Corresponding author: Catherine A. Tauber (
Academic editor: A. Contreras-Ramos
Here,
As a result of this study, three generalizations appear: (1) The genital morphology of both males and females of
Tauber CA, Sosa F, Albuquerque GS, Tauber MJ (2017) Revision of the Neotropical green lacewing genus
The genus
Despite the above, the systematics of the genus has been confusing, and attempts to identify specimens to species can be exasperating. A number of factors underlie the difficulties. (1) The original seven
Here, we present an historical account of the genus, and for each named species, we provide images and/or information on the type as well as our evaluation of the species’ taxonomic status. For some species, we were able to borrow or photograph the types and the information is detailed; for others it is less detailed. In addition, for those species that we can identify confidently, and for which we have sufficient material, we offer notes and images of recently collected (non-type) material for use in identifications. To facilitate comparisons, we redescribed all previously described species in the same format as the new species descriptions. This includes species recently described by
We based this review on four resources: published literature, type specimens, unpublished notes compiled by Phillip A. Adams, and non-type specimens from numerous museums.
Most specimens were studied by C. A. Tauber; those examined only by F. Sosa are noted as (FS). In some cases, specimens were also examined or photographed by G. S. Albuquerque (GSA) and/or M. J. Tauber.
We examined (or refer to) specimens from the following museums:
*
Phillip A. Adams’ notes consist of sketches and brief descriptive statements regarding the specimens (mostly types) that he examined during his trips to museums in Europe, South America, and North America. These were very valuable in identifying the location and condition of types, and in providing his thoughts on the similarities and differences between selected species. Upon Adams’ death, his notes became the property of the California Academy of Sciences; sections were made available to CAT and MJT by N. D. Penny (
At the time during which most of the notes were written, Adams considered the species that now are included in
The
Banks, too, often used more than one specimen in his descriptions, but he usually labeled specific specimens as holotypes, or otherwise indicated which specimen he considered to be the primary type. Thus, if we were reasonably certain that his labels indicated his intent to identify a holotype, we considered it as such. If the labeling left us uncertain, we designated a lectotype.
We made selected measurements on the wing and head. Our techniques are illustrated in Figs
In addition to the abbreviations for the museums above, we used several other abbreviations, as follows. For morphological structures: Wing veins (forward to hind) – C = Costa,
To categorize literature: biol = biological information; catalog = species included in a catalog; desc = description of adult; dist = distribution records; larval desc = description of larva; list = species name included in a list of species; redesc = redescription of adult; tax = taxonomic information; type(s) = information on type specimen(s).
Measurements of head, dorsal.
Wing structures and measurements; upper, forewing; lower, hindwing. Veins (black lettering):
Measurements of male abdomen; upper, lateral; lower, dorsal.
Originally
Later,
Males have distinctive abdominal and genitalic features that differentiate them from males of other chrysopid genera (see below). In contrast, females share a very simple set of reproductive structures that occurs repeatedly among chrysopid genera [
Externally,
From the beginning of the study, we noted that specimens fell into two distinct and non-overlapping groups. Now at the conclusion of the study, we recognize: (1) seven species in southern South America (Argentina: six species; southern Brazil: one species) and (2) 18 species in more northern regions of South America (Peru, Brazil, Bolivia, Ecuador, Colombia, Venezuela: 15 species), and extending into Central America (Costa Rica: 2 species) and southern Mexico (Chiapas: 1 species). The species from the southern region were by far the more problematic, and now that our study is complete, we are unsatisfied with our understanding of this group of species. Clearly, additional specimens and more intense examination of the types from this part of the continent are necessary for a satisfactory classification. In contrast, although the availability of specimens from the more northern regions was relatively scanty, the species were more readily distinguishable on the basis of external characteristics.
For convenience, we divided our treatment of the genus along geographic lines. We begin with the northern group and then deal with the species from Argentina and Brazil separately in the latter part of the revision. We do not expect that the relatively distinct geographic separation that we found among the
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1 | Frons with dark (brown, red or black) mesal spot (Figs |
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1’ | Frons without mesal spot |
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2(1) | Antenna with flagellum black at least basally or with base black (Figs |
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2’ | Antenna with flagellum cream-colored throughout |
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3(2) | Frons with prominent mesal protuberance below frontal spot (at least in male); forewing ~ 10 mm long, with ~ 3–4 gradate veins per series (Fig. |
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3’ | Frons without protuberance; forewing ~13–14 mm long, with ~ 5–7 gradate veins per series (Figs |
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4(2) | Forewing long: ~17–18 mm long, with 8–11 gradate veins per series (Fig. |
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4’ | Forewing shorter: ~11–15.5 mm long, with 4–8 gradate veins per series; U-shaped mark on vertex separated mesally, but arms visible |
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5(4) | Hindwing with gradate veins dark (brown or black) (Fig. |
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5’ | Hindwing with gradate veins pale (green) (Figs |
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6(5) | Dorsal surface of scape with brown distal stripe (Fig. |
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6’ | Dorsal surface of scape tinged with red, but without distinct stripe (Fig. |
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7(1) | Flagellum black or partially black |
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7’ | Flagellum cream-colored, unmarked |
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8(7) | Scape with dorsal surface unmarked, ventral surface with large black mark distally; foretibia without black spot (Fig. |
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8’ | Scape with dorsal surface almost entirely black, ventral surface unmarked (Fig. |
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9(7) | Prothorax with lateral stripe red, extending only along anterior half of segment (Fig. |
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9’ | Prothorax with lateral stripe red to brown, extending full length of segment |
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10(9) | Dorsum of scape with distinct reddish, brown, or black stripe or mark |
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10’ | Dorsum of scape cream-colored or slightly tinged with red, without marks |
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11(10) | Mark on vertex reduced to pair of small or diffuse red marks |
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11’ | Mark on vertex distinct, robust, with crescent-shaped arms at least anteriorly, either diffuse or heavy posteriorly |
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12(11) | Forewing venation (males) robust, green except gradate veins dark, with at least some brown suffusion on surrounding membrane (Fig. |
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12’ | Forewing with venation thin, delicate; longitudinal veins mostly green; most crossveins, including gradates reddish, without suffusion (Fig. |
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13(12) | Gena and palpi light or with no marks; region between lateral margin of eyes and vertex marked with red (Fig. |
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13’ | Gena and palpi heavily marked with black or reddish black; region between lateral margin of eyes and vertex cream-colored, unmarked (Figs |
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14(11) | Forewing with crossveins mostly brown or black, some with suffusion on surrounding membrane; dorsum of head with prominent reddish brown mark between eyes and margin of raised vertex |
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14’ | Forewing with crossveins mostly reddish, none with suffusion; area between eyes and lateral margin of vertex without prominent mark |
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15 (14) | Head cream-colored to yellow; gena, lateral margin of clypeus with black stripe; antenna with dorsal stripe extending full length of scape and onto antennal fossa (Fig. |
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15’ | Head orange, gena, clypeus pale, without black stripe; scape with dorsal stripe extending ~3/4th distance from apex to base, not on antennal fossa (Fig. |
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16(14) | Dorsum of scape with dark brown distal stripe; U-shaped mark on vertex brown (Figs |
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16’ | Dorsum of scape with red or reddish brown distal stripe (Fig. |
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17(10) | Gena unmarked (Fig. |
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17’ | Gena with black, brown or reddish mark; some veins of forewing with brown marks (small or extensive) |
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18(17) | Vertex with U-shaped mark prominent, thick, with mesal arm extending laterally to eyes (Fig. |
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18’ | Vertex with U-shaped mark reduced to a pair of small, red spots on anterolateral margin, far removed from eye (Fig. |
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(Figs
The species is named for Phillip A. Adams, an intense and talented contributor to systematics and evolutionary studies of
Head cream-colored with dark to light reddish black markings; vertex with anterior marking dark, prominent, separated or contiguous mesally, with diffuse coloration, not extending anteriorly toward antennal base; lateral marks on vertex diffuse, extending from anterior dark mark to base of vertex; dorsal fossa with small red stripe mesally (usually hidden by scapes); posterior parts of vertex unmarked; frons swollen distally (male), smooth distally (female), with large, prominent mesal mark, otherwise whitish (male), cream-colored with inverted V-shaped mark extending from mesal spot (female); gena, lateral margin of clypeus with black stripe throughout; tentorial pits amber-colored. Antenna: scape cream-colored, dorsum with diffuse, light reddish marks; pedicel amber or with amber-colored ring, flagellum cream-colored; maxillary palp with basal two segments pale, three distal segments black, articulations pale; labial palp with basal segment pale, distal two segments marked with black, articulation pale. Corner of cervix with large, dark brown to black mark.
Thorax with distinct longitudinal cream-colored stripe mesally. Prothorax with pair of broad, dark reddish brown stripes laterally; small transverse furrow in posterior region of segment, extending to pair of pale spots within sublateral stripe; setae mostly elongate, pale. Measurements: head width: 1.3 mm; ratio head width : eye width: 2.7–2.8 : 1; prothorax width: 0.6–0.8 mm, length: 0.9–1.0 mm.
Forewing somewhat broad, apex broadly subacute to rounded, membrane clear, hyaline, with light to very light suffusion of brown surrounding gradate veins; stigma very slightly opaque; longitudinal veins green; transverse veins brown or marked with brown; forewing with veins robust to slender, not crassate;
Male: Abdomen with large spiracles, elongate segments (e.g., A7: spiracle diameter ~0.14-0.18x length of sternite); T9+
PERU: Department of Cuzco.
(in addition to the holotype; all paratypes). Peru; Cuzco, Machu Picchu museum, 1,385 m., 11-14/VIII/1971 at light, C. & M. Vardy. B.M. 1971-533 (M,
This species is distinguished by pronounced, dark brown body markings; dark brown or black marking on frontal base of foretibia; tall wings with rounded apices, forewing with very dark venation, hindwing venation less dark; stark, shiny, dark brown to black markings on dorsum of scape and vertex; moderately enlarged abdominal spiracles; distal setae of S8+9 larger than those on ventral or proximal region, but not exceptionally robust; those on narrow strip of distolateral margin flanged. Forewing 13.0 mm long, 5.5 mm wide; hindwing 12.1 mm long, 4.4 mm wide.
Head cream-colored, with vertex smooth, shiny, with inverted U-shaped marking prominent, dark brown, broad, separated anteromesally, not extending anteriorly to area between scapes; dorsal antennal fossa pale; area between eyes and posterior half of vertex cream-colored, unmarked; frons, clypeus cream-colored, without markings; gena with large, brown mark from eye onto clypeus; tentorial pits amber. Antenna with scape cream-colored, unmarked ventrally, laterally, entirely dark brown dorsally; pedicel, flagellum dark brown to black; maxillary palp, labial palp with basal segments pale, ultimate segment dark brown.
Prothorax with pair of wide, brown stripes laterally, extending mesally and becoming reddish along transverse furrow, pale green mesally; transverse furrow in posterior region, not reaching lateral margins of segment, golden to dark brown setae throughout. Mesothorax, metathorax with pair of broad, dark brown stripes laterally, pale green mesally. Legs pale, unmarked except for small dark brown spot near anterior base of foretibia. Measurements: head width: 1.5 mm; ratio head width : eye width: 3.0–3.2 : 1; prothorax width: 1.2 mm, length: 0.9 mm.
Forewing, hindwing broad, with round apices. Forewing with venation heavy, especially at base of R,
Male: Abdomen with slightly enlarged spiracles (e.g., A7: spiracle diameter ~0.10× length of sternite); T9+ectoproct relatively long (~two thirds times length of T7), with dorsal invagination deep (~0.6× dorsal length of T9+
The thickness and the depth of the brown coloration of the veins on the forewing were considerably more robust in the male specimen than in the female.
BOLIVIA: Departments of La Paz, Santa Cruz.
(in addition to type above). Bolivia. Santa Cruz, Florida, 11 km. N.E. Achira, 1800 m, 3/XI/1999, cloud forest, Malaise trap, C. Porter & L. Stange (1F,
(based on holotype). Head: vertex cream-colored, with large, shiny, dark brown marks; posterolateral region of head probably with brown stripe near margin of eye; frons cream-colored with single brown spot mesally, below scapes, with distinctly protruding, mesal lobe; clypeus cream-colored, anterior margin thickened, raised above labrum; labrum cream-colored, with anterior margin indented mesally; distal edge of gena, lateral margins of clypeus with dark brown stripe. Antenna with dorsal surface of scape dark brown, frontal surface probably cream-colored; pedicel, flagellum dark brown throughout.
Prothorax probably light brown, with narrow, brown, longitudinal stripe laterally, with transverse furrow in central region, ending at two small pale spots interior to lateral margins, with short, dark setae throughout. Mesothorax, metathorax light to dark brown, probably with darker brown markings [as described by Banks, but faded now]. Measurements: head width: 1.2 mm; ratio head width : eye width: 2.7 : 1; prothorax width, length: not measured.
Forewing, hindwing broad, rounded apically, with robust venation, with following veins crassate: R-Cux1, base of
Male and female. Abdominal characteristics unknown.
COLOMBIA: Department of Tolima.
Holotype only. Also see below.
The characteristics of
Head, thorax as in
Male: Abdomen yellowish green. Tergites (T) and sternites (S) densely covered with both elongate, thin and short, thickened setae. Spiracles small. T9+
Venezuela.
Adults of this species are pale green, with a dorsal yellow stripe extending from the head through the abdomen; they have the following distinctive head and wing features. The frons is cream-colored or pale yellow; the genae and lateral margins of the clypeus have a black stripe; the scape is pale green with a red dorsolateral stripe extending from the tip to the base or almost to the base. The inverted U-shaped mark on the vertex that typifies most species of
Head cream-colored to yellowish, with vertex smooth, shiny; inverted U-shaped marking on vertex faint, reddish, when visible, separated mesally, not extending anteriorly to antennal fossae or scapes, broader posteriorly than anteriorly; area between eyes and vertex with reddish marks or coloration; frons unmarked, flat in female, broad (frontal view) and raised mesally (lateral view) in male; gena with long, brown stripe extending from near base of eyes continuing through basolateral part of clypeal margin; tentorial pits amber to pale. Antenna cream-colored to yellow, dorsum of scape with diffuse reddish, longitudinal stripe laterally, flagellum with pale bristles; maxillary palp with basal two segments pale, distal three segments dark brown to black laterally; labial palp with basal two segments pale, distal one with light brown at tip.
Thorax mostly green (living specimens) to cream-colored (preserved specimens). Prothorax flat, with broad, red to reddish brown stripes laterally, light cream-colored mesally; transverse furrow relatively deep, in middle region of segment, ending laterally in broad, whitish spot on lateral stripe; setae on dorsal, lateral surfaces long, slender, yellow to reddish, dense laterally. Mesothorax, metathorax with broad band of reddish to reddish brown laterally. Legs pale, cream-colored, without marks. Measurements: head width: 1.3 mm; ratio head width : eye width: 2.4–2.6 : 1; prothorax width: 1.0–1.1 mm, length: 0.6–0.7 mm.
Forewing, hindwing slender; forewing rounded apically, hindwing broadly acute; membrane clear, without fumose areas except around gradates. Forewing with veins crassate (male), without thickening (female); stigma transparent to slightly opaque, with 4 (female) to 6 (male) subcostal crossveins below, without marks; longitudinal veins, crossveins mostly light green, pinned specimens (male) with radial crossveins slightly darkened; gradate veins reddish brown, with some suffusion on surrounding membrane. Hindwing venation light green. Forewing 10.4–12.0 mm long, 3.5–3.9 mm wide (ratio, L : W = 2.9–3.1 : 1), height of tallest costal cell 0.8 mm (cell number 6–7); length of first intramedian cell 0.7–0.8 mm; 10 radial cells (closed cells between R and
Male: Abdomen with enlarged spiracles (e.g., A7: spiracle diameter ~0.23–0.26× length of sternite); abdominal sternites with small to no dorsal depression; subanal plate large, triangular, with ~twelve midsized setae; T9+ectoproct dome-shaped, with dorsal invagination shallow, extending approximately one half distance to anterior margin of T9, lateral margins of invagination slightly convex, base U-shaped; dorsal margin of ectoproct rounded throughout, with posteroventral margin extended distomesally in husky, rounded knob; ventral margin sclerotized posteriorly to base of callus cerci; callus cerci oblong, with ~ 30 robust microtrichia; circumference sclerotized heavily on posterior, anterior, ventral margins, lightly on dorsal margin; sclerotization contiguous with that on ventral margin of ectoproct, with narrow band of light sclerotization extending dorsally for very short distance from posterodorsal margin of callus cerci. S8+9 fused, with line of fusion not demarcated, but with slight fold on teneral specimen; S9 narrowed, considerably more heavily sclerotized than S8; dorsal margin relatively well sclerotized, including basally; terminus up-turned posteriorly, extending distally well beyond T9+
Female: See
The coloration of the body, markings, and wing venation deteriorates quickly and substantially after death.
VENEZUELA: States of Aragua, Falcón, Mérida, Táchira.
[in addition to those listed by
Head cream-colored to yellow, with vertex smooth, shiny; inverted U-shaped marking on vertex, light to dark red, narrow to separated mesally, not extending anteriorly to antennal fossae or scapes, broader posteriorly; area between eyes and vertex unmarked; frons unmarked; gena with long, black stripe extending from base of eyes continuing through lateral margin of clypeus; tentorial pits amber to light brown marginally. Antenna cream-colored to yellow, dorsum of scape with diffuse reddish, longitudinal stripe, flagellum with light brown to amber bristles; maxillary palp with basal two segments pale, distal three segments dark brown to black; labial palp with basal two segments pale, distal one with dark brown.
Thorax mostly green (living specimens) to cream-colored (preserved specimens). Prothorax short, flat, with broad, red to reddish brown, lateral stripes; transverse furrow shallow in middle region, with yellowish spots at lateral margins; setae on dorsal, lateral surfaces long, slender, yellow to reddish, dense laterally. Mesothorax, metathorax with reddish to reddish brown laterally. Legs pale cream-colored, without marks. Measurements: head width: 1.3–1.4 mm; ratio head width : eye width: 2.2 : 1; prothorax width: 1.0–1.1 mm, length: 0.4–0.5 mm.
Forewing rounded apically; hindwing broadly acute; membrane clear, without fumose areas. Forewing with veins uninflated, except base of
Male: Abdomen with large spiracles (e.g., A7: spiracle diameter ~0.24× length of sternite); subanal plate rectangular, with ~twelve robust setae; T9+ectoproct short, with dorsal invagination shallow, extending approximately one half the distance to anterior margin of T9, lateral margins of invagination fairly straight; dorsal margin of segment slightly rounded, with posteroventral margin extended distally in husky, rounded knob; ventral margin heavily sclerotized well beyond callus cerci; callus cerci large, oblong, with ~ 30 robust microtrichia; circumference lightly sclerotized, sclerotization separate from that on ventral margin of ectoproct, without dorsal extension. S8+9 fused, with no line of demarcation; S9 considerably more heavily sclerotized than S8; dorsal margin sclerotized mesally, lightly so basally; terminus rounded, blunt, not up-turned posteriorly, extending distally well beyond T9+
Female. See Fig.
The darkness and size of the head markings on the vertex vary among specimens.
VENEZUELA (northeastern): States of Aragua, Mérida, Trujillo.
[in addition to those listed by
The marking on the distolateral corner of the ventral surface of the scape varies from absent (some Venezuelan specimens) to large, dark and shiny (Bolivian specimens). Indeed the dorsal head markings of the Bolivian population are so dark and shiny that they appear similar to
Head golden to cream-colored, with vertex smooth, shiny; curved marking on vertex dark brown to black, large, prominent, broadly connected mesally, extending anteriorly to antennal fossae, but not between scapes; area between eyes and vertex unmarked; frons unmarked; gena with brown mark distally, not touching eyes, continuing along lateral margin of clypeus, but not reaching tip; tentorial pits with light brown margins. Antenna with scape golden to cream-colored, ventral surface with large, dark brown to black mark distolaterally; pedicel with brown ring; flagellum cream-colored to light tan dorsally, dark brown to black ventrally; maxillary palp with basal two segments pale, distal three segments dark brown; labial palp with basal segment pale, distal two segments with dark brown.
Prothorax cream-colored mesally, with wide, shiny, dark brown, lateral stripes, very thin, brown mesal line; transverse furrow shallow, in posterior region, ending laterally in slightly lightened circular area within dark lateral stripe; dorsal surface apparently with waxy covering, with fine pale setae dorsally, darker, heavier setae laterally. Mesothorax, metathorax cream-colored to tan mesally, marked with light brown to brown laterally. Legs pale, cream-colored, without marks. Measurements: head width: 1.4–1.5 mm; ratio head width : eye width: 2.5–2.7 : 1; prothorax width: 1.1–1.2 mm, length: 0.5–0.7 mm.
Forewing, hindwing rounded apically; membrane clear, largely without fumose areas. Forewing with veins uninflated, except basal section of
Male. Abdomen with unenlarged spiracles (e.g., A7: spiracle diameter ~0.05x length of sternite); subanal plate large, with ~ten robust setae; T9+ectoproct short, with dorsal invagination shallow, extending approximately one half the distance to anterior margin of T9, lateral margins of invagination slightly convex; dorsal margin of segment rounded distally, with posteroventral margin extended distally in husky, rounded knob; ventral margin sclerotized well beyond callus cerci; callus cerci large, oblong, with ~ 30–36 robust microtrichia; circumference sclerotized, sclerotization contiguous with that on ventral margin of ectoproct, with narrow band of light sclerotization extending dorsally from dorsal margin of callus cerci to top of T9+
Female. See
The darkness and sizes of the head and appendage markings on this species show considerable variability. Dorsal head markings range from faint, diffuse to dark, shiny brown or black. The basal flagellomeres range from entirely pale, to light brown on ventral and lateral surfaces only, to shiny dark brown or black on all surfaces.
BOLIVIA: Department of Cochabamba. VENEZUELA: States of Aragua, Lara.
[in addition to those listed by
This species is distinguished from other Andean
Among all
Body color: brown to brownish, sometimes with yellow mesally. Head cream-colored, with vertex smooth, often shiny; inverted U-shaped marking on vertex brown to reddish brown, usually prominent, narrowing and sometimes separated mesally, not extending anteriorly to area between scapes; antennal fossa, area between eyes and posterior half of vertex unmarked; frons often with brown, small to large, triangular marking centrally; gena with broad, brown stripe extending from eye along lateral margin of gena, clypeus; tentorial pits amber-colored. Antenna pale, dorsum of scape with brown longitudinal stripe distally, extending onto dorsal surface of pedicel; maxillary palp, labial palp with basal two segments pale, distal segments dark brown.
Prothorax yellowish mesally, with broad, brown to reddish brown, longitudinal, submesal stripes almost reaching lateral margin; transverse furrow in posterior region, not reaching lateral margins; dorsal surface with thin, pale setae, sparse mesally, denser laterally. Mesothorax, metathorax brown to yellowish brown laterally, yellow mesally. Measurements: head width: 1.2–1.3 mm; ratio head width : eye width: 2.4 : 1; prothorax width, 1.0 mm; length: 0.5–0.6 mm.
Forewing, hindwing narrow, rounded apically; membrane clear, hyaline, without fumose areas, with slender venation; stigma lightly opaque to clear, with four to five light to dark brown subcostal crossveins below stigma, area surrounding crossveins unmarked; longitudinal veins mostly pale, slightly darker at intersections, base of
Male. Abdomen with long and short setae, especially dense posteriorly, on A7–A9; spiracles moderately enlarged (e.g., A7: spiracle diameter ~0.15x length of sternite). T9+ectoproct rounded, sloping ventrally (lateral view), with dorsal invagination rounded, shallow (extending approximately one half distance to anterior margin of T9), posteroventral knob well defined, heavily sclerotized, bending mesally; ventral margin lightly sclerotized beyond callus cerci; callus cerci large, ovate, circumference lightly sclerotized; subrectal plate bearing field of ~15 short setae. S8+9 fused, with line of fusion not demarcated; dorsal margin sclerotized, especially basal 2/3rds; terminus rounded, extending distally only slightly beyond the tip of T9+
Among the specimens we examined, there was considerable variation in the size and degree of separation of the dorsal head markings, in the presence or absence and size of a frontal marking, and the darkness and amount of brown coloration on the wing veins.
BOLIVIA (central): Department of Cochabamba. COLOMBIA (west to central): Departments of Cundinamarca, Quindio, Valle del Cauca.
(in addition to types above). Bolivia.
(Figs
The species name “
The Andean species
Externally, this species also resembles the Argentinian
Head white to cream-colored with dark brown to black markings; vertex smooth, often shiny; inverted U-shaped marking dark brown, prominent but small, dot-like, narrowing and separated mesally, not extending anteriorly to area between scapes; antennal fossa, area between eyes and posterior half of vertex unmarked; frons unmarked, slightly swollen laterally in males; gena with dark brown to black stripe extending from near base of eye along lateral margin of gena, most of clypeus; tentorial pits amber-colored. Antenna pale, dorsum of scape with short, brown longitudinal stripe distally, not extending onto dorsal surface of pedicel; maxillary palp with basal two segments pale, three distal segments dark brown; labial palp with basal segment pale, middle segment light brown, distal segment dark brown.
Prothorax yellowish mesally, with broad, diffuse, reddish brown, longitudinal, lateral stripes, extending to lateral margin; transverse furrow in mesal region, almost reaching lateral margins; dorsal surface with thin, pale setae, sparse mesally, denser laterally. Mesothorax, metathorax marked with reddish brown laterally, yellow mesally; both with pair of brown spots on margin between prescutum and scutum (smaller on metathorax), pair of small brownish spots laterally. Measurements: head width: 1.5 mm; ratio head width : eye width: 2.3 : 1; prothorax width: 1.0 mm; length: 0.5 mm.
Forewing with apex rounded, hindwing acute; membrane clear, hyaline, without fumose areas, with venation slender (female) to very slightly crassate (male); stigma lightly opaque to clear, with three to four light brown subcostal crossveins below stigma, area surrounding crossveins unmarked; longitudinal veins light green, all costal, radial crossveins brown to brownish; transverse veins in posterior sector of wing brown to pale; gradates dark brown without suffusion. First gradate vein meeting
Male. T9+
COLOMBIA: States of Antioquia, Valle del Cauca.
(in addition to holotype). 1F, same data as holotype (paratype,
This species and
Adults of
Pale green, with dorsal surface mostly yellowish from vertex to tip of abdomen. Head with vertex raised, smooth, shiny, with pair of red to reddish brown spots or crescent shaped marks laterally; frons smooth, flat, with or without small mesal mark anteromesal to base of antennae; gena, lateral edge of clypeus with reddish to brownish stripe; maxillary palp with distal three segments dark brown, pale at articulations, basal segments unmarked; labial palp unmarked except tip of distal palpomere dark brown; antenna light yellow basally, becoming slightly brownish distally; scape, pedicel without marks. Prothorax length and width about equal, narrowing anteriorly, unmarked or slightly tinged with red. Legs yellow-green, with similarly colored setae; tarsus paler; base of claws strongly dilated. Measurements: head width: 1.2–1.4 mm; ratio head width : eye width: 2.2–2.4 : 1; prothorax width: 0.9–1.0 mm, length: 0.6–0.7 mm.
Forewing, hindwing elongate, with broadly acute tips; membrane mostly clear, hyaline, with fumose areas around brown sections of transverse veins, gradate veins; stigma clear to slightly opaque; longitudinal veins mostly green; transverse veins with brown at most intersections; gradate veins, first two r-m crossveins, last crossvein of distal b’ cell entirely brown, with surrounding membrane suffused with brown. Forewing 13.5–15.9 mm long, 4.6–5.3 mm wide (ratio, L : W = 2.9–5.0: 1); height of tallest costal cell 0.8–1.2 mm (cell number 8–9); width of first intramedian cell 0.8–1.0 mm; 11–13 radial cells (closed cells between R and
Male. Abdomen with small spiracles (e.g., A7: spiracle diameter ~0.02x length of sternite); T9+ectoproct relatively long (~2/3× length of T7), with dorsal invagination shallow (~1/3 dorsal length of T9+
Among the specimens we examined, there was considerable variation in the size and depth of the reddish markings on the frons mesal to the eyes and below the base of the antenna. The specimens from the unknown type locality in Colombia, Azogues in south-central Ecuador, and Cajamarca in Peru had distinct markings, whereas the specimens from Cuenca, Huancavelica, and Matucana, Lima, in Peru, had no mark or only a speck of red on the frons mesal to the eyes.
COLOMBIA (no locality). ECUADOR (south central): Provinces of Cañar, Huancavelica. PERU (central coast and north): Provinces of Lima, Cajamarca, Cuzco.
(in addition to types above). ECUADOR.
(also see
Head cream-colored to yellow with red markings; vertex smooth, shiny, with U-shaped marking reduced to two arms or two prominent spots separated at center; area between antennae unmarked; red stripe between eye and sides of vertex, extending posteriorly; dorsal antennal fossa entirely red or with thin reddish stripe mesally, not extending between antennae. Frons cream-colored, without markings; clypeus cream-colored, with black stripe on lateral margin, contiguous with black genal mark, which reaches almost to eye. Antenna mostly cream-colored, scape, pedicel with broad dorsolateral red stripe; flagellum yellow with black bristles; maxillary palp with two basal segments pale, distal three segments black dorsally, with intersections, tip of distal palpomere pale; labial palp with basal two segments pale, ultimate segment light brown to black shading laterally.
Prothorax green with cream-colored stripe mesally, pair of broad, dark red stripes sublaterally; transverse furrow in posterior region, not deep, reaching lateral margins of segment, long, cream-colored to golden setae throughout. Mesothorax reddish brown centrally, green, marked with red laterally; metathorax green with reddish brown markings. Legs light green, unmarked. Measurements: head width: 1.4–1.5 mm; ratio head width : eye width: 2.7: 1; prothorax width: 0.9–1.2 mm, length: 0.7–0.8 mm.
Forewing, hindwing with moderate width. Forewing with rounded apex, robust, but without heavy venation, with very slight swelling at furcation of
Male. Abdomen with large spiracles (e.g., A7: spiracle diameter ~0.2× length of sternite); A7-A9 with numerous setae extending from robust setal bases; T9+ectoproct relatively long (~0.6× length of T7), with dorsal invagination deep (~0.5× dorsal length of T9+
Female. See
Oviposition of stalked, clustered eggs (probably infertile) (see images,
VENEZUELA: States of Aragua, Portuguesa.
[in addition to those listed by
(Figs
The holotype (a female) is the only specimen of this species that we have seen, and we were reluctant to describe it as new on the basis of such limited material. However, the specimen is very well preserved, and its external features (head and body coloration and markings, wings) are notable. The abdomen is cleared, stained, and in a vial attached to the specimen. Because of the specimen’s importance as the northernmost record for the genus, we describe it to facilitate future identifications.
The type locality of this species is in the highlands of southern Mexico (State of Chiapas). Its species name,
The female holotype expresses many of the characteristics that typify most
Head cream-colored with red markings; vertex smooth anteriorly, shiny, with U-shaped marking large, robust, lateral arms separated anteriorly, meeting posteriorly, with lateral margins marked with red adjacent to eyes, posterior section unmarked; dorsal fossae probably with small, red mesal mark; area between antennae unmarked; frons cream-colored, unmarked; clypeus with lateral margin red; tentorial pits amber-colored; gena with bold, black stripe throughout, contiguous with red clypeal mark. Antenna: cream-colored, dorsal surface of scape possibly with tinge of red, but no stripe, flagellar setae pale; maxillary palp pale, distal segments with thin black lateral stripe; labial palp pale throughout.
Thorax mostly cream-colored; prothorax with pair of broad lateral bands – red anteriorly, green posteriorly; transverse furrow slightly posterior to middle of segment, not extending to lateral margins; setae pale. Legs cream-colored, unmarked. Measurements: head width: 1.4 mm; ratio head width : eye width: 2.3: 1; prothorax width: 1.0 mm, length: 0.8 mm.
Forewing, hindwing somewhat broad, apex slightly acute; membrane clear, hyaline. Forewing with veins robust, not crassate;
Male. Unknown.
MEXICO: Central Highlands region in state of Chiapas.
. Holotype only.
This species is distinguished by a golden yellow head and a round, black spot on the frons, and dark wing venation. It differs from other species that have a prominent black frontal spot and cream-colored antennae (
Head: As described by
Forewing, hindwing with moderate width. Forewing with rounded apex, hindwing with apex slightly acute; venation not heavy, very slight swelling at furcation of
Male. Abdomen with small spiracles (e.g., A7: spiracle diameter ~0.05× length of sternite); A7-A9 with numerous setae, those extending from pleural region robust; without microtholi; T9+ectoproct relatively long (~0.5× length of T7), with dorsal invagination deep (~0.5× dorsal length of T9+
Female. See
COLOMBIA (central): Department of Colombia. VENEZUELA (northeastern): States of Trujillo, Táchira, Mérida, Lara.
[in addition to those listed by
In the original description of
At this time,
See
Forewing, hindwing delicate; forewing rounded apically; hindwing tapered, but rounded apically; membrane entirely clear, without markings or fumose areas; stigma clear; all veins light green, unmarked. Forewing with
Male (paratype, slightly teneral). Abdomen with unenlarged spiracles (e.g., A7: spiracle diameter ~0.06× length of sternite); subanal plate obscured; T9+ectoproct moderately long (~0.5× length of T7), with dorsal invagination deep (~0.8× dorsal length of T9+
COSTA RICA: Province of Puntarenas. PERU: Province of Lambayeque [tentative].
Two paratypes with same data as holotype (1M, 1F,
Note 1: The above male specimen from Peru expresses almost all of the characteristics that distinguish
Note 2: We have seen female specimens from Bolivia that resemble
(Figs
The species is named for Norman D. Penny of the California Academy of Sciences, who died during the preparation of this revision. Norm was a world-renown Neuropterist and valued colleague. We (CAT) had some disagreements with Norm, but our shared attachment to the
The
Head badly discolored, probably cream-colored with red markings; vertex smooth anteriorly, shiny, with a pair of red spots in place of U-shaped marking, pair of larger red marks around posterolateral margin of raised vertex; dorsal fossae either pale or reddish; no apparent markings between antennae; frons with hint of mesal red spot below each antenna; reddish marking(s) near front of frons; gena without visible coloration; clypeus, labrum without marks. Antenna: dorsal surface of scape with broad red longitudinal stripe laterally, frontal surface with indistinct dark markings mesally; pedicel, flagellum pale, with pale setae basally, darker setae distally; maxillary, labial palpi pale, without discernible marks.
Thorax without distinct coloration or marks; prothorax short, with pale setae; mesothorax perhaps with pair of dark spots on frontal surface of scutum. Legs pale, unmarked, with pale setae. Measurements: head width: 1.3 mm; ratio head width : eye width: 2.4 : 1; prothorax width: 0.9 mm, length: 0.5 mm.
Forewing rounded apically; hindwing slightly acute, both with venation robust; membrane mostly clear, dull; stigma opaque, brownish, with three faint subcostal crossveins, basal one with brown suffusion. Forewing: 10.9 mm long, 3.7 mm wide (ratio, L : W = 3.0 : 1);
Male. Abdomen with very large spiracles (e.g., A7: spiracle diameter ~0.25× length of sternite); subanal plate small, with ~4 setae; T9+ectoproct long (~0.7 length of T7), with dorsal invagination deep (~0.6x dorsal length of T9+
COSTA RICA: Provinces of Cartago, Puntarenas.
(in addition to holotype above). Puntarenas, Costa Rica: Finca Las Cruses near San Vito, Puntarenas Province, Costa Rica 1800m elev. 13.III.69 at black light, J. Sheldon #1969-140 (paratype, F,
(Figs
The type locality of this species is the Andean valley of Cochabamba in central Bolivia. The locality is also known by its Quechua name – Quchapampa. Both the Spanish and the Quechua names are derived from the Quechua words “qhucha” (meaning “small lake”) and “pampa” (meaning “open plain”) (
Head cream-colored with dark brown to black markings; vertex with inverted U-shaped marking large, robust, with lateral arms meeting anteriorly, extending forward into dorsal fossa, between antennae; mesal side, posterior margin of dorsal fossa marked with dark brown; posterior parts of vertex unmarked; frons with large, prominent mesal mark; clypeus with lateral margin black; tentorial pits surrounded by dark brown; gena with black stripe from eye, contiguous with black mark on clypeal margin. Antenna: scape cream-colored, dorsum with large, brown longitudinal mark; pedicel, flagellum dark brown to black basally, becoming lighter distally, with cream-colored intersegmental membrane; maxillary palp with basal two segments pale, three distal segments black, articulations pale; labial palp with basal segment pale, distal two segments marked with black.
Thorax with distinct longitudinal cream-colored stripe mesally; prothorax with pair of broad, dark reddish brown stripes laterally; transverse furrow in posterior region of segment, extending to lateral margins; setae mostly elongate, pale. Measurements: head width: 1.3–1.5 mm; ratio head width : eye width: 2.1–2.6 : 1; prothorax width: 1.1–1.2 mm, length: 0.6–0.7 mm.
Forewing, hindwing somewhat broad, apex round; membrane clear, hyaline, without fumose area; stigma slightly opaque to light brown; longitudinal veins green, marked with brown at intersections with transverse veins; transverse veins mostly brown or marked with brown; forewing with veins robust, not crassate;
Male. Abdomen with small spiracles (e.g., A7: spiracle diameter ~0.04× length of sternite); subanal plate substantial, with ~20 setae in a well defined group; T9+
BOLIVIA (central): Department of Cochabamba (Provinces of Carrasco and Chapare).
(in addition to holotype). Same data as holotype (1F, paratype,
This species is recognized by the following set of features: body mostly cream-colored to green; head cream-colored, vertex with robust reddish brown, inverted U-shaped mark with lateral bar extending to mesal margin of eyes; frons with broad, reddish brown, transverse band above clypeus; antenna with basal two segments reddish, flagellum pale; thorax probably with yellowish mesal stripe dorsally, reddish brown stripe sublaterally, legs pale, unmarked, with pale setae; wings hyaline, lightly fumose around gradate veins, with veins green, except costal crossveins, gradate veins black. In the male, the spiracles are not enlarged; the setae on the pleural membrane of A7, A8 have slightly enlarged bases; the terminus of S8+9 is without flanged setae; and the gonarcus, mediuncus, and gonosaccus are as in Figs
Head cream-colored with reddish brown to dark brown markings; vertex smooth, shiny; inverted U-shaped marking prominent, reddish brown, broad, connected anteromesally above antennae, with lateral bar extending to midregion of eye; posterior half of vertex unmarked; dorsal antennal fossa with thin reddish brown stripe mesally, extending between antennae to reddish brown anterior fossa. Frons with narrow, elongate, brown stripe between antennal fossae (“Y-shaped” mark of Navás), anterior margin of frons with pair of heavy reddish brown transverse bands from midregion to genal mark; clypeus with pair of large reddish brown marks, separated mesally, extending to margin laterally; gena with large, dark brown to black mark extending from eyes, contiguous with dark mark on lateral margin of clypeus; tentorial pits surrounded by reddish brown. Antenna with scape, pedicel reddish, unmarked; flagellum cream-colored; maxillary palp, labial palp with two basal segments pale, other segments dark brown.
Prothorax cream-colored mesally, with pair of wide, reddish brown stripes sublaterally, reddish brown extending mesally along transverse furrow; transverse furrow in posterior region, not reaching lateral margins of segment, long, cream-colored to golden setae throughout. Mesothorax, metathorax probably cream-colored mesally, with pair of broad, reddish brown stripes laterally. Legs entirely cream-colored, unmarked. Measurements: head width: 1.3–1.6 mm; ratio head width : eye width: 2.0–2.3 : 1; prothorax width: 0.8–1.1 mm, length: 0.6–0.7 mm.
Forewing, hindwing of moderate width, with apex rounded or slightly subacute. Forewing with robust, but not heavy venation, some swelling at furcation of
Male. Abdomen with small spiracles (e.g., A7: spiracle diameter ~0.03× length of sternite); A6–A9 with numerous robust, elongate setae extending from large setal bases, especially notable on pleural regions; T9+ectoproct relatively long (~0.6× length of T7), with dorsal invagination deep (~0.7× dorsal length of T9+
Among the specimens we examined, there was variation in the depth and size of the reddish brown markings on the vertex and frons.
PERU: Region of La Libertad, Province of Lima (Districts of Chancay, San Isidro).
(in addition to types above). Peru.
(Figs
The species is named for Lionel A. Stange, a life-long student of the systematics and natural history of myrmeleontoid
Like
The three male specimens of
Head yellow to whitish cream-colored with red to reddish black markings; vertex smooth, often shiny, with inverted U-shaped marking reduced to two anterolateral red spots; fossae, posterior parts of vertex, area between antennae unmarked; frons cream-colored, with dark red markings laterally between base of antenna and margin of eyes, small to large red mesal spot between base of antennae and clypeus; clypeus with dark red mark on lateral margin; tentorial pits amber-colored; gena with reddish black to black stripe. Antenna with scape, pedicel, flagellum cream-colored, unmarked; maxillary palp with basal two segments pale, three distal segments black with articulations pale; labial palp with basal segment pale, distal two segments with black.
Thorax green, with distinct longitudinal yellow stripe mesally; prothorax with transverse furrow in central region of segment, extending almost to lateral margins; setae mostly elongate, pale. Measurements: head width: 1.5–1.6 mm; ratio head width : eye width: 2.3–2.8 : 1; prothorax width: 1.0–1.1 mm, length: 0.8 mm.
Forewing, hindwing elongate, with acute tips; membrane mostly clear, hyaline, with brown, fumose areas around brown veins or brown markings on veins; stigma clear to slightly opaque; longitudinal veins mostly green; transverse veins mostly green, with brown at intersections; gradate veins, r-m crossveins, distal crossveins of most b’ cells entirely brown. Forewing 16.0–17.6 mm long, 5.0–6.0 mm wide (ratio, L : W = 2.9–3.0 : 1); height of tallest costal cell 1.3–1.5 mm (cell number 5–6); width of first intramedian cell 0.9–1.0 mm; 14–18 radial cells (closed cells between R and
Male. Abdomen with small spiracles (e.g., A7: spiracle diameter ~0.03× length of sternite); T9+ectoproct relatively long (~half length of T7), with dorsal invagination moderately deep (~0.4x dorsal length of T9+
BOLIVIA: State of La Paz. ECUADOR: Province of Pichincha. PERU: Province of Andahuaylas. All localities at ~3000 meters elevation.
(in addition to holotype). 4M, 1F, one without abdomen, all with same data as holotype (all paratypes,
The specimens from Bolivia and Ecuador vary somewhat in the depth of coloration in their markings and in the size of their wings, but all exhibit all of the markings mentioned in the description. In comparison, the marks on the sole specimen from Peru are light, and its wings are slightly shorter; most notably, it appears to lack a marking on the frons. We suspect that the natural coloration of this specimen has faded; but, we cannot exclude the possibility of individual variation or a pale colored population. This specimen is the only male that is not teneral.
Previously, this species was known from the single (slightly teneral) female holotype (
Head yellow dorsally; vertex smooth, shiny; inverted U-shaped marking prominent, dark red, broad posteriorly, separated or lightly connected anteromesally, not extending anteriorly to area between scapes; dorsal antennal fossa pale, with dark reddish stripe mesally, reaching mesal base of scape; area between eyes and posterior half of vertex cream-colored, unmarked; frons, clypeus yellow, without markings; gena with large, black mark from eyes onto clypeus; tentorial pits amber. Antenna with scape cream-colored, with broad, diffuse dark red stripe extending full length, onto dorsal surface of pedicel; pedicel with frontal surface pale, dorsal surface dark red; flagellum yellow, except basal flagellomere brownish; maxillary palp with basal two segments pale, distal three segments black; labial palp with basal segment pale, distal two segments black.
Prothorax green, with pair of broad, dark red stripes along entire lateral length, with pair of pale, circular spots at level of transverse furrow; transverse furrow in posterior region of prothorax extending laterally only to pale spots. Mesothorax, metathorax green, with ivory mesally; mesonotum with pair of reddish spots on anterior margin of scutellum; metanotum unmarked. Legs pale, unmarked. Measurements: head width: 1.4–1.6 mm; ratio head width : eye width: 2.3 : 1; prothorax width: 1.1–1.4 mm, length: 0.7–0.8 mm.
Forewing, hindwing elongate; forewing with round apex; hindwing with slightly acute apex. Forewing with venation not inflated, except at base of
Male. Unknown.
Female. See
No variation of note.
VENEZUELA: State of Amazonas.
(in addition to holotype). Venezuela,
The
1 | Longitudinal veins of forewing with conspicuously alternating sections of white and dark brown or black; mark on vertex V-shaped (not rounded) anteriorly, extending forward toward scapes |
|
1' | Longitudinal veins of forewing mostly light green to tan, sometimes with brown at junctions with transverse veins, or brown for long stretches; mark on vertex in shape of inverted U (rounded anteriorly), not extending forward toward scape |
|
2 (1) | Mark on vertex extending anteriorly between scapes; frons usually with pair of light brown spots mesally (Fig. |
|
2' | Mark on vertex extending towards scapes, but usually not between them; frons without mesal mark (Fig. |
|
3 (1) | Palpomeres marked with black or dark brown laterally; forewing with most transverse veins green, sometimes brown; male abdominal spiracles enlarged or not |
|
3' | Palpomeres cream-colored, without marks (Fig. |
|
4 (3) | Forewing with longitudinal veins regularly marked with brown at intersections, costal crossveins and sections of other crossveins marked with brown; male abdominal spiracles slightly to moderately enlarged |
|
4' | Forewing with longitudinal veins (except sometimes the anal veins) and all crossveins entirely green or with very slight darkening at intersections with transverse veins; male abdominal spiracles may be greatly enlarged |
|
5 (4) | Head with prominent, black stripe on gena and clypeus, from base of eyes at least part way along lateral clypeal margin (Fig. |
|
5' | Head with small genal mark, not in contact with base of eye or margin of clypeus (Fig. |
|
6 (4’) | Forewing small (no more than 13.5 mm long, 4.8 mm wide); 10-12 radial cells; 5-6 inner gradates, 5-6 outer gradates (Figs |
|
6' | Forewing large (over 14.5 mm long, 5.3 mm wide); 13 radial cells; 7 inner gradates, 9 outer gradates; male unknown (Fig. |
|
This species, along with
Head: vertex cream-colored to tan, with dark reddish brown “U-shaped marking” prominent, usually broken mesally, with pigmented coloration extending forward from tips of marking to area between scapes, not extending to frons; anteromesal margin of antennal fossa also with elongated, reddish brown to brown marking extending to mesal margin of scapes; gena, lateral margins of the clypeus with dark brown stripe. Antenna with scape cream-colored to tan with diffuse, brown, longitudinal mark or marks laterally, pedicel mostly brown or light brown, flagellum tinged with brown basally, tan distally; basal palpomeres pale, mesal palpomeres brown, terminal segments pale with brown basally.
Prothorax brown to dark brown laterally, tan to light brown mesally, with transverse furrow in posterior region, not reaching lateral margins, with short, dark setae throughout. Mesothorax, metathorax brown to dark brown, with darker brown markings. Measurements: head width: 1.3–1.4 mm; ratio head width : eye width: 2.3–2.5 : 1; prothorax width: 1.0–1.2 mm; prothorax length: 0.7 mm.
Forewing, hindwing broad to very moderately narrow, with apices rounded (forewing) to rounded or slightly subacute (hindwing), venation slender to slightly robust; alar membrane hyaline, with suffusion light, often obscured by reflection; stigma light brown, opaque, with four to five black crossveins below surrounded by dark brown marks; longitudinal veins with alternate creamy and dark markings (usually dark brown at intersections with dark veins and cream-colored between intersections); transverse veins, gradate veins, crossveins dark. Forewing 10.2–11.6 mm long, 3.7–4.3 mm wide (ratio, L : W = 2.7–3.0 : 1); height of tallest costal cell 0.7–0.9 mm (cell number 3–6); first intramedian cell ovate, 0.8–0.9 mm long; 9–10 radial cells (closed cells between R and
Male. Abdomen with greatly enlarged spiracles, with large flaps on opening (e.g., A7: spiracle diameter ~0.30× length of sternite); T9+ectoproct short, rounded posterodorsally, with dorsal invagination extending ~3/4 distance to anterior margin of T9, lateral margins of invagination straight; ventral margin of T9+
The
Unknown.
ARGENTINA (northwest to central east): Provinces of Buenos Aires, Catamarca, Cordoba, Entre Rios, Salta, Tucumán.
(in addition to types above). Argentina.
The above specimen from La Rioja (
The lectotype of
Head: vertex light brown to brown, with U-shaped marking prominent to somewhat obscured, brown coloration extending anteriorly between the scapes, usually to the frons, forming an X-shaped frontal marking below antennal fossae; antennal fossa with large, dark brown mark extending around dorsal margin, between antennae, and around frontal margin; gena, lateral margin of clypeus with dark brown stripe. Antenna cream-colored; scape with diffuse, brown, longitudinal mark or marks; pedicel mostly brown; flagellum tinged with brown. Basal two labial palpomeres pale, distal three brown; membrane between palpomeres cream-colored; distal maxillary palpomere light brown, basal two cream-colored to tan.
Prothorax with brown, longitudinal stripes sublaterally, laterally; transverse furrow in posterior region, not reaching lateral margins; short, dark setae throughout. Mesothorax, metathorax with lighter, more diffuse brown markings. Measurements: head width: 1.3–1.4 mm; ratio head width : eye width: 2.4–2.6 : 1; prothorax width: 1.0 mm; prothorax length: 0.5–0.7 mm.
Forewing, hindwing moderately narrow, with well-rounded apices, robust venation; alar membrane clear, with light to dark brown suffusion adjacent to dark veins; stigma usually light brown, cloudy to opaque, with three to four black crossveins below, surrounded by dark brown fumose to solid marks; longitudinal and transverse veins with alternate creamy and dark markings (usually dark brown at intersections with crossveins and cream-colored between intersections); gradate veins, crossveins dark. Forewing 8.6–11.1 mm long, 2.8–3.8 mm wide (ratio, L : W = 2.9–3.1 : 1); height of tallest costal cell 0.6–0.8 mm (cell number 5–6); first intramedian cell ovate, 0.6–0.8 mm long; 8–9 radial cells (closed cells between R and
Male. Abdomen with spiracles small (e.g., A7: spiracle diameter ~0.04x length of sternite); T9+ectoproct short, rounded posteriorly, with dorsal invagination deep, almost reaching anterior margin of T9, lateral margins of invagination straight to convex; callus cerci large, ovate, entire margin lightly sclerotized, sclerotization extending distally along posteroventral margin of segment; subanal plate small, delicate, with patch of one to eight irregularly placed setae. S8, S9 fused, fusion demarcated; dorsal margin with light sclerotization anteriorly, heavier throughout terminal ~1/5th of S8+9; terminal setae slightly robust, otherwise unmodified. Gonarcus with bridge slender, lateral apodemes slightly expanded, blunt, process on sides of lateral apodemes curved smoothly, perpendicularly (lateral view) and slightly inward (frontal view) from gonarcal arm; mediuncus long, narrow, with dorsal margin slightly bowed; gonosaccus distinctly bilobed, each lobe with large patch of robust gonosetae on enlarged bases – anterobasal gonosetae smaller, shorter, on smaller bases than posterodistal gonosetae; hypandrium internum U-shaped with arms bending outward distally, comes elongate, irregular.
Female. Spermatheca round, pillbox-shaped, velum obscured, invagination present but size not clear, spermathecal duct with U-shaped bend and curve; subgenitale with short knob-like protrusion.
The clarity of the alternate white and dark markings on the wing veins is quite variable; white areas are often dull. Similarly, the size, shape, and prominence of the head and facial marks can also vary; occasionally the paired brown frontal marks are absent, but the mark on the vertex usually extends anteriorly between the scapes and onto the top of the frons.
We examined one particularly small male specimen (Argentina: La Rioja, Guadacol, 1–3 December 1983, Luis F. Pena,
ARGENTINA (northwest to central east): Provinces of Buenos Aires (
(in addition to type). Argentina.
Adults of
Several series of specimens in the
This species resembles
Head: vertex smooth, slightly depressed mesally, inverted U-shaped marking on vertex reddish brown, usually prominent, often separated mesally but with pigmented marking joining mesally, extending anteriorly toward but not reaching area between scapes; anteromesal margin of dorsal antennal fossa also with dark reddish brown mark; area between eyes and posterior half of vertex cream-colored, unmarked; frons cream-colored with elongate, narrow, linear marking from between antennae to center of frons, anterior margin of frons with or without reddish brown, transverse stripe; clypeus cream-colored to tan; gena, posterolateral margin of the clypeus with dark brown stripe; tentorial pits with brown margins. Antenna with scape cream-colored to light tan, unmarked; pedicel cream-colored to tan, with distal ring of brown; flagellum cream-colored to light tan basally, becoming darker distally; maxillary palp with basal two segments pale, distal segments brown; labial palp with basal segment pale, distal two segments brown.
Prothorax green mesally, with wide, brown, lateral stripes, thin, brown mesal stripe; transverse furrow in posterior region, not reaching lateral margins; short, dark setae throughout. Mesothorax, metathorax yellowish brown with light brown markings. Measurements: head width, 1.2–1.4 mm; ratio head width : eye width, 2.4–2.6 : 1; prothoracic length, 0.5–0.8 mm; prothoracic width, 0.8–1.0 mm.
Forewing, hindwing clear, hyaline, without fumose areas, with slender venation; stigma lightly opaque, with three to six light to dark brown subcostal crossveins below stigma surrounded by dark brown marks; all veins, except anal veins, cream-colored to light green, without suffusion. Forewing 9.8–13.4 mm long, 3.3–4.8 mm wide (ratio, L : W = 2.8–3.1 : 1); height of tallest costal cell 0.6–1.0 mm (cell number 6–8); width of first intramedian cell 0.7–0.9 mm; 10–12 radial cells (closed cells between R and
Male. Abdomen with greatly enlarged spiracles having large flaps on opening (e.g., A7: maximum spiracle diameter ~0.23–0.30× length of sternite); T9+ectoproct short, with dorsal invagination extending almost to anterior margin of T9, lateral margins of invagination straight to convex; dorsal margin rounded distally; posterior half of ventral margin lightly sclerotized – sclerotization, contiguous with and extending distally from sclerotization around circumference of callus cerci, extending posteriorly in small, rounded knob; callus cerci oblong; subanal plate small, with ~ eight setae. Sternite S8, S9 fused, with line of fusion distinct, terminating in sclerotized, elongate, plate-like extension; dorsal margin sclerotized for entire length, heavier throughout terminal ~one third; terminal setae robust, flanged. Gonarcus with bridge slender, lateral apodemes narrow (lateral view), slightly expanded, rounded distally; process on side of lateral apodeme extending dorsally (lateral view), turning inward distally (frontal view) from gonarcal arm; mediuncus long, narrow, with dorsal margin straight, terminus with small knob; bilobed gonosaccus with large patch of well-spaced gonosetae on enlarged bases – anterobasal gonosetae slightly smaller, shorter, on smaller bases than posterodistal gonosetae; hypandrium internum U-shaped with arms straight (frontal view), bending upward (lateral view), comes delicate.
The species shows large variation in both size and coloration. For example, forewing length ranges from 9.8 to 13.4 mm, and the number of radial cells from 10-12 (forewing) and 9-12 (hindwing). In addition, the enlarged spiracles on the abdomen vary in size from ~0.23 to 0.30. Some of the variation in dorsal and frontal markings on the head is illustrated in Fig.
ARGENTINA (northwest): Provinces of Catamarca, Córdoba, La Rioja, Mendoza, Salta, Tucumán.
(in addition to the types above). Argentina.
A series of 11 unidentified female specimens (Figs
Externally, the dorsal head markings and largely green longitudinal veins marked with brown at intersections distinguish
Head cream-colored, with vertex smooth, often shiny; inverted U-shaped marking on vertex reddish brown, usually prominent, narrowing and sometimes separated mesally, not extending anteriorly to area between scapes; anteromesal margin of dorsal antennal fossa with reddish brown mark (sometimes pale); area between eyes and posterior half of vertex cream-colored, unmarked; frons cream-colored, with markings absent or light reddish brown stripe mesally, or reddish tinge along anterior margin; clypeus cream-colored to tan, with dark brown spot laterally; gena with brown mark not touching eyes, approaching clypeus; tentorial pits with light brown margins. Antenna with scape cream-colored to light tan, unmarked; pedicel cream-colored to tan, with distal ring of brown; flagellum cream-colored to light tan basally, becoming darker distally; maxillary palp with basal two segments pale, mesal two segments brown dorsally, laterally, distal segment brown; labial palp with basal segment pale, distal two segments with brown.
Prothorax relatively short, green mesally, with wide, brown, lateral stripes, thin, brown mesal stripe; transverse furrow in posterior region, not reaching lateral margins; with short, dark setae throughout. Mesothorax, metathorax yellowish brown with light brown markings. Measurements: head width: 1.2–1.3 mm; ratio head width : eye width: 1.7–2.3 : 1; prothorax length: 0.4–0.6 mm; prothorax width: 0.9–1.1 mm.
Forewing, hindwing clear, hyaline, without fumose areas, with slender venation; stigma lightly opaque to clear, with four to five light to dark brown subcostal crossveins below stigma, area surrounding crossveins unmarked; longitudinal veins usually mostly green, with brown at intersections, sometimes with extensive brownish sections; transverse veins, crossveins mostly brown to golden brown. Forewing 10.3–12.4 mm long, 3.6–4.4 mm wide (ratio, L : W = 2.8–2.9 :1); height of tallest costal cell 0.6–0.8 mm (cell number 6–8); length of first intramedian cell 0.7–1.0 mm; 9–12 radial cells (closed cells between R and
Male. Abdomen with slightly enlarged spiracles (e.g., A7: spiracle diameter ~0.10× length of sternite); T9+ectoproct short, with dorsal invagination shallow, extending approximately one half the distance to anterior margin of T9, lateral margins of invagination straight to slightly convex; dorsal margin rounded distally, with posteroventral margin extended distally in well defined knob; ventral margin lightly sclerotized beyond callus cerci; callus cerci large, ovate, circumference lightly sclerotized, sclerotization contiguous with that on ventral margin of ectoproct. S8+9 fused, with line of fusion not demarcated; setae on ventral surface relatively long, not dense; dorsal margin lightly sclerotized, except near base; terminus extended, upturned distally, flat, plate-like, well sclerotized, bearing numerous, large, flanged setae along dorsal margins and heavy, unflanged setae distomesally. Gonarcus with broad, flat bridge (posterior view), arms broad, rounded throughout, with digitiform process extending forward and mesally almost perpendicularly from arm (lateral view); mediuncus long, narrow, straight, with base containing two elongate rods; gonosaccus bilobed, with each lobe bearing single, discrete patch of large gonosetae well separated from opposite lobe, with gonosetae probably facing mesally when unexpanded; gonosetae arising from enlarged setal bases, interior ones slightly smaller than distal ones; hypandrium internum narrow, U-shaped, with hooked comes (Fig.
Among the specimens we examined, there was variation in the depth and size of the reddish brown head markings. The body color of most specimens had faded. Also, there was considerable variation in the amount of brown coloration on the venation of the wings – from very little (almost as in
The larvae of
ARGENTINA (north & central western to central eastern): Provinces of Buenos Aires, Catamarca, Córdoba, La Rioja, Mendoza, Rio Negro, Salta, San Juan, Tucumán.
(in addition to types above). Argentina.
(Figs
This species is named in memory of Professor Sanford S. Elberg (1913-2011), University of California, Berkeley. Dean Elberg was Professor of Bacteriology and Public Health and served for 17 years as Dean of the Graduate School. He was a remarkable researcher, teacher, administrator, and friend.
A prominent black stripe on the gena that extends from the base of the eyes through the entire gena and along the lateral margin of the clypeus of
Head cream-colored, with vertex smooth, often shiny; inverted U-shaped marking on vertex reddish brown, prominent, narrowing mesally but contiguous, not extending anteriorly to area between scapes; anteromesal margin of dorsal antennal fossa pale or with reddish brown mark; area between eyes and posterior half of vertex cream-colored, unmarked; frons whitish to cream-colored, sometimes with transverse reddish band along anterior margin; other markings absent; clypeus cream-colored to tan, black laterally; gena with black stripe extending from base of eyes, along lateral margin of clypeus; tentorial pits cream-colored. Antenna with scape cream-colored to light tan, unmarked; pedicel cream-colored to tan, with distal ring of brown; flagellum cream-colored to light tan basally, becoming darker distally; maxillary palp with basal two segments pale, distal three segments dark brown to black, pale at junctions; labial palp with basal two segments pale, distal one dark brown to black.
Prothorax variable in length, green mesally, with wide, brown, lateral stripes, very thin, brown mesal stripe; transverse furrow slightly posterior to middle, reaching lateral margins; with pale, fine setae throughout. Mesothorax, metathorax yellow mesally, reddish brown laterally. Measurements: head width: 1.3 mm; ratio head width : eye width: 2.3–2.5 : 1; prothorax length: 0.6–0.7 mm; prothorax width: 1.0 mm.
Forewing, hindwing clear, without fumose areas, with slender venation, except R,
Male. Abdomen with moderately enlarged spiracles (e.g., A7: spiracle diameter ~0.13× length of sternite); T9+ectoproct short, dome-like, with dorsal invagination shallow, extending approximately one half the distance to anterior margin of T9, lateral margins of invagination straight to slightly convex; dorsal margin rounded distally, with posteroventral margin extended distally in well defined knob; ventral margin well sclerotized along entire length; callus cerci large, ovate, circumference lightly sclerotized, sclerotization contiguous with that on ventral margin of T9+ectoproct. S8+9 fused, with line of fusion not demarcated, but with distal tip well sclerotized (but not as heavily as that on
ARGENTINA (northwestern): Provinces of Jujuy, La Rioja, Tucumán.
(in addition to holotype above, all paratypes). Argentina.
Among
Head with vertex yellow, raised, smooth, shiny; inverted U-shaped mark red to reddish brown, broken anteromesally, not extending forward between antennae. Frons yellow, without marks; gena light reddish brown; maxillary palp pale, ultimate palpomere 1.5 times longer than anterior one; scape yellow with dorsomesal red mark that extends onto pedicel, base of flagellum; flagellomeres yellowish brown, with rows of black setae.
Thorax yellowish green, with distinct longitudinal cream-colored stripe mesally. Prothorax with pair of broad, dark reddish brown stripes laterally; transverse furrow in posterior region of segment, extending to lateral margins, with brown to black setae on areas marked with brown, golden setae on green areas. Mesonotum, metanotum with diffuse brown lateral marks. Measurements: head width: 1.3 mm; ratio head width : eye width: 2.3 : 1; prothorax width: 1.0 mm, length: 0.6 mm.
Forewing, hindwing with rounded tip; membrane clear, hyaline, without fumose areas; stigma slightly opaque to light brown. Forewing with longitudinal veins green, marked with brown at intersections; transverse veins dark brown to black; transverse costal, radial, gradate veins black, without suffusion. Forewing with
Abdomen yellowish green, without spots. Male. T9+
Female. See
Immatures were reared, and the three instars are described (
BRAZIL: State of Minas Gerais.
. Same data as type (1M, 1F, offspring from original collection of type series,
We have seen only one other specimen, a somewhat discolored female from Jujuy Province that resembles the type of
Head: vertex smooth, slightly depressed mesally; inverted U-shaped marking on vertex reddish, large, probably joined mesally, not extending between scapes; anteromesal margin of dorsal antennal fossa dark reddish brown; area between eyes and posterior half of vertex probably reddish; frons cream-colored with broad reddish brown coloration along anterior margin; clypeus cream-colored to tan, reddish brown laterally; gena, posterolateral margin of the clypeus with dark brown stripe; tentorial pits with reddish brown marking. Antenna with scape cream-colored to light tan, perhaps with some reddish coloration dorsally; pedicel cream-colored to tan, without distal ring; flagellum cream-colored basally, becoming darker distally; maxillary palp with basal two segments pale, distal segments brown, articulations pale; labial palp with basal segment pale, distal two segments brown.
Prothorax wide, probably with pair of broad, brown, lateral stripes, thin, brown mesal stripe; transverse furrow in mesal region, not reaching lateral margins; with short to long, mostly pale setae throughout. Mesothorax, metathorax greenish with yellow mesally. Measurements: head width, 1.4 mm; ratio head width : eye width, 2.5 : 1; prothoracic length, 0.8 mm; prothoracic width, 1.1 mm.
Forewing, hindwing clear, hyaline, without fumose areas, with slender venation; stigma lightly opaque, with five brown to dark brown subcostal crossveins below stigma; most veins of forewing cream-colored to light green, basal subcostal crossvein, base of
Male. Unknown.
Female. See Fig.
Wings of three
The female specimen from Jujuy, Argentina, is similar in size to
ARGENTINA: Provinces of Tucumán and possibly Jujuy.
(in addition to the type): Argentina. Jujuy, no locality, 20/II/1955, Bought F. H. Walz, Phillip A. Adams Collection 1998 bequest to Calif. Acad. Sci. (F,
The following species, at one time or another, were considered to be synonyms of
Based on name changes listed above, we conclude that a lectotype should be designated and the taxonomic status of the lectotype should be substantiated with supporting evidence.
We have not seen types or identified specimens of the following two species. The first one was assigned to
Type labels from
Type labels from
First, we acknowledge the work of Phillip A. Adams (deceased); his extremely rich collection of
We are especially grateful to the curators and museum staff members who graciously helped us borrow or study specimens in their care; they are listed at the beginning of the article in the Materials and Methods section. In addition, the following Neuropterists made special efforts to gather
We thank John D. Oswald (TAMU) for developing and providing access to the
Over the years, our systematic work on the