectoproct (Fig. 10). We suspect that these features (abdominal expansion and integumental sclerotization) are at least partially a function of age and developmental or reproductive maturation.

Two features, specific to males, also show considerable variation. First, the breadth of the elongate mediuncal “rods” varies. For example, the Chrysopa internata type [= Leucochrysa (Leucochrysa) varia ] has very broad mediuncal rods (Fig. 3), whereas the Chrysopa varia type [= Leucochrysa (Leucochrysa) varia ] has considerably narrower ones (Fig. 1). Other specimens express a range in size between these two specimens (Fig. 11). Finally, the pattern of microtholi on the third abdominal sternite varies among males. In one specimen from Ecuador, the microtholi were restricted to a relatively small lateral patch; they were absent from the mesal region of the sternite. In another specimen, this one from Brazil, microtholi were present along the entire lateral region of the sternite and also in a strip along the posterior margin. Whether this variation has a geographical component is unknown.

Material examined

(in addition to the types listed above). Argentina. Salta: Cafayate, ii–iv.1968, Hayward (2♂, 3♀, USNM). Brazil. Mato Grosso: Itiquira [17°12'S, 54°09'W], 522 m, 17.i.1996, 2♂, V. Cruz; idem, 26.viii.1996, 2♂, V. Cruz; idem, 5.x.1996, 1♂, V. Cruz; 18.x.1996, 1♂, 3♀, V. Cruz; idem, 31.xii.1996, 2♂, V. Cruz; idem, 13.i.1997, 1♀, S. Freitas (all FCAV-UNESP). Minas Gerais: Caratinga, 19.ix.2011, S. Freitas & F. Sosa (2♂, 2♀, UCOB). Pará: Rio Xingu Camp, ca 60 km S. Altamira [3°39'S, 52°22'W], 2-8.x.1986, malaise trap, night collection, P. Spangler & O. Flint (1♀, USNM). Bahia: Camacan, Reserva Serra Bonita, Fazenda Paris, 200 m, 6.x.2005, G. S. Albuquerque, M. J. Tauber, C. A. Tauber Expedition, Oct. 2005 (15♂, 11♀, UCB). Distrito Federal: CENARGEN Farm nr. Núcleo Bandeirante, 20.iii.1999, M. J. & C. A. Tauber (2♂, 3♀, UCB); idem, 9.v.2002 (2♂, UCB). Rio de Janeiro: Parque Estadual do Desengano, Santa Maria Madalena, Terras Frias, 30.iii.1999, M. J., C. A. & P. J. Tauber, G. S. Albuquerque (3♂, 2♀, UCB); idem, 15.v.2002, M. J., C. A., P. J. & A. J. Tauber, G. S. Albuquerque, E. A. Silva (9♂, 6♀, UCB); idem, 28.x.2003, G. S. Albuquerque, M. J. Tauber, C. A. Tauber Expedition, Oct-Nov 2003 (1♀, UCB); idem, 22.iv.2004, G. S. Albuquerque, M. J. Tauber, C. A. Tauber Expedition, April 2004 (16♂, 10♀, UCB); Parque Estadual do Desengano, Campos dos Goytacazes, Babilônia, 26.iii.1999, M. J., C. A. & P. J. Tauber, G. S. Albuquerque (9♂, 6♀, UCB); idem, 21 & 27.iii.2001, M. J., C. A. & P. J. Tauber, G. S. Albuquerque (5♂, 3♀, UCB); idem, 20.v.2002, M. J., C. A., P. J. & A. J. Tauber, G. S. Albuquerque, E. A. Silva (6♂, UCB); Parque Estadual do Desengano, Campos dos Goytacazes, Santo Antônio do Imbé, 24 & 31.iii.1999, M. J., C. A. & P. J. Tauber, G. S. Albuquerque (8♂, 2♀, UCB); idem, 27.iii.2001, M. J., C. A. & P. J. Tauber, G. S. Albuquerque (1♂, 1♀, UCB); idem, 1 & 5.v.2003, G. S. Albuquerque, M. J. Tauber, C. A. Tauber Expedition, April-May 2003 (2♂, UCB); Parque Estadual do Desengano, Campos dos Goytacazes, Sossego, 25.iii.1999, M. J., C. A. & P. J. Tauber, G. S. Albuquerque (1♂, 1♀, UCB); Parque Estadual do Desengano, Campos dos Goytacazes, Fazenda Pedrinho, 3.v.2003, G. S. Albuquerque, M. J. Tauber, C. A. Tauber Expedition, Apr.-May 2003 (1♂, UCB); near Parque Estadual do Desengano, Campos dos Goytacazes, Fazenda Boa Vista, M. J., C. A. & P. J. Tauber, G. S. Albuquerque (8♂, 6♀, 1 teneral, UCB); Campos dos Goytacazes, Dist. de Morangaba, Fazenda São Julião, 18.x.2005, M. J. & C. A. Tauber, G. S. Albuquerque (2♂, 1♀, UCB); Campos dos Goytacazes, Conceição de Macabu, Santo Agostinho, 21.v.2002, M. J., C. A., P. J. & A. J. Tauber, G. S. Albuquerque, E. A. Silva (1♂, UCB); Campos dos Goytacazes, Conceição de Macabu, Fazenda Carrapeta, 29. iv– 6.v.2003, G. S. Albuquerque, M. J. Tauber, C. A. Tauber Expedition, Apr.-May 2003 (1♂, UCB); idem, 23.iv.2004, G. S. Albuquerque, M. J. Tauber, C. A. Tauber Expedition, April 2004 (11♂, 4♀, UCB); Ilha Grande, 01.vi.2002, G. S. Albuquerque (1♂, UCB). São Paulo: São Vicente [23°57'S, 46°23'W], 916 m, 21.x.2009, S. Freitas (1♀, FCAV-UNESP). Rondônia: 62 km s. Ariquemes, 8-20.xi.1994, W. J. Hanson (1♀, USU); idem, 22-31.x.1997 (1♀, USU). Ecuador. Orellana [Napo*]: Yasuní Res. Sta. [0°38'S, 6°36'W [Sic!, probably 76°36'], 250 m, 19-30.x.1996, W. J. Hanson (1♀, USU); idem, 18-30.x.1998, W. J. Hanson (1♀, USU); Misahualli nr. Tena, 3-8.x.1999, S. R. Keller (1♂, USU); idem, 26.viii-2.ix.2000, S. & P. Keller (1♀, USU); idem, 6-19.x.2001, C. Branimer (1♀, USU); Yasuni Biol. Res. Sta. [ca 0°40'S, 76°24'W] 1-7.xi.2002, E. M. Fisher (1♂, 1♀, CAS); Coca [= Puerto Francisco de Orellana] [0°03'S, 79°40'W], v.1965, L. C. Peña (1♂, CAS). Peru. Madre de Dios: Tambopata, 15 km NE Pto. Maldonado, 200 m, 25.vi.1989, R. A. Leschen #177, ex malaise trap (1♀, SEM); Río Tambopata Res., 30 km (air) sw Pto. Maldonado [12°50'S, 69°20'W], 290 m, 08.xi.1984, T. L. Erwin, canopy fogging 04/01 (1♀ alcohol, USNM); idem, 02.v.1984, T. L. Erwin et al., canopy fogging project 02/03 (1♀ alcohol, USNM); Pakitza, Zone 02 [11°55'48"S, 71°35'18"W], 9.ix.1988, insecticidal fog, canopy/pouteria, BIOLAT 02180413, T. L. Erwin (1♀ alcohol, USNM).

[Note: * The Yasuní Research Station is located in Orellana (a province that was separated from Napo province in 1998). Ecuador’s entire Yasuní Biosphere Reserve (established by UNESCO in 1989) encompasses a large area between the Napo River in the north and west and the Curaray River in the south and east; in terms of biological diversity, it is an extremely rich area.]