Revision of the Neotropical green lacewing genus Ungla (Neuroptera, Chrysopidae)

Abstract Here, Ungla Navás, 1914, a poorly known Neotropical genus is reviewed. Twenty-five valid species are recognized; seven of them are new to science: Ungla adamsi sp. n., U. elbergi sp. n., U. grandispiracula sp. n., U. mexicana sp. n., U. pennyi sp. n., U. quchapampa sp. n., U. stangei sp. n.; and five are transferred to Ungla from other genera: U. bolivari (Banks), U. chacranella (Banks), U. siderocephala (Navás), U. steinbachi (Navás), and U. banksi Tauber, new replacement name. In addition, ten new synonymies are identified. For each species, a full nomenclatural history, diagnosis, description or redescription with images, literature citations, and available information on the distribution and biology are provided. Name-bearing types were examined for each species, and images of most are included. Keys based on external features are provided for species identifications. As a result of this study, three generalizations appear: (1) The genital morphology of both males and females of Ungla species is very conserved. All species express a common structural pattern, the components of which vary only slightly among species. (2) Ungla species appear to fall into two geographically distinct groups: about one third (n=7) of the species are recorded from southern South America (specifically Argentina and Brazil) and the other approximately two thirds of the species (n=18) from more northern regions of Neotropical America [Andean and Caribbean regions, Central America, and southern Mexico (Chiapas)]. None of the species from either of the regions is known to overlap into the other region. (3) Available information on the immature stages and natural history of species in Ungla is meagre.


Introduction
The genus Ungla Navás, 1914 (Neuroptera: Chrysopidae: Chrysopinae: Chrysopini) is an intriguing and yet neglected group of Neotropical green lacewings. It was described for a single species, Ungla annulata Navás, and for ~85 years, the number of species assigned to the genus grew very slowly; by 1998 it included only seven species. All were from southern and western South America (Argentina, Colombia, Peru). Subsequently, two additional Ungla species were reported -one from Costa Rica and another from Brazil (Penny 1998, Freitas 2007. And very recently, seven new species were described from Venezuela (Sosa 2015). As a result of this relatively recent work, the genus Ungla now ranks as a diverse, broad-ranging, and important component of the Neotropical chrysopid fauna.
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 Ungla species were described on the basis of external features (e.g., coloration, venational differences) that either fade with age and/or that show significant individual variation.
(2) Subsequent studies that examined genital characteristics demonstrated very little interspecific variation, especially among females.
(3) Many of the type specimens are females and their coloration is faded; thus they were not readily distinguished from other species. (4) The type specimens of currently described Ungla species, as well as those of species that should be transferred to Ungla, are dispersed among museums in the New and Old Worlds (including South America); they have received little comparative study. Thus, when we initiated this revision we recognized that we were faced with serious constraints and that our goals should be appropriately targeted.
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 Sosa (2015), but for these species, reference is made to the earlier work to avoid redundancy. Because female genital characteristics show only subtle interspecific variation, in the interest of time, we did not include them. Finally, the results presented below are largely meant as an aid in identifying species and as a springboard for future studies. The species-delineations are our current interpretations; they are proposed for future studies to challenge and correct.

Materials and methods
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.

Notes of Phillip A. Adams
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 (CAS).
At the time during which most of the notes were written, Adams considered the species that now are included in Ungla to belong in the Old World genus Suarius. In all cases, unless specifically noted otherwise, we verified his observations by our own (CAT) examination of the types.

Lectotype designations
The Ungla species that were described before Penny (1998), as well as all of the species later transferred to Ungla were described by either L. Navás (Spain) or N. Banks (USA). Each of these authors had a distinct manner of treating types. Navás often described species on the basis of several specimens, and he often labeled several of these specimens as "Typus" without mentioning in the description how many specimens he had or which one he considered to be the primary type. In some cases, subsequent authors (e.g., Stange 1967, González Olazo 1997 considered single known types of Navás as holotypes. However here, consistent with our previous treatments of his types (e.g., Legrand et al. 2008, Tauber et al. 2017, unless Navás specifically mentioned having a single specimen, we assumed that he had others. Thus, to be in accord with Article 74.6 of the International Code of Zoological Nomenclature, rather than identifying holotypes, we designated lectotypes.
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.

Measurements, terminology, abbreviations
We made selected measurements on the wing and head. Our techniques are illustrated in Figs 1-3.
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, Sc = Subcosta, R = Radius, Rs = Radial sector, M = Media, Psm  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).
Taxonomic placement. Subfamily -Chrysopinae; tribe -Chrysopini. Based on male morphological characteristics, the genus has been suggested as related to Kostka and Chrysopidia (Brooks 1997: 273). It has not yet been included in published phylogenetic studies using molecular methods.

Taxonomic history
Originally Navás (1914) established the genus Ungla based on a single specimen that had an extremely unusual type of foreleg -one that he had not seen on a chrysopid previously. He stated: "The peculiar form of the anterior leg with long and thin tibial spurs and claws, a thing which is not seen in any other genus of chrysopids; the same with the elongate tarsus, is characteristic of this new genus" (translation by Adams 1975). He described the specimen with the unusual foreleg as Ungla annulata Navás, 1914 and designated that species as the type species of the genus. Navás (1914) did not mention the origin of the genus name "Ungla". Because the unusual leg on the type specimen had long spurs and claws, we suspect that it probably originated from the Catalan word "ungla", which means nail or claw (information provided by R. Pantaleoni).
Ungla persisted as a monotypic genus until the mid-1970s when Adams (1975) demonstrated that the leg that Navás described actually belonged to another insect (an antlion). As a result of his discovery, Adams designated the genus name as invalid; in the same paper he also considered the type species as synonymous with another Argentinian species described earlier (Hypochrysa argentina Navás, 1911). He transferred that species to the Old World genus Suarius Navás, 1914, and for many years he continued to identify Neotropical species that express similar features as New World representatives of Suarius.
Later, Brooks and Barnard (1990) re-evaluated Adams' generic determinations, and they decided that the New World species were distinct; as a result, they reinstated Ungla as a valid genus [for nomenclatural specifics, see Oswald and Penny (1991: 59)]. They described distinctive diagnostic characteristics for the genus and listed four species and one synonym [they missed one synonym previously proposed by Adams (1975)]. During the fifteen years that followed the publication of Brooks and Barnard (1990), three species previously assigned to Chrysopa were moved to Ungla (González Olazo 1996), and one new species was described (Penny 1998). Thus, when work on this study began (~2006), the genus contained eight recognized species. During the course of the study, Freitas (2007) described a species from Brazil, two species previously assigned to Chrys-opa were moved to Ungla (Legrand et al. 2008), and seven species were described from Venezuela (Sosa 2015). Now, at the conclusion of our study here, the genus contains 25 described species that we consider to be valid biological entities and 15 new synonymies.

Generic characteristics
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 [Chrysoperla, Pseudomallada, Meleoma, others; see Brooks and Barnard (1990)]. The larvae of two species have been described, but generic level features have not yet been identified.
Externally, Ungla adults generally express the following traits: adult body color green to brown, anterior wing length 8-18 mm; head with vertex raised, up-folded posteriorly, surface smooth, without striations, usually with red, brown, or black, inverted Ushaped mark; mark sometimes divided mesally or reduced to a pair of spots; gena usually marked with black, brown, or occasionally red, rarely unmarked; area between antennae sometimes marked; scape broad, strongly curved mesally; palpi flattened apically, usually marked with black, pale at joints; maxillary palpi usually with basal two palpomeres pale, distal three black or marked with black; labial palpi usually with basal palpomere pale, distal two black; cervix usually with dark lateral mark covered by pronotum; pronotum usually with longitudinal, reddish or brownish lateral band, usually bearing long pale setae, sometimes short, black setae; mesonotum, metanotum marked or unmarked; legs usually unmarked, with short pale to black hairs, claws without dilatation at the base. Forewing rounded to acute, without marks but sometimes with brown suffusion around crossveins and gradate veins and in pterostigma, first gradate vein often not meeting Psm, intramedian cell usually ovate, rarely quadrate, with three intracubital cells, distal one open. Hindwing narrow, usually without marks, usually acute, occasionally rounded, with two intracubital cells; venation usually green, occasionally with some veins dark.
Female genitalia. Spermatheca round, pillbox-shaped, with or without identifiable invagination; velum tall, opening directly to bursa via elongate dorsal slit or opening on the side; bursa unenlarged, simple, without obvious glands; spermathecal duct of moderate length, with two to three curves, distal end with setae-like, filamentous ducts; praegenitale absent; subgenitale with base broad, weakly to well sclerotized, with protrusion short to slightly elongated, bilobed distally, with simple, broad pocket or lobe below, sometimes invaginated; colleterial complex delicate, with elongate gland, small reservoir; transverse sclerite somewhat broad, slightly bowed to flat, with elongate filamentous projections. Adams (1975) reported "slender dorsal bursal glands" on female Ungla argentina (as Suarius), but in his notes for this species, he states that there are "no bursal glands" and none are shown in his sketch. We have not found any on this species or other Ungla species.

Geographic distribution
Ungla, which now contains 25 valid species, is known to occur from southern Mexico (Chiapas) and Central America (Costa Rica) throughout most of South America. It is not yet known from the Caribbean islands or Chile. And, it appears most diverse in northern South America (Venezuela, Ecuador, Peru, Bolivia); however, this pattern may be the result of bias in collecting. For example, the fauna of Argentina and parts of Brazil has not been explored well and we suspect that undescribed Ungla species occur there.
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 Ungla species we studied will persist when additional specimens become available. Thus, we encourage readers to refer to both sections below when identifying or otherwise studying Ungla species.

6'
Dorsal surface of scape tinged with red, but without distinct stripe (Fig. 4e, f); female with frontal marking extended transversely into an inverted V-shape (Fig. 4b) Etymology. The species is named for Phillip A. Adams, an intense and talented contributor to systematics and evolutionary studies of Neuroptera. His publications, collection, and unpublished work on the lacewings of Latin America were of great help to this study, especially during the early stages. Indeed, Adams identified this species as new, thus the name "adamsi". From his notes, it is clear that he, as did we, often found working with this genus to be perplexing.
Diagnosis. Ungla adamsi, currently known only from the Andean archeological site of Machu Picchu, Peru, has several distinguishing features. Most notably, the males and females are strongly dimorphic in head markings and frontal modifications. In males the frons is enlarged and inflated above the clypeus, and it is marked with a prominent dark, central spot. Females also have a dark frontal spot, but in the female, the spot extends transversely towards the distal margin of the frons, forming a thick, inverted "V-shaped" mark. The female frons is not enlarged.  Tauber Ungla adamsi resembles three other Ungla species that have frontal spots and cream-colored flagella -U. stangei sp. n., U. favrei, and U. nigromaculifrons. However, none of these species express sexual dimorphism in the frons. Ungla adamsi also differs in that it lacks the features that distinguish the other three species. That is, it has diffuse reddish marks on the scape, not a distinct stripe like the one that typifies U. favrei. It has moderately sized forewings (< 15 mm), with 7-8 gradate veins per series, not the very large wings that distinguish U. stangei; and the veins on its hindwings are green, unlike the very dark transverse veins on the fore and hindwings of U. nigromaculifrons. Description. 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.
tral margin of T9+ect, with three arms projecting dorsally (mature): first arm extending along posterior margin of segment, second arm contiguous with sclerotization around callus cerci, third arm extending dorsally distal to callus cerci; callus cerci large, slightly ovate, with discrete, well separated trichobothria. S8+9 fused, with line of fusion not detectible; ventral apodeme extending along dorsal margin for full length of A8; dor- sum of S8+9 tapering gradually, then forming concave ledge at terminus; terminus blunt (lateral view), extending slightly beyond distal margin of T9+ect; setae slender, mostly long, simple, those along distolateral margin large, flanged. Subanal plate fairly large, triangular in shape, with ~9 setae of medium length. Gonarcus broadly arcuate, rounded mesally, with bridge relatively slender, arms elongate, narrow throughout (lateral view), extending straight downward from gonarcal bridge, margin rounded distally, mesal section with angular enlargement, distinct digitiform process extending posteriorly, inward; mediuncus with quadrate base closely attached to gonarcal bridge, narrow distally, with terminus spoon-shaped, slightly curved downward throughout (lateral view); gonosaccus bilobed, with lobes closely aligned, when unexpanded forming a triangular envelope around tip of mediuncus, when expanded consisting of two lobes, rounded dorsally, mesally, laterally, but flat distally; each lobe with large, dense patch of gonosetae arising from prominent setal bases facing inward when uneverted, outward when everted; gonosetae moderate length (shorter than length of mediuncus); hypandrium internum small, with two short, dense arms, with narrow, rounded junction.
Known distribution. PERU: Department of Cuzco.
The species is a valid biological entity within the genus Ungla; its original name (Nothochrysa tibialis Banks, 1914) was preoccupied . Because the name is a primary homonym, Articles 57.2 and 60.1 of the International Code of Zoological Nomenclature require the designation of a replacement name. In recognition of Nathan Banks' discovery and original description of the species, we offer the name Ungla banksi as the replacement.
Diagnosis. 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.
Redescription. Head cream-colored, with vertex smooth, shiny, with inverted Ushaped marking prominent, dark brown, broad, separated anteromesally, not extending anteriorly to area between scapes; dorsal antennal fossa pale; area between eyes    Tauber 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.
Variation. 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.
Known distribution. BOLIVIA: Departments of La Paz, Santa Cruz. Specimens examined (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,FSCA). (Banks, 1913), comb. n.  . Although the abdomen is missing, the frontal horn and crassate wing venation are typical of a male. The head and body are flattened; thus many structures could not be observed and/or measured accurately. It seems unlikely that Banks had specimens other than this one when he described the species; it is labeled as the type in his hand. Thus, we consider it to be the holotype by original designation. San Antonio, the type locality, is a small city in Tolima Department, Colombia (altitude, ~1500 m). Support for generic placement. Currently, this species is known only from the holotype, which is lacking its abdomen. Here, we base our generic identification on several external features that characterize Ungla species: vertex with pair of large, dark marks, distal segments of the maxillary and labial palpi dark brown, and a basal inner gradate that does not meet the Psm.

Ungla bolivari
Diagnosis. Based on the type specimen, it appears that U. bolivari can be distinguished from other Ungla species by the following suite of external features: small, rounded wings, with tan to light brown longitudinal veins, darker brown transverse veins; several basal veins crassate; large, shiny, dark brown marks on the vertex; frons with single, brown mesal spot and with a small mesal lobe protruding below the scapes (probably males only); clypeal region enlarged (boxy) frontally; antenna with flagellum and pedicel dark brown, scape dark brown dorsally. Redescription (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.
Male and female. Abdominal characteristics unknown. Known distribution. COLOMBIA: Department of Tolima. Specimens studied. Holotype only. Also see below.
Possible sibling species from Venezuela. The characteristics of U. bolivari are known with certainty only for the type specimen from Colombia. However, one female and two male specimens from Venezuela that resemble the U. bolivari holotype were studied. These specimens express many features that characterize U. bolivari (e.g., dark basal flagellomeres and dark brown to black markings on the vertex, gena, and lateral margins of clypeus). However, several features of the wings (size, number of gradate veins) lead us to believe that they are not conspecific with U. bolivari. Thus, we are not naming it here. However, to facilitate comparison with future specimens, here we include a description and images of the Venezuelan specimens.
Diagnosis. 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 Ungla is reduced to small red markings on and around the vertex and fossae. The forewing has pale green veins; however, the gradate veins, which are parallel, are reddish brown to brown, with light suffusion of brown on the surrounding membrane. In males, most of the longitudinal veins of the forewing are crassate, and the abdominal spiracles are enlarged, but the dorsomesal margins of the abdominal sternites are only slightly indented. [Note: On many pinned specimens, the head markings are faded or discolored and the costal and radial crossveins are either partially or completely darkened to light brown (compare Redescription. 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.
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+ect, with distal setae slightly enlarged, with row of heavy, flanged setae along dorsodistal margin. Gonarcus flattened, narrow in lateral view, thick in frontal, posterior views; bridge straight for short distance mesally, then arms extend-ing downward at sharp angle; arms very slender in lateral view, broad, with rounded tips in posterior, frontal views, with digitiform process extending forward, not inward from distal margin of apodeme; process with wide base, tapering to acute tip distally; mediuncus long, narrow, straight dorsally (lateral view), rounded, with broad hook distally; bilobed gonosaccus, each lobe with single, large patch of heavy gonosetae facing mesally when unexpanded; dorsal setae smaller than ventral ones, all arising from bulbous setal bases; hypandrium internum attached closely to base of gonosaccus, robust, broadly V-shaped, with lightly sclerotized, hooked comes.
Variation. The coloration of the body, markings, and wing venation deteriorates quickly and substantially after death. Diagnosis. Ungla demarmelsi is the only known Ungla species that has the following set of features: a yellow head with raised vertex bearing diffuse to dark reddish, inverted U-shaped mark, scape with red mark dorsoapically, pronotum with broad, red, lateral stripes, and mesoscutum marked with red laterally. Males have enlarged spiracles, and the dorsal margins of the sternites are clearly concave in both sexes. The subanal plate is small; it bears about ten rather small setae. The tip of the male S8+9 is concave and extends beyond T9+ect; its terminus bears enlarged setae on robust bases; approximately ten pairs of these setae are also flanged basally above the setal base. The gonarcal bridge is round and smoothly arched throughout. Redescription. 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.
Female. See Fig. 30 here, and Sosa (2015). Variation. The darkness and size of the head markings on the vertex vary among specimens. Diagnosis. Ungla diazi is typified by (i) a yellow flagellum marked with black ventrolaterally, (ii) scape unmarked dorsally, except sometimes with a small to large mark distolaterally on the ventral surface, and (iii) gena cream-colored to golden below the eyes, with a black spot on the pleurostomal margin. The spiracles on the male abdomen are not enlarged, and sternites S4-S6 are densely covered with microsetae. 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 U. banksi and U. quchapampa. Hand-written labels on some specimens of U. diazi from Bolivia indicate that Adams too thought the species was related to U. banksi (then known as Nothochrysa tibialis). In addition to the traits above, U. diazi adults can be distinguished from U. banksi by their more elongate wings and lighter brown veins, largely lacking suffusion. They can be differentiated from U. quchapampa by their lack of a large marking on the frons.
Redescription. 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.
Variation. 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.  Banks referred to a single specimen; thus it is the holotype. We could not find a "Rio Ognacatal" in South America; we assume that the term refers to Río Aguacatal, a stream in Valle del Cauca Department, Colombia (located between El Centenario and Barrio Terrón Colorado. Rio Aguacatal is in the Western Cordillera of the Andes at an elevation of approximately 2,000 meters. Support for synonymy. The types of both species are females and discolored with age. A large series of specimens (CAS, see below) collected in Cochabamba Department, Bolivia, allowed us to associate males and females of the species and to identify diagnostic external and male genitalic characters that confirm the synonymy.
Diagnosis. This species is distinguished from other Andean Ungla species by a brown, inverted U-shaped mark that is broken mesally, a white to cream-colored face, frons usually with brown mesal spot of variable size and darkness, antenna cream-colored, with longitudinal brown mark on the distal, upper surface of the scape that extends onto the pedicel, and wings with pale longitudinal veins and numerous brown crossveins. The male abdomen has moderately enlarged spiracles and dense setation; the gonarcal bridge has a ledge that extends forward and receives the base of the mediuncus. Among all Ungla species, U. favrei appears most similar to U. elbergi sp. n., both externally and in some male abdominal features (e.g., the moderately enlarged spiracles (A7: ~0.15× length of sternite). However, its male abdominal segments are taller in height and shorter in length than those of U. elbergi, and its genital structures (gonarcus, mediuncus and gonosaccus) differ in shape. It also resembles U. grandispiracula sp. n., which can be differentiated by its larger spiracles (A7: 0.25× length of sternite), more robust gonosetae, gonarcal bridge without an enlarged mesal platform, and scape with brown dorsal mark that does not reach the pedicel.
Redescription. 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.
Variation. 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.     Tauber Etymology. The species name "grandispiracula" (Latin, neuter, plural) refers to the large spiracles that distinguish males of this species from those of U. favrei, another Andean species of Ungla with which it shares many features. The word is a compound noun in apposition to the genus name (grandis, meaning "large"; spiracula, meaning "spiracles").

Ungla grandispiracula
Diagnosis. The Andean species U. grandispiracula and U. favrei are very similar externally and in many of their male abdominal features. Both have a brown or red, inverted U-shaped mark that is broken mesally, a white to cream-colored face, creamcolored antenna with longitudinal brown mark on the distal, upper surface of the scape that extends onto the pedicel, and wings with pale longitudinal veins and numerous brown crossveins. They also have enlarged abdominal spiracles and similar genitalia. However, there are subtle differences in the male abdomen that distiguish the two species: the U. grandispiracula spiracles are larger (A7: 0.25x length of S7 versus 0.15x in U. favrei), its gonosetae are more robust, and its gonarcal bridge is narrow, uniformly rounded, and it lacks the mesal ledge that occurs in U. favrei. Females of the two species are difficult to separate; in U. grandispiracula the frons is unmarked (variable in U. favrei) and the stripe on the dorsal surface of the scape does not extend onto the pedicel as it does in U. favrei.
Externally, this species also resembles the Argentinian U. elbergi, sp. n. However, the U. grandispiracula spiracles are larger than those of U. elbergi; and, unlike on U. elbergi, the two lobes of the gonosaccus are well separated mesally, and the gonosetae are borne laterally, on somewhat flattened plates.
Description. 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 be- tween 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.
Known  Legrand et al. 2008: 149). The type locality is an unspecified location in Colombia. Chrysopa aroguesina . Rev. chil. Hist. nat. ("1928  small, crescent-shaped marks on the vertex (in place of the inverted U-shaped mark that typifies most Ungla species). The wings of both species have slightly acute tips; the longitudinal and transverse veins are largely yellow to light green; most transverse veins are pale mesally with small brown marks at both ends where they intersect with longitudinal veins. The gradate veins, the first two r-m crossveins, and the last crossvein of the distal b' cell are entirely brown and are bordered with brown suffusion. Adults of U. laufferi can be distinguished from U. stangei by their smaller wing size (forewings = 13.5-15.9 mm, 11-13 radial cells, 6-7 inner gradate veins, 6-8 outer gradates), and 6-7 gradate cells that are long and narrow. Also, in the U. laufferi male, the S8+9 is elongate and its dorsal surface tapers evenly throughout the entire segment; the gonarcus is arcuate and has elongate gonarcal arms that extend downward from the gonarcal bridge (smoothly or slightly angled); they do not recurve markedly below the gonarcal bridge and gonosaccus. Redescription. 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 Rs); third gradate cell 1.9-2.2 mm long, 0.4-0.5 mm wide (ratio, L : W = 3.8-5.0 : 1); fourth gradate cell 1.9-2.4 mm  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+ect), margins of invagination rounded throughout; dorsal margin of T9+ect rounded distally (above anus), often compressed, thus ap- pearing straight (especially in teneral specimens); posterior margin of ectoproct relatively straight, posteroventral corner slightly extended distally, without knob; dorsal apodeme along ventral margin sinuous, with arms extending dorsally on each side of callus cerci; callus cerci large, ovate, margin unsclerotized at top. S8+9 fused, with line of fusion perceptible; dorsum tapering gently throughout, with margin slightly irregular; ventral apodeme on margin of S8 lightly, irregularly sclerotized, ventral apodeme on margin of S9 more densely sclerotized; terminus extended distally, but not beyond T9+ect, flat, distal margin upturned, plate-like, well sclerotized (setae mostly missing). Gonarcus U-shaped or slightly acute (probably teneral), with bridge slender, apodemes elongate, extending below bridge then bending posteriorly, with quadrate unarticulated process mesally; mediuncus relatively broad-based, with pair of straight, sclerotized rods, extending from base along sides of mediuncus, coalescing distally in rounded, down-turned beak; gonosaccus bilobed, each lobe with single, relatively sparse patch of gonosetae arising from enlarged setal bases, lobes evertible (setae extended) or withdrawn (patches of setae opposite each other); hypandrium internum V-shaped, with flat base, mature specimens: arms expanded, scalloped, comes hook-shaped.   Variation. 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.   (2015); for labels, see Fig. 143e here. Support for synonymy. The original description of U. rubricosa was based on a single teneral male and two female specimens from Venezuela (Sosa 2015). Additional Venezuelan specimens used in the study here, allowed us to reexamine the pattern of variation among a larger group of specimens and to establish that the two species indeed are the same.
Diagnosis (also see Sosa 2015). This phenotypically variable species is recognized by its green body, marked with deep red; light yellow to cream-colored head; U-shaped marking on vertex either broken mesally or connected via thin mesal line; margins of head adjacent to eyes marked with red band; scape and pedicel with red dorsal stripe; flagellum cream-colored; frons unmarked; palpi marked with black. The wing venation, with dark brown transverse veins and fumose R-Rs crossveins and gradates, and the male genitalia are also distinctive.
Ungla martinsi resembles another Venezuelan species U. curimaguensis, in size, male abdominal and genital characteristics, and perhaps head markings. However, the two species differ notably in that U. curimaguensis has largely green forewing venation and it lacks the dark genal marks and marked maxillary and labial palpi of U. martinsi.
Redescription. 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; clyp eus 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.
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.
Etymology. The type locality of this species is in the highlands of southern Mexico (State of Chiapas). Its species name, mexicana (Latin, adj., fem.), recognizes that it is the first, and at this time the only, species of Ungla reported from Mexico.
Diagnosis. The female holotype expresses many of the characteristics that typify most Ungla species: a U-shaped marking around the margins of the vertex, dark genal markings, dark [albeit, thin] markings on the palpi, and a typical doughnut shaped spermatheca.
Ungla mexicana is recognized by its green body with reddish markings; prothorax with a distinctive red lateral band that extends only about one half the length of the segment (from the anterior margin to the midlength of the segment), and mesothorax with a pair of bold black markings anteriorly. The U. mexicana forewing venation is green, with most longitudinal and transverse veins marked with dark brown at intersections; crossveins are dark brown, with the gradate veins having brown suffusion on the adjacent membrane. This species most closely resembles U. martinsi; however, the two differ as follows. In addition to the above features, the U-shaped marking on the U. mexicana vertex is much larger, the scape is without a distinct dorsal stripe, and the distal three maxillary palpomeres have a thin lateral stripe rather than being black throughout.
Description. 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.
Male  (2015); for labels, see Fig. 143f here. Much of the abdomen of the holotype is lost; the original description illustrates the structures that are present: T9+ect (lateral), S8+9 (ventral) and parts of the gonarcus.
Diagnosis. 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 (U. stangei, U. favrei, and U. adamsi) in that the scape is tinged with red, but otherwise unmarked (distinct stripe in U. favrei), the wings are smaller and with at most 4 to 8 gradate veins per series (8 to 11 for U. stangei), the clypeus and frons do not exhibit sexual dimorphism (male clypeus swollen and female frons with transverse marking in U. adamsi), and finally the hindwing has dark veins (unlike all the other species). Redescription. Head: As described by Sosa (2015). Maxillary palp with basal two segments cream-colored, distal three segments entirely black, intersections creamcolored; labial palp with basal segment pale, distal two entirely black. Thorax: Prothorax with transverse furrow in the posterior section of segment, marked with narrow, dark red transverse stripe extending almost to center of segment; setae long, golden. Mesonotum, metanotum green mesally, with pair of broad, dark red stripes laterally. Measurements: head width: 1.5 mm; ratio head width : eye width: 2.1-2.2: 1; prothorax width: 0.8-1.1 mm, length: 0.6-1.0 mm.
Female. See Sosa (2015). In the original description of U. nigromaculifrons, a specimen from Lara State in Venezuela was reported as a paratype (Sosa: 2015); we now believe that this specimen was misidentified. It is a species that resembles U. bolivari (see above). Penny, 1998 Figs 68-70, 143h, 143i Ungla pallescens Penny, 1998 Fig. 143h, i). Originally, this species was described from a series of specimens collected in Costa Rica by N. D. Penny; these remain the only specimens thus far reported for this species.
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+ect), margins of invagination almost straight, base rounded; dorsal margin of T9+ect convex, curving downward, melding into descending distal margin; posterior margin of ectoproct slightly rounded, tip subacute, without protruding knob; ventral margin of T9+ect convex, with distinct, curved apodeme extend- ing along entire length; posterior margin of T9+ect straight; callus cerci large, ovate, margin sclerotized throughout; sclerotization contiguous with that on ventral margin of ectoproct. S8+9 fused, with line of fusion apparent; dorsal margin (lateral view) without apodeme, with very slight taper, small hump, then stronger taper distally; terminus rounded (lateral view), slightly convex (ventral view), not extending much beyond T9+ect; terminal setae long, some along upper lateral edge of S8+9 with small flanges, otherwise simple. Gonarcus (teneral) arcuate, U-shaped, with bridge slender, curved throughout, arms elongate, extending ventrobasally from gonarcal bridge, rounded distally, mesal section with elongate digitiform process extending posteromesally; mediuncus broad basally, tapering to straight, rounded projection, Note 1: The above male specimen from Peru expresses almost all of the characteristics that distinguish U. pallescens (gena unmarked, distal palpomeres dark, abdominal spiracles small, venation pale and genital characteristics). The one exception is that the markings on the vertex are small and spot-like. Unfortunately, the wings are broken at about midlength and the genitalia are teneral (as they are on the male paratype of U. pallescens that we studied). Thus, at this time we consider our identification of this specimen (and the range extension for the species) as tentative.
Diagnosis. Ungla pennyi is one of three Ungla species reported from an area north of South America; the other two species are U. pallescens from Puntarenas, Costa Rica, and U. mexicana, sp. n., described above, from Chiapas, Mexico. Ungla pennyi differs from these two species in its smaller size (forewing length = 10.9 mm, vs 13.1-14.1 mm for U. mexicana and U. pallescens), and cream-colored, unmarked palpi. Also, in contrast to U. pallescens, the male U. pennyi has enlarged abdominal spiracles. [The male of U. mexicana is unknown.].
The U. pennyi holotype closely resembles specimens of the Venezuelan species U. curimaguensis and U. martinsi, in size and especially in the male abdominal and genital characteristics. Thus, it is possible that our specimen represents a differentiated population of one of these two species. However, it lacks the dark genal marks and marked maxillary and labial palpi of both species, and the largely green forewing venation of U. curimaguensis. In U. curimaguensis, the C-Sc crossveins, R-Rs crossveins and the transverse branches from the Rs are entirely green, whereas in U. pennyi and in U. martinsi they are dark brown with surrounding membrane suffused with brown coloration. At this time, we consider that the set of features expressed by the U. pennyi specimen indicate species-level differentiation; we await confirmation or correction of our opinion when additional specimens from Central America become available.
Description. 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.
Known distribution. COSTA RICA: Provinces of Cartago, Puntarenas. Specimens examined (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, CAS). Tauber Etymology. 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") (Ajacopa 2007). We use the name "quchapampa" as a compound, proper noun (Latinized, fem.), in apposition to the genus name.

Ungla quchapampa
Diagnosis. Ungla quchapampa is one of several species that has a prominent dark mark in the center of the frons, but it is the only one that has a dark flagellum and also lacks a prominent protuberance on the frons. The size and shape of the marks on the vertex are also distinctive: they are very large, dark brown, contiguous with each other, and extend anteriorly to the mesal margins of the antennal base. In more than half of the specimens we examined, the first intramedian cell (im1) of the forewing was quadrate (unlike the commonly triangular im1 cells of other Ungla species).
Ungla quchapampa closely resembles U. diazi in many features -notably size, wing features, and male abdominal characters (external and internal). In addition, the two species are known to occur sympatrically; however, U. quchapampa can be differentiated by its prominent frontal marking, sometimes quadrate im1 cell in the forewing, and the absence of markings on the ventral surface of the scape.
Description. 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 creamcolored 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.
Known distribution. BOLIVIA (central): Department of Cochabamba (Provinces of Carrasco and Chapare).

Figs 81-87
Chrysopa siderocephala Navás, 1933. Rev. R. Acad. Ciencias exactas fis. Nat. Madrid (1933a 30: 306-307; "Perú: Lima, 4.X.1932". Penny 1977 (Figs 83, 84). MZBS, male. Specimen discolored, with abdomen in poor condition. Confusion concerning the type is unlikely; however, to assure stability, here the specimen in the MZBS is designated as the lectotype (des. CAT). Support for synonymy. The types of both C. siderocephala and C. lambda were collected at the same locality and on the same day, and in his original description, Navás (1933a) noted the similarity between the two. However, the notable variation in the shape of the frontal markings (from an inverted V to an inverted Y) caused Navás to believe that he had more than one species. Additional specimens are now available. Phillip Adams recognized the synonymy and labeled the C. lambda specimen as the junior synonym, but he did not publish his conclusion.
Here, we formalize the synonymy. The main structural difference that we noticed between the two specimens is in the intersection of the basal inner gradate with the Psm: on both wings of the C. lambda type the basal inner gradate veins meet the Psm, whereas on the C. siderocephala type they do not. We attribute this difference to intraspecific variation. Otherwise, the venation of the two specimens is almost identical, and the size of the wings is very similar: C. lambda, forewing 12.2-12.3 mm long, 4.2 mm wide, hindwing 11.0-11.1 mm long, 3.3-3.4 mm wide; C. siderocephala, forewing 12.3-12.4 mm long, 4.3-4.4 mm wide, hindwing 11.1-11.3 mm long, 3.3-3.6 mm wide.
Diagnosis. 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 82, 84, 87. Redescription. 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.
Variation. Among the specimens we examined, there was variation in the depth and size of the reddish brown markings on the vertex and frons. Holotype (Figs 88a-e, 90d, 92d, 144f ). CAS, male. Bolivia, La Paz, Rio Zongo, 3200 m, 24-30/XI/1984, L. E. Pena. Etymology. The species is named for Lionel A. Stange, a life-long student of the systematics and natural history of myrmeleontoid Neuroptera, an enthusiastic, generous collector of Neuropteran specimens, colleague, and friend.
Diagnosis. Like U. laufferi (above), adults of this species are recognized by their light green body color, yellow longitudinal stripe dorsally, and distinctive pair of red spots on the vertex in place of the inverted U-shaped mark that typifies most Ungla species. Despite the similarities of the two species, U. stangei is distinguished by several notable external features: (1) larger wings (forewing length = 17.1-17.6 mm, versus 13.5-15.9 mm in U. laufferi), (2) wings with a greater number of gradate cells, all of which are long and narrow (n = 9-10 gradate cells; third gradate cell 2.23-2.74 mm long; fourth gradate cell 2.39-2.74 mm long), versus a smaller number of gradate cells, all of which are shorter and wider in U. laufferi (n = 6-7 gradate cells; third gradate cell  1.88-2.22 mm long; fourth gradate cell 1.86-2.36 mm long), (3) heavy setation on the pleural region of the terminal abdominal segments (A7, A8+9) (U. stangei: dense covering of elongate setae arising from enlarged setal bases; U. laufferi: a sparse covering of long setae arising from small setal bases).
The three male specimens of U. stangei that we examined appeared somewhat teneral. In all three, the abdomen and genitalia were soft, flexible, and very lightly sclerotized; the arms of the gonarcus were elongate and flexible -extending fully around and below the gonosaccus; and the mediuncus was broad, rounded, and somewhat soft at the base. We are hesitant to ascribe taxonomic significance to these features.
Description. 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, un- marked; 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.
Known distribution. BOLIVIA: State of La Paz. ECUADOR: Province of Pichincha. PERU: Province of Andahuaylas. All localities at ~3000 meters elevation.
Variation. 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. Diagnosis. Previously, this species was known from the single (slightly teneral) female holotype (Sosa 2015). Subsequently, additional female specimens became available for comparison; unfortunately, the male remains unknown. This species resembles both U. favrei and U. diazi in that adults have a golden head, black stripe on the gena and basolateral section of the clypeus, a pair of broad, curved, brown marks on the vertex, and broad dark brown or reddish brown lateral stripes on the prothorax. However, a pale flagellum distinguishes U. yutajensis from U. diazi, and the dark crossveins and the four dark marks in the pterostigma differentiate it from U. favrei. Redescription. 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.

Part 2. Ungla species from southern South America (Argentina, southern Brazil)
The Ungla species of southern South America (Argentina, Brazil) comprise a grouping of seven valid species, none of which is known to also occur outside of the region. Six of the species (U. annulata, U. argentina, U. chacranella, U. confraterna, U. elbergi, U. ivancruzi) are small bodied (forewing length < 13.5 mm), and they resemble each other in a number of features. We suspect that at least some of them may be closely related to each other. The other species, U. steinbachi, is larger bodied (forewing length > 14.5 mm); it is known only from the type (a female). Possible relationships of U. steinbachi with other species are not apparent. Mark on vertex extending anteriorly between scapes; frons usually with pair of light brown spots mesally (Fig. 103a-c); forewings with relatively dull membrane, with prominent fumose markings surrounding gradate veins and other crossveins (Fig. 104); veins extending from radial sector to apex of wing usually marked with black and white; male abdominal spiracles not enlarged (Fig. 105a, b)  Mark on vertex extending towards scapes, but usually not between them; frons without mesal mark (Fig. 98b, c, d); forewing membrane shiny, hyaline, with fumose markings surrounding gradate veins and other crossveins often masked by reflection (Fig. 95b, e, f); veins extending from radial sector to apex of wing usually entirely black (Fig. 99c); male abdominal spiracles enlarged (Fig. 100a)  Palpomeres cream-colored, without marks (Fig. 134b); forewing with most transverse veins and crossveins brown or black (Fig. 134d); male abdominal spiracles not enlarged (Fig. 135a)  Head with prominent, black stripe on gena and clypeus, from base of eyes at least part way along lateral clypeal margin (Fig. 130d, f); male abdominal spiracles moderately enlarged (width = ~0.13x length of segment) (Fig.  132a-c)

Ungla annulata Navás, 1914 Figs 95-101
Ungla annulata Navás, 1914. Broteria (Zool.) Monserrat (1985: 240) reported a type from the MZBS; it is a female, studied by Adams (1975: 169). We (CAT) examined the specimen in 2010; to help stabilize the nomenclature of Ungla, here this specimen is designated as the lectotype (des. CAT). Type locality: The type locality "Huasán" is located in northwestern Argentina -Andalgalá, Catamarca (27°34'S, 66°19'W, ~1100 m). Cintameva lurida Navás, 1930. Rev. Chil. Hist. Nat. 34: 65-66; "República Argentina: La Paz (Dep. San Javier), Córdoba 1-20 de Enero de 1929. Col Bruch". Stange 1967Penny 1977: 19 (list, as Chrysopa); Brooks and Barnard 1990: 280 (list, as " 'Chrysopa' incertae sedis"); González Olazo 1996: 378 (catalog, as synonym of U. argentina); Oswald 2015 (catalog, as synonym of U. argentina). syn. n. Syntypes (Fig. 97). MACN. Several specimens from the type locality and the approximate collection dates (Dec. 15-31, 1928, Jan 20, 1929 are in the MACN. Any of those specimens that carry dates from Jan 1 to Jan 20 could be considered syntypes. Currently, one of the specimens carries lectotype and type labels, but none of these labels are in Navás' hand, and it is not clear which of the specimens in the MACN Navás may have used. Stange (1967: 35) noted a male and González Olazo (1996: 378) noted a female that they each considered as the holotype. Given that Navás clearly had more than one specimen, we suggest that a lectotype be designated after the full series of syntypes has been studied carefully. Support for synonymy. González Olazo (1996: 378) recognized that this species belongs in the genus Ungla, and he listed it as synonymous with U. argentina. However, it is not clear what specimens or information he considered to support his synonymy. Here, in the absence of a confirmed type specimen for C. lurida, we used Navás' description of the external features and Adams' unpublished notes and drawings from specimens in the MACN. These specimens had been collected at the type locality during the period mentioned in the original description and during the month earlier. Adams apparently considered these specimens as representative of the C. lurida type series.
Diagnosis. This species, along with U. argentina, can be differentiated from most other Ungla species by a number of features: brownish coloration, longitudinal wing veins with alternating white or cream-colored and dark brown or black, often with dark patches at intersections. It differs from U. argentina in usually expressing the following external features: vertex with distinct, inverted U-shaped marking, usually divided mesally, with anterior markings running forward, but not extending beyond scapes to the frons; frons pale (sometimes whitish), unmarked; wing membrane irridescent, wings with longitudinal and some transverse veins alternately white and black, distal ones, dark (see Figs 98, 99c, d, for comparison of distal venation with U. argentina). In the males of both species, the terminus of S8+9 has large, but simple (unflanged) setae, and the gonosaccus is bilobed, with lobes broadly connected. In U. annulata, the abdominal spiracles are greatly enlarged -a trait that clearly separates it from U. argentina. Also, the ventral apodeme of the U. annulata ectoproct extends distally beyond the terminus of the segment, a feature that does not occur in U. argentina. Redescription. Head: vertex cream-colored to tan, with dark reddish brown "Ushaped 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 Rs); third gradate cell 1.0-1.2 mm long, 0.4-0.5 mm wide (ratio, L : W = 2.6-3.2 : 1); fourth gradate cell 0.9-1.1 mm long, 0.4-0.5 mm wide (ratio, L : W = 2.2-3.2 : 1); 3-4 Banksian cells (b cells), 4 b' cells; 3-5 inner gradates, 5-6 outer gradates. Hindwing 9.1-10.4 mm long, 3.0-3.4 mm wide (ratio, L : W = 3.0-3.1 : 1), 9-10 radial cells, 3 b (Banksian) cells, 4 b' cells, 3-5 inner gradates, 4-6 outer gradates.
Variation. The U. annulata specimens that we examined varied in size and the depth of the dark markings on the head and body and the degree of white and black on the wing veins. The male abdominal and genital characteristics were consistent among specimens.
Larvae and biology. Unknown. The above specimen from La Rioja (MSNG) bears a red type label with the name "Cintameva giacamellina Nav." handwritten by Navás. Apparenly, the name was never published; it is not listed by . We identified the specimen as a female of U. annulata.
Diagnosis. The lectotype of Hypochrysa argentina expresses a set of external characters that we concluded are typical for the species: wings with membrane somewhat shiny to slightly dull, veins robust; both longitudinal veins and transverse veins marked alternately with white (or cream-color) and black (or dark brown); gradate veins dark with prominent fumose marks on surrounding membrane; dorsum of head (vertex) brown, with relatively weak, inverted U-shaped marking, with paired darker brown markings extending anteriorly from the vertex, between the scapes, onto the frons and around the lower margins of the antennal fossae. The U. argentina male abdomen has small, unmodified spiracles; the mediuncus is relatively short and slightly sinuous dorsally; gonosetae are present in a single patch on the upper and outer surfaces of each lobe of the bilobed gonosaccus; and setae at the tip of S8+9 are simple, i.e., they are not flanged. Redescription. 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.
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.
Variation. 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, CAS) that, based on abdominal features, we believe is U. argentina. However, it expresses very unusual features: extensive regions of white on the transverse veins, many R-Rs crossveins missing (especially on the hindwing), and the presence of only one series of gradate veins (the distal series) on the hindwing. This specimen remains an enigma.
Larvae and biology. Monserrat and Freitas (2005) reported on some aspects of the developmental cycle and the third instar; most notably, they stated that the third instar carries debris, and eggs are laid in clusters. Unfortunately, voucher specimens for their study are not available. Thus, we cannot confirm the species identification for the report. Interestingly, the authors did not mention enlarged spiracles on the male abdomen (thus, this character could be consistent with U. argentina). However, the report does not indicate that there were markings on the forewing longitudinal veins ing the subcostals, are green. Totally green venation is typical of U. chacranella. Second, the head and facial coloration, markings, size, etc. of the C. mendocensis type are indistinguishable from those of U. chacranella. Similarly, because of the above features, we disregarded González Olazo's label of "Ungla argentina (Navás)" on the type specimen. Chrysopa plesia Navás, 1918. Rev. Soc. Arg. Cienc. nat., 4: 86-87 ;Oswald 2015 (catalog). syn n. Lectotype (Fig. 110). MLPA, male (examined). Head missing, genitalia in a vial with glycerine. Stange (1967: 36) reported a male, and González Olazo (1996: 380) reported  a single specimen (sex unspecified). We examined the external features briefly and the abdomen in detail. The locality data are as reported in the original description, and the identification label is in Navás' hand. The specimen carries an additional identification label (Ungla argentina) by González Olazo (Fig. 110a). Here, this specimen is designated as the lectotype of C. plesia (des. CAT). Support for synonymy. Adams' (unpublished notes) stated that this specimen resembles "C. binaria" (now a synonym of U. confraterna). However, he noted several differences; for example, he considered the venation to be paler than that of "C. binaria", and he noted that the vertex of the C. plesia type has a pair of reddish marks posterolater-   ally, which are not present on the C. binaria type. It is surprising that he made no mention of the enlarged abdominal spiracles, which are prominent on the C. plesia type. All of the above features [pale wing venation, reddish marks on vertex, and large abdominal spiracles (males)] are typical of U. chacranella. Chrysopa metanotalis Navás, 1924. Rev. Chil. Hist. nat. 27: 114 ("1923 Oswald (2015, as Chrysopa). syn n. Lectotype (Figs 111, 112). MACN, male (examined). González Olazo (1996: 380) recognized the type as a species of Ungla, and he attached an "Ungla argentina" identification label to the specimen, but he did not formalize the synonymy. To help stabilize the nomenclature of the genus, here this specimen is designated as the lectotype of C. metanotalis (des. CAT). The type locality, Hualfín, is in northeastern Argentina: Belén, Catamarca (~1850 m). Support for synonymy. Unlike Navás (1923) in his original description, we did not observe dark cross veins on the wings; they were pale as in U. chacranella. We base the synonymy on the shared head and metanotal markings, similarities in the wings (size, shape, venation, and pale veins), the enlarged abdominal spiracles, and the male genitalia. Chrysopa villica Navás, 1929. Rev. Soc. Entomol. Arg. 2: 222;"República Argentina;La Granja, Alta Gracia, 30 de febrero de 1924. Col. Bruch". Stange 1967Penny 1977: 21 (list); Brooks and Barnard 1990: 280 (list, as " 'Chrysopa' incertae sedis"); Oswald (2015, as Chrysopa). syn n. Lectotype (Figs 113,114). MACN, male (examined). Stange (1967: 38) and Adams (unpublished notes) reported that a type, from the province of Córdoba, is in the MACN; to stabilize the nomenclature of the genus, here this specimen is designated as the lectotype of C. villica (des. CAT). When we examined the specimen (a male), parts of the abdomen were cleared and in a vial of glycerine; unfortunately, the genitalia had been lost sometime after Adams examined the specimen. [Apparently Stange's (1967: 38) note of the type as a female is in error.] The original description mentions both La Granja and Alta Gracia as the type locality, and the type bears a label with these same two localities. Both places are in the Province of Córdoba, within ~73 km of each other: La Granja is a town in the Department of Columbus (~730 m), and Alta Gracia is a city in the Department of Santa Maria (~550 m). Support for synonymy. We based the synonymy on our examination of the external features of the type and the structures that are present on what remains of the abdomen, supplemented with Adams' unpublished notes on the genitalia.

Diagnosis.
Adults of U. chacranella are distinguished by the following external features: dark brown to black genal marks; antenna, including pedicel, pale, without dark ring; wing veins, including crossveins and gradates, light green (an occasional anal vein may appear light brown to brown); wings relatively wide, with rounded to subacute tips. Several series of specimens in the CAS and FSCA allowed us to associate males and females of this species. Males have abdominal spiracles greatly enlarged; S8+9 elongate, broad (ventral view), shallow, protruding well beyond T9+ect (lateral view); tip of S8+9 with flanged terminal setae; gonarcus with narrow bridge (dorsal view), slender apodemes with long, digitiform mesal process, elongate, narrow mediuncus, gonosaccus bilobed, each lobe with field of robust, widely spaced, terminally bent, gonosetae with large chalazae. The female has spermatheca with a large velum, small invagination; spermathecal duct elongate, curved without tight coils; and subgenitale (ventral view) short, stubby, and with well rounded, separated lobes. This species resembles U. confraterna. However, U. confraterna has at least some dark crossveins and slightly acute wing tips. Males of U. confraterna also lack the enlarged spiracles, and the dorsal (mesal) portion of the gonarcal bridge is robust and relatively straight, whereas in U. chacranella males the spiracles are greatly enlarged and the gonarcal bridge is slender and more rounded dorsally.
Redescription. Head: vertex smooth, slightly depressed mesally, inverted Ushaped 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 creamcolored, 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.
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, platelike 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 postero-   (Figs 122, 123). MLPA, male (examined). The type locality "Rio Negro" probably refers to the "Territorio del Rio Negro" which at the time of the collection was included within the Province of Patagonia. Today, Rio Negro is a separate province in the northern part of the Patagonian Region. Because Navás did not state how many specimens he had, and to preclude any confusion, the specimen that we photographed and that bears the number "3409" is hereby designated as the Chrysopa scalai lectotype (des. CAT). A red lectotype label has been applied to the specimen. Note: The specimen number (#3407) that González Olazo (1996: 381) reported for C. scalai is in error; it should read "3409". We examined the specimen briefly during a visit to the MLPA, and subsequently examined the cleared abdomen in detail. The abdomen is in fairly good condition except that most of the setae are absent and the gonarcus is distorted. Support for synonymy. Although González Olazo placed an U. argentina identification label on the C. scalai type, he did not publish a synonymy (see González Olazo (1996: 381). Our examination of the lectotype showed that the specimen has significant differences that preclude a synonymy with U. argentina. Notably, the posterovental corner of the ectoproct has a small knob that protrudes posteriorly, the gonarcus is thin and bears a long process, and the mediuncus is long, narrow, and straight (without a sinuous dorsal margin). All of these features are characteristic of U. confraterna. The specimen also differs from C. chacranella in that the spiracles are not enlarged, and the wing venation has notable brown areas. Chrysopa binaria . Arxius Inst. Ciènc., Inst. Est. Catalans, Sec. Ciènc. ("1919 Stange (1967: 31) and González Olazo (1996: 378) reported a female type in the MACN. We have seen this specimen and we consider it to be a paralectotype (Fig. 125a, b); it is not missing its head as reported by Legrand et al. (2008: 119). Its locality label has been changed from the original, and it does not include a collection date. However, it bears an origi- nal determination label in Navás' hand. Also, in the MACN, there is another specimen from the type locality, determined by the collector of the type specimen, C. Bruch (labels, Fig. 125c). It bears the same collection label (without date) as the paralectotype; we found no evidence that this specimen was seen by Navás, and we do not consider it to be part of the type series. Support for synonymy. The C. binaria type shares external and internal features, especially patterns of wing venation and coloration, head and body markings, with those of U. confraterna females. Also, we found nothing notable in the female abdomen or genitalia that would differentiate the species.
Diagnosis. Externally, the dorsal head markings and largely green longitudinal veins marked with brown at intersections distinguish U. confraterna from most of the other small Argentinian species (U. annulata, U. argentina, and U. chacranella). Its lack of a prominent stripe on the gena and clypeal margin distinguishes it from U. elbergi. However, the most reliable characters for identifying any of these species are in the male abdomen and genitalia. For example, U. confraterna is notable because of the unique size of its abdominal spiracles (diameter ~ 0.08 mm, ~0.1× length of segment) -slightly larger than those of most chrysopine adults, including U. argentina (diameter ~0.03 mm, ~0.03× length of segment), and considerably smaller than those of the other small Argentinian species, all of which have spiracles with a diameter of > 0.15 mm, > 0.2× length of segment. The shapes of its terminal segments and gonarcal complex are also distinctive. Redescription. 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.
Variation. 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 U. chacranella) to extensive (large sections of the longitudinal veins and all transverse veins brown or brownish).
Etymology. This species is named in memory of Professor Sanford S. Elberg , 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. Diagnosis. 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 U. elbergi is unique among all the small Argentinian species of Ungla. In addition, the head markings, largely green longitudinal veins, and reddish brown to brown transverse veins also help to sort U. elbergi and U. confraterna from other Argentinian species. The most reliable species-specific characters are found in the male. The R and Rs veins of the forewing are heavy in males; the moderately enlarged abdominal spiracles (diameter of A7 spiracle: 0.11 mm, > 0.13× length of segment) are larger than those of U. confraterna (diameter ~ 0.08 mm, ~0.1× length of segment) and U. argentina (diameter ~0.03 mm, ~0.03× length of segment), and yet considerably smaller than those of U. chacranella and U. annulata (diameter > 0.15-0.18 mm, > 0.2× length of segment). The shape of the U. elbergi gonosaccus and the large number of gonosetae evenly distributed over the entire dorsal surface of the gonosaccus are also unique.
Description. Head cream-colored, with vertex smooth, often shiny; inverted Ushaped 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.
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 U. confraterna); setae on ventral surface short, numerous; dorsal margin heavily sclerotized throughout; terminus slightly extended distally, flat, scooplike, bearing numerous, heavy, flanged setae along dorsal margins and heavy, unflanged setae distomesally; without the upturned tip of U. confraterna. Gonarcus with relatively flat, narrow bridge (dorsal, ventral views), arms extending downward, bowed in broad horseshoe shape; gonarcal arms narrow, rounded distally, with digitiform process extending forward and mesally at an acute angle from arm (lateral, dorsal views); mediuncus medium-length, narrow, convex dorsally, with base containing two elongate, juxtaposed rods; gonosaccus bilobed, with lobes contiguous and gonosetae in contiguous patch basally, lobes juxtaposed and with gonosetae in more discrete, separated patches distally; gonosetae all of somewhat similar size, arising from relatively small setal bases, probably facing mesally when lobes unexpanded, facing upward when lobes inflated; hypandrium internum narrow, broadly U-shaped, with comes faint.

Diagnosis.
Among Ungla species U. ivancruzi is unique in that the male has a broadly wedge-shaped S8+9 that is more typical of Chrysopodes than other Ungla males, which have S8+9 elongate and tapering. Nevertheless, we consider U. ivancruzi a good species of Ungla; it expresses most of the diagnostic features of the genus, including the distinctive inverted U-shaped mark on the vertex, dorsal apodeme on the male T9+ect reduced so that it does not extend beneath T8, gonarcal arch rounded, with a narrow, elongate mediuncus attached dorsally, and a bilobed gonosaccus, with each lobe bearing a patch of elongate gonosetae. Ungla ivancruzi can be distinguished from other Ungla by its (i) light colored (not black or dark brown) palpi, (ii) reddish genal markings, (iii) narrow gonarcal arch with thin, elongate arms lacking a prominent frontal process, and (iv) gonosaccus with gonosetae relatively well-spaced, not abutting each other. At this time, U. ivancruzi is the only Ungla species known from Brazil. See original description by Freitas (2007) for additional images. Redescription. 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.
Female. See Freitas (2007). Larvae and biology. Immatures were reared, and the three instars are described (Freitas 2007 (Figs 137,138a,139). ZMB, female (examined). Specimen in good condition, but discolored with age; abdomen in vial with glycerin. Stange (1967) listed it as a holotype; however Navás did not mention how many specimens he had; thus the specimen in the ZMB is here designated as the lectotype (des. CAT). A red lectotype label has been applied to the specimen. Support for generic placement. Currently, this species is known only from the lectotype, which is a female. Here, we base its generic placement on several features that characterize Ungla species: vertex with U-shaped dorsal marks; dark markings on the terminal segments of the maxillary and labial palpi, and a basal inner gradate that does not meet the Psm. The female genitalia are also typical of Ungla species.
Diagnosis. Ungla steinbachi is a pale-bodied Argentinian species known with certainty only from the type. It closely resembles U. chacranella in the following external features: vertex with small reddish mark, dark brown to black genal marks; antenna, including pedicel, pale, without dark ring; wings with rounded to subacute tips and with veins, including the crossveins and gradates, light green (an occasional anal vein may appear light brown to brown). However, three features lead us to recognize the two species as distinct from each other. First, the difference in size between the C. steinbachi type and the U. chacranella specimens that we studied is substantial; the forewings and hindwings of the C. steinbachi type are each more than 10% greater in length and width than those of the largest U. chacranella specimen we measured (For comparison, see Fig. 138). Second, the number of transverse veins is greater in the C. steinbachi type than in the U. chacranella specimens: radial cells in both the forewing and hindwing (13 versus a maximum of 12 in U. chacranella), inner and outer gradate veins (one more each than the maximum found in U. chacranella). Third, on the C. steinbachi type, the cells on either side of the inner gradate veins are longer and narrower than on the U. chacranella specimens.
We have seen only one other specimen, a somewhat discolored female from Jujuy Province that resembles the type of U. steinbachi in size and probable coloration of veins; however its pattern of venation differs considerably, and we cannot identify it with confidence. Given the small number of specimens (the type female) underlying our retention of U. steinbachi as a valid species separate from U. chacranella, we hope that additional pale-bodied Argentinian Ungla species will be collected and studiedespecially in regard to the range and pattern of morphological variation.
Redescription. Head: vertex smooth, slightly depressed mesally; inverted Ushaped 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.

Chrysopa reboredina
Type. MACN (IIES), sex unknown. Stange (1967: 36) reported the type was deposited at "San Miguel"; González Olazo (1996: 380) reported that it was at "INESALT" and that the abdomen and one hindwing were absent. Later, it was found in the Museo de Ciencias Naturales, Salta, Argentina (IIES) (von Ellenrieder 2009: 259), but now is in the MACN (von Ellenrieder, personal comunication). Apparently it is in a vial; its abdomen and one wing are missing, but the other three wings are present. We have not seen it.
Status of species. Probably a species of Ungla, in need of verification. In our opinion, the illustration and description by Navás are consistent with an Ungla species, and Adams' notes indicate that he considered the species to be associated with other species that later were included within Ungla. Apparently, he did not see the type. . Arxius Inst. Ciènc., Inst. Est. Catalans, Sec. Ciènc. ("1919 Type. Apparently destroyed. According to the original description, the type was deposited in the Navás collection (MZBS), but it is not there now (Monserrat 1985: 237). Adams' notes state that he saw a series of eight females and two males that were collected from La Paz (Dep. San Javier) Córdoba between January 15 and 31, 1928. These specimens are unlikely to be from the same series as the type. Adams also mentioned a male specimen that was collected from the type locality about a month later than the type; it carries a Navás determination label. At the time of our visit to the MACN, we were unaware of the specimens, and we did not see them in the collection.