Neuroptera: Chrysopidae)

A new genus of Neotropical Chrysopini (Neuroptera: Chrysopidae)

FRANCISCO SOSA1 & SERGIO DE FREITAS2 1 Universidad Centroccidental “Lisandro Alvarado”, Museo Entomológico “Dr. José Manuel Osorio” (UCOB), Barquisimeto, Lara, . E-mail: [email protected] 2 Universidade Estadual Paulista, Jaboticabal, São Paulo, Brazil (deceased)

Abstract

Titanochrysa Sosa & Freitas is a new genus of Neotropical Chrysopini (Chrysopidae: Chrysopinae) recorded from Costa Rica, Venezuela and Brazil. Titanochrysa gen. nov. shares several external and genitalic characters with Ceraeochrysa Adams, 1982; Chrysopodes Navás, 1913; Cryptochrysa Freitas & Penny, 2000; Parachrysopiella Brooks & Barnard, 1990 and Ungla Navás 1914. It may be distinguished from those genera by its very long sternite 8+9, sternites 2–8 usually with microtholi, male genitalia with the dorsal surface of the arcessus striated, gonosaccus well-developed, bearing elongate gonosetae and microsetae, and a spoon-like gonapsis. Herein, Titanochrysa circumfusa (Burmeister, 1939) [= Chrysopodes circumfusa (Burmeister)] comb. nov. and Titanochrysa pseudovaricosa (Penny) [= Ceraeochrysa pseudovaricosa Penny, 1998] comb. nov. were identified; Titanochrysa ferreirai Sosa & Freitas sp. nov. and Titanochrysa trespuntensis Sosa & Freitas sp. nov. were described. The external morphology, and male and female genitalia of all these species are described. Titanochrysa circumfusa (Burmeis- ter, 1939) comb. nov. is recorded for the first time from Venezuela.

Key words: Chrysopinae, Chrysopini, green lacewing, predator

Resumen

Titanochrysa Sosa & Freitas es un nuevo género Neotropical de Chrysopini (Chrysopidae: Chrysopinae) con registros para Costa Rica, Venezuela y Brasil. Titanochrysa gen. nov. comparte varias características externas y de la genitalia con Ceraeo- chrysa Adams, 1982; Chrysopodes Navás, 1913; Cryptochrysa Freitas & Penny, 2000; Parachrysopiella Brooks & Barnard, 1990 y Ungla Navás, 1914. Se diferencia de aquellos géneros por presentar el esternito 8+9 muy largo, esternitos 2–8 general- mente con microtholi, genitalia del macho con superfície dorsal del arceso estriada, gonosaco desarrollado con largas gonosetas y microsetas, gonapsis dilatada basalmente con ápice serreado. Aquí: Titanochrysa circumfusa (Burmeister, 1939) [= Chrysop- odes circumfusa (Burmeister)] comb. nov. y Titanochrysa pseudovaricosa (Penny, 1998) [= Ceraeochrysa pseudovaricosa Penny, 1998] comb. nov. fueron identificadas y Titanochrysa ferreirai Sosa & Freitas sp. nov., y Titanochrysa trespuntensis Sosa & Freitas sp. nov. descritas. La morfología externa, así como la genitalia del macho y la hembra son descritos. Titano- chrysa circumfusa (Burmeister, 1939) comb. nov. es reportada para Venezuela por primera vez.

Palabras clave: Chrysopinae, Chrysopini, crisópidos, depredator

Introduction

Adams (1978) organized the higher classification of Neotropical Chrysopidae including genera within tribes and subfamilies. Later, Adams (1982a, b) described Plesiochrysa Adams and Ceraeochrysa Adams, respectively. Brooks & Barnard (1990) identified 31 genera in Chrysopini, and Freitas & Penny (2000) added two new genera. At present, this tribe has 33 genera: nine are present in the Neotropics. Recently, we discovered that several species have traits that do not conform to any of the described Neotropical genera; they appear to belong in a distinct, previously undescribed genus. We herein describe this genus, Titano- chrysa gen. nov. We transfer two previously described species—Chrysopodes circumfusus (Burmeister, 1853) and Ceraeochrysa pseudovaricosa Penny, 1998—to the new genus, and we describe two new Titanochrysa species.

Accepted by A. C.-Ramos: 8 May 2012; published: 19 Jun. 2012 1 The four species share several characteristics: i) head with red occipital marks; ii) scape with dark red or black, longitudinal, lateral lines; iii) gena and lateral clypeus black or dark red tinged; iv) forewing generally with wide costal area; v) sternite 8+9 (S8+9) elongate, generally with microtholi; vi) gonapsis short, spoon-shaped; vii) females with S7 almost two times longer than tall; and viii) spermatheca with pillbox-shape. These characters indi- cate that the four species belong in Chrysopini, but not in either Chrysopodes Navás, 1913, Ceraeochrysa, or any other described Neotropical genus in this tribe. However, these species share characteristics with Ceraeochrysa; Chrysopodes; Cryptochrysa Freitas & Penny, 2000; Parachrysopiella Brooks & Barnard, 1990 and; Ungla Navás 1914 (see characteristics described by Brooks & Barnard, 1990; Penny, 1996; Freitas & Penny, 2000). The character combination described above precludes placement of these species in any described genus and warrants the new genus description for Neotropical Chrysopinae.

Material and methods

Specimens examined in this study are deposited in the following institutions: California Academy of Sciences Col- lection (CAS); Entomology Museum “Dr. José Manuel Osorio”, Universidad Centroccidental “Lisandro Avarado”, Barquisimeto, Lara, Venezuela (UCOB), and the Sergio de Freitas Entomological Collection FCAV-UNESP, Jabot- icabal, São Paulo, Brazil (SFCC). Type specimens will be deposited in the Museum of Zoology University of São Paulo, Brazil (MZUSP). Nomenclature for the terminalia and genitalia (male and female) was based in Adams & Penny (1985) and Brooks & Barnard (1990); nomenclature for the wings followed Tillyard (1916), complemented by Brooks & Bar- nard (1990), Nel et al. (2005) and Tauber (2008); methodology for the measurements and maceration is presented in Sosa & Freitas (2011).

Results and remarks

Titanochrysa Sosa & Freitas, gen. nov.

Type species: Titanochrysa circumfusa (Burmeister, 1939) comb. nov.

Generic characters. Vertex smooth or striated; scape and pedicel with a dark red, longitudinal stripe laterally; flagellum pale green, generally covered with black bristles, as long or longer than forewings; face elongate, labrum slightly incised; genae and lateral clypeus tinged with dark red or black; maxillary palpi black; labial palpi varia- bile, with black tinge; mandibles symmetrical or asymmetrical. Forewings generally with wide costal area, crossveins generally shaded. Male terminalia: ninth tergite and ectoproct (T9+ect) densely covered by stalked setae, base acute or truncated; callus cerci round or slightly ovate; sternites 8 and 9 (S8+9) elongate, fused; microtholi absent or present (Figs. 3A, 7A). Female terminalia: seventh sternite (S7) at least 2.0 times longer than wide; posterior margin of T9+ect usually with a medial cleft beneath level of callus cerci (Figs. 8A, 12A, 16A). Male genitalia: tignum, parameres, entoprocessus, gonocornua and spinellae absent; gonarcus U-shaped, gonarcal bridge narrow; arcessus usually with dorsal striation, apex trilobate, terminus with mesal beak flanked by lateral lobes; gonosaccus with long gonosetae generally on prominent papillae; microsetae present or absent. Gonapsis spoon- shaped, wide anteriorly, narrow posteriorly, short in relation to S8+9. Female genitalia: spermatheca pillbox- shaped; ventral impression shallow or deep; velum short; spermathecal duct elongate; bursa copulatrix sacular, with short, filamentous bursal gland; subgenitale cordate with dorsal section lobate. Other species: Titanochrysa pseudovaricosa (Penny, 1998) comb. nov; Titanochrysa ferreirai Sosa & Freitas sp. nov., and Titanochrysa trespuntensis Sosa & Freitas sp. nov. Etymology: The name Titanochrysa refers to the “Titans”, powerful dieties that had a large role in Greek mythology [= large size, in reference to the elongate S8+9] and to “Chrysos” meaning golden, as in the original description of green lacewings. The word is considered feminine. Generic relationships. Titanochrysa Sosa & Freitas gen. nov. shares external and genitalic characteristics with Ceraeochrysa, Chrysopodes, Cryptochrysa, Parachrysopiella and Ungla, Only Ceraeochrysa, Cryptochrysa

2 · Zootaxa 3351 © 2012 Magnolia Press SOSA and Parachrysopiella have a gonapsis, although Adams & Penny (1985) and Tauber (2003) recorded a short gonapsis for some species of Chrysopodes, but in accordance with Brooks & Barnard (1990) this structure should be absent in this genus. Ceraeochrysa adults are characterized by pale green coloration, pronotum generally with a red, longitudinal, lateral stripe; scape and pedicel with dorsal, lateral, or both dorsal and lateral longitudinal, red lines; frons and genae generally green, black in a few species [see Freitas et al. (2009) and Sosa & Freitas (2010, 2011)]. The following features of Ceraeochrysa adults distinguish them from Titanochrysa gen. nov. male sternites without microtholi, gonarcal complex with wide gonarcal bridge, well developed gonocornua, long or short arcessus with large median plate, gonapsis elongate (usually as long as S8+9), narrow, and bifurcate apically; female spermatheca usually elongate, J- or U-shaped (Adams 1982b; Brooks & Barnard 1990; Freitas et at. 2009). Cryptochrysa (known from one species) has the following adult characters that distinguish it from Titano- chrysa Sosa & Freitas gen. nov.: pronotum with a brown (not red), longitudinal, lateral stripe; scape at least 2.0 times longer than wide; forewing with narrow costal area; S8+9 elongate, but without microtholi; gonarcal complex with a narrow gonarcal bridge, gonocornua present as a pair of flattened plates, and arcessus spiculose subapi- cally; the gonapsis is elongate, with hook-shaped, lateral projections anteriorly (Freitas & Penny 2000). Parachrysopiella adults differ from Titanochrysa Sosa & Freitas gen. nov. in that the body is small, marked with a longitudinal, median brown stripe on the pronotum; eyes very small compared with head width; ectoproct extended posteriorly and covered by coarse spine-like setae apically, gonarcal complex with short, narrow gonarcal bridge, arcessus with dorsal surface smooth, apex hook-shaped, not flanked by lateral lobes; the gonapsis is flat, broad and crown-shaped apically (Brooks & Barnard 1990; Penny 1996). This genus is known only from the Andean region. Titanochrysa Sosa & Freitas gen. nov. and Ungla are very similar in their female genitalia. However, the wider costal area in the forewings, basal inner gradate meeting the Psm, T9+ect with dorsal apodemes, S8+9 generally with microtholi and male genitalia with short, wide and generally striate arcessus, observed in Titanochrysa Sosa & Freitas gen. nov. distinguish it from Ungla, where the costal area is narrow, the basal inner gradate generally does not meet the Psm, the T9+ect is without an apodeme, S1–8 without microtholi, and the arcessus is longer than wide, without dorsal striations. The wide costal area on the forewings of the Titanochrysa Sosa & Freitas gen. nov. is shared with many species of Chrysopodes, but the elongate S8+9, gonosaccus bearing elongate gonosetae and microsetae, spoon-shaped gonapsis, and pillbox-like spermatheca of Titanochrysa distinguish it from this genus. Geographical distribution. Costa Rica, Venezuela and Brazil.

Titanochrysa circumfusa (Burmeister, 1839) comb. nov.

Synonymy Chrysopa circumfusa Burmeister, 1939: 980. Chrysopa burmeisteri Navas, 1929: 858. Banks 1944: 10. Chrysopodes (Chrysopodes) circumfusa (Burmeister). Brooks & Barnard, 1990: 272, Figs. 331–332.

Diagnosis. Adults green with longitudinal, yellow stripe dorsally; antennae longer than forewings; base of T9+ect truncated; microtholi absent. Inner gradate veins in irregular series; outer gradates in regular series. Mandibles symmetrical. Redescription. Head. Vertex yellow, smooth; occiput with dark red spots (Fig. 1A); scape and pedicel yellowish green with reddish brown longitudinal stripe laterally, extending onto antennal base (Fig. 1B); flagellum pale, longer than forewing; frons and mesal clypeus white; gena and lateral clypeus dark red; maxillary palpus black; labial palpus pale, with last segment black (Fig. 1C); mandibles both with prominent basal tooth (Fig. 17A). Thorax. Yellowish green, with longitudinal yellow band; pronotum square-shaped, with narrow, diffused lateral red stripe (Figs. 1A, 1C); meso- and metanota yellowish green, without marks. Pleura, sternal areas, legs pale green. Wings (Fig. 2): forewing with longitudinal veins green; crossveins green (except basal subcostal vein black), all except the costal crossveins with black markings on surrounding membrane. Inner gradates spaced irreg- ularly, 2nd and 3rd sometimes aligned; outer gradates regularly spaced. First radial crossvein originating after origin of radial sector, meeting Psm after apex of intramedian cell; intramedian cell triangular. Hindwing with longitudinal veins, crossveins pale green, gradate series parallel, with lightly fumose borders in some exemplars.

A NEW GENUS OF NEOTROPICAL CHRYSOPINI Zootaxa 3351 © 2012 Magnolia Press · 3 FIGURE 1. Titanochrysa circumfusa (Burmeister, 1839) comb. nov. (Male, Sanare, Lara, Venezuela). A) Head and pronotum, dorsal. Note the yellow mesal band; B) Head and pronotum, lateral. Note the dark red marks on the lateral scape, base of antenna, gena and lateral clypeus; as well as the pale basal labial and maxillary palpomeres; C) Head, frontal.

FIGURE 2. Titanochrysa circumfusa (Burmeister, 1839) comb. nov. Wings. bsx = basal subcostal crossvein; c.a = costal area; cx = costal crossvein; ig = inner gradates; im = intramedian cell; og = outer gradates; r-m1 = first radial crossvein; Rs = radial sector.

FIGURE 3. Titanochrysa circumfusa (Burmeister, 1839) comb. nov. (Male, Sanare, Lara, Venezuela). A) Terminalia, lateral; B) gonarcal complex, dorsal; C) gonarcal complex, lateral; D–E) Gonarcal complex, frontodorsal; F) Gonarcal complex, frontal. Note the prominent the elongate gonosetae on papillae; G) Gonapsis, frontal, frontolateral and lateral. ap-gs = lateral apodemes of gonarcus; ar = arcessus; cc = callus cerci; d.ap = apodeme of T9+ect; gp = gonapsis; gs = gonarcus; gsac = gonosaccus; gst = gonosetae; l.lb = lateral lobe; st = setae; S8+9 = fused sternites 8 and 9; T9+ect = tergite 9 and ectoproct; v.ap = apodeme of S8+9.

Abdomen. Yellowish green, with longitudinal yellow band dorsally; pleura with dark red spots laterally; sternites without microtholi. Male terminalia (Fig. 3A): T9+ect elongate, basally truncated, with scattered stalked setae anteriorly, densely setose posteriorly; dorsal apodeme simple, slightly curved, not reaching callus cerci; callus cerci round with 28–30 trichobothria; S8+9 fused, with truncate apex; S8 ca. two times longer than wide, with dorsal margin straight; ventral apodeme short, narrow, extending along dorsal margin of S8. Gonarcus broadly truncate, U-shaped (Fig. 3B); lateral arms round, with anterior projection decurved ventrally, hook-like (Fig. 3C–E); arcessus short, broad, with dorsal rods on median plate forming an X-shape, apex decurved, trifurcated with dorsal striations, inner area beneath arcessus with short setae in two lateral series (Fig. 3B–E); gonosaccus long, with numerous gonosetae on papillae (Fig. 3C, F). Gonapsis spoon-shaped basally, hooked apically (Fig. 3G). Female terminalia (Fig. 4A): S7 ca. three times longer than wide, tapering apically; T9+ect without cleft on posterior margin; callus cerci round, with ca. 26 trichobothria; subgenitale with anterior section prominent, widely lobate (Fig. 4A–B); spermatheca pillbox-shaped, ventral impression deep; spermathecal duct elongate (Fig. 4C–E). Measurements. Male (n=2): Head: width 1.4–1.5 mm. Pronotum: length 0.9–1.0 mm, width 0.9–1.0 mm. Forewing: length 14.0–14.7 mm, width 4.9–5.1 mm, length/width ratio = 2.9–2.9: 1. Six to seven inner, six to seven outer gradates. Hindwing: length 12.6–13.1mm, width 3.9–4.0 mm, length/width ratio = 3.2–3.3: 1. Five to seven inner, five to seven outer gradates. Female (n=1): Head: width 1.5 mm. Pronotum: length 1.0 mm, width 1.0 mm. Forewing: length 15.6 mm, width 5.5 mm, length/width ratio= 2.8: 1. Seven inner, seven outer gradates Hind- wing: length 14.3 mm, width 4.3 mm, length/width ratio = 3.3: 1. Seven inner, seven outer gradates.

A NEW GENUS OF NEOTROPICAL CHRYSOPINI Zootaxa 3351 © 2012 Magnolia Press · 5 FIGURE 4. Titanochrysa circumfusa (Burmeister, 1839) comb. nov. (Female, Sanare, Lara, Venezuela). A) Terminalia, lateral; B) Terminalia lateral, note detail of subgenitale; C–D) Spermatheca, lateral; E) Spermatheca, dorsal. cc = callus cerci; gl = gonapophysis lateralis; S7 = sternite 7; sg = subgenitale; sp = spermatheca; sp.d = spermathecal duct; T9+ect = Tergite 9+ectoproct; v = velum; v.i = ventral impression.

Material examined. VENEZUELA. Lara: Sanare [9º42’N / 69º36’W, 1753 m], 14.i.2008, F. Sosa & M. Roa Legs (UCOB); Same, 18.i.2009, 1♂, 1♀, F. Sosa & F. Díaz Legs (UCOB); Same, 18.i.2009, 1♂, F. Sosa & F. Díaz Legs (MZUSP); Same, 14.i.2008, 2♀, F. Sosa & M. Roa, (UCOB); Same, 16.ii.2008, 1♂, F. Sosa & F. Díaz Legs (UCOB), head macerated. Species relationships. Titanochrysa circumfusa comb. nov. differs from other species of the genus by the absence of microtholi from S2–8. The forewing venation is fumose like that of Chrysopodes (Chrysopodes) brevi- ata Adams & Penny, 1985, Chr. (Chr.) limbata (Navás, 1926), and Chr. (Chr.) nebulosa Adams & Penny, 1985. However the dark red, longitudinal, lateral line on the scape, dark genae, elongate S8+9, male genitalia with developed gonosaccus bearing elongated gonosetae on prominent papillae, and spoon-like gonapsis separate Titano- chrysa circumfusa comb. nov. from those species. Remarks. i) The lectotype of Chrysopa circumfusa is in the Museum für Naturkunde [Humboldt-Universität zu Berlin, Berlin, ZMBH]; sex unknown. Its labels read: [1] “S. João d. Rey, Sellow”; [2] “circumfusa Burm. / Sch- neider.*”; [3] “338”; [4] “Chrysopodes”; [5] “LECTOTYPE / Chrysopa circumfusa / Burmeister desig. / by P. A. Adams 1985 / det. C. A. Tauber 2011”. The type locality is probably São João del Rei in the state of Minas Gerais, Brazil. [Identification of the species and information on the Lectotype were provided by C. A. Tauber, Entomology, Cornell University.]; ii) The mandibles and galea of Chr. (Chr.) circumfusa illustrated by Brooks & Barnard (1990: Figs 331–332) differ from those of our specimens; it appears that their drawings were based on an erroneous identification of the lectotype (C. A. Tauber, personal communication); iii) Banks (1944) reported Chrysopa circumfusa from Paramaibo, Suriname and mentioned that Chrysopa burmeisteri Navás, 1929 is very similar or identical to Chrysopa circumfusa; his statement is taken as a synonymy (C. A. Tauber, personal communication). The type of Chrysopa burmeisteri was from Brazil (Espiritu Santo). Therefore, the species has been reported from Brazil and Suriname. Geographical distribution. Brazil, Suriname and Venezuela (new record).

Ceraeochrysa pseudovaricosa Penny, 1998: 62, Figs. 18–28. Penny, 2002: 216, Figs. 233–238. Freitas et al. 2009: 569, Fig. 41.

FIGURE 5. Titanochrysa pseudovaricosa (Penny, 1998) comb. nov. (Puntarenas, Costa Rica). A) Head and pronotum, dorsal. Note the narrow, yellow, dorsal band; B) Head and pronotum, lateral. Note the black marks on the lateral scape, gena and lateral clypeus; C) Head, frontal.

FIGURE 6. Titanochrysa pseudovaricosa (Penny, 1998) comb. nov. Wings. bsx = basal subcostal crossvein; c.a = costal area; cx = costal crossveins; ig = inner gradates; im = intramedian cell; og = outer gradates; r-m1 = first radial crossvein; Rs = radial sector. The arrow indicates the dark reticulate network of short crossveins covered by round black spot.

Diagnosis. Adults pale green, with narrow, yellow, longitudinal band medially. Pronotum wider than long, with dark red, longitudinal stripe laterally. Inner gradate series of forewing with dark reticulate network of short veins covered by black medial spot. Microtholi present on A2–8. Mandibles asymmetrical.

A NEW GENUS OF NEOTROPICAL CHRYSOPINI Zootaxa 3351 © 2012 Magnolia Press · 7 Redescription. Head. Vertex with light, transverse striate; occiput with dark red spot laterally (Fig. 5A); scape and pedicel pale green, with reddish brown longitudinal stripe laterally, extending onto antennal base, with short apical dorsal line; flagellum pale, slightly shorter than forewing, covered by black bristles (Fig. 5B); frons white with diffuse, red-wine spots beneath antennae; clypeus white, marked laterally with dark red; gena reddish brown; maxillary palp black, labial palp pale, darker at tip (Fig. 5B–C); mandibles with basal tooth on left mandible, without indication of basal tooth on right mandible (Fig. 17B).

FIGURE 7. Titanochrysa pseudovaricosa (Penny, 1998) comb. nov. (Male, Puntarenas, Costa Rica). A) Terminalia, lateral; B) gonarcal complex, dorsal; C) gonarcal complex, lateral; D) Gonarcal complex, ventral; E) Gonarcal complex, frontal; F) Gonapsis, lateral. ap-gs = lateral apodemes of gonarcus; ar = arcessus; cc = callus cerci; d.ap = apodeme of T9+ect; l.lb = lateral lobe; gs = gonarcus; gsac = gonosaccus; gst = gonosetae; S8 = sternite 8; S9 = sternite 9; v.ap = apodeme of S8+9.

Thorax. Pro, meso- and metanota pale green, with longitudinal, yellow dorsal band medially; pronotum wider than long, with broad, longitudinal, dark red stripe laterally (Fig. 5A). Pleura, sternal areas, and legs pale green. Wings (Fig. 6): forewing with longitudinal veins pale green; crossveins pale green, with darkening at intersections with longitudinal veins; gradate veins dark, with some infuscation of surrounding membrane, especially on inner gradates; inner gradates with dark reticulate network of short veins covered by rounded black spot on medial area. First radial crossvein arising after origin of radial sector, reaching Psm before apex of intramedian cell; intramedian cell triangular. Hindwing with all veins pale green, except for intersections of crossveins with subcosta and first radial vein; apex acute; inner and outer gradates in regular, slightly diverging series. Abdomen. Dorsum with longitudinal yellow median stripe, green laterally, pale yellow ventrally; dark pleural spots on apices of basal segments. Male terminalia: T9+ect elongate, basally tapering to acute anterior margin, covered by scattered, stalked setae; callus cerci ovate, with ca. 25–27 trichobothria; dorsal apodeme simple, elongate, reaching callus cerci (Fig. 7A); S8+9 covered by numerous setae with thickened bases; S8 as long as wide, with dorsal margin domed, densely covered with microtholi; ventral apodeme short (Fig. 7A). Male genitalia: gonarcal

8 · Zootaxa 3351 © 2012 Magnolia Press SOSA complex with narrow bridge (Fig. 7B), lateral apodemes wide posteriorly, tapering anteriorly (Fig. 7C); arcessus short, broad, decurved, trifurcate apically, lightly striated dorsally (Fig. 7D); gonosaccus simple, with few, thin, scattered gonosetae, microsetae present (Fig. 7E); gonapsis elongate with apex lightly serrated apically, with lateral projection anteriorly (Fig. 7F). Female terminalia (Fig. 8A): S7 ca. 3.4 times longer than wide, with dorsal margin bilobed, densely covered by thick setae; T9+ect with dorsal margin straight, forming ca. 90o angle with corner of posterior margin; posterior margin with ventral cleft beneath the callus cerci; callus cerci ovate with ca. 24 trichobothria; spermatheca pillbox-shaped, with ventral impression deep; spermathecal duct elongate (Fig. 8B–C). Sub- genitale cordate, longer than wide, with broad medial notch.

FIGURE 8. Titanochrysa pseudovaricosa (Penny, 1998) comb. nov. (Female, Puntarenas, Costa Rica). A) Terminalia, lateral; B) Spermatheca, lateral; C) Spermatheca, dorsal. S6 = sternite 6; S7 = sternite 7; sp = spermatheca; sp.d = spermathecal duct; T9+ect = Tergite 9+ectoproct; v = velum; v.i = ventral impression.

Measurements. Allotype. Female. Head: width 1.3 mm. Pronotum: length 0.6 mm, width 1.0 mm. Forewing: length 14.6 mm, width 5.1 mm, length/width ratio = 2.9:1. Five inner, seven outer gradates. Hindwing: length 13.0 mm, width 4.3 mm, length/width ratio = 3.0: 1. Four inner, six outer gradates. Male (n=2): Head: width 1.2–1.3 mm. Pronotum: length 0.5–0.6 mm, width 0.9–0.9 mm. Forewing: length 11.9–12.7 mm, width 4.1–4.4 mm, length/width ratio = 2.9–2.9: 1. Four to five inner, five to six outer gradates. Hindwing: length 10.5 mm, width 3.8 mm, length/width ratio = 2.8: 1. 5 inner, six outer gradates. Female (n=1): Head: width 1.2 mm. Pronotum: length 0.5 mm, width 0.9 mm. Forewing: length 12.8 mm, width 4.4 mm, length/width ratio = 2.9:1. Five inner, six outer gradates. Hindwing: length 11.2 mm, width 3.5 mm, length/width ratio = 3.2:1. Four inner, five outer gradates. Material examined. Allotype: COSTA RICA: Puntarenas, La Amistad National Park (8º57’N / 82º50’W, 1500 m), 18.ii.1991, Norman D. Penny Leg (MZC). Paratypes: COSTA RICA: Puntarenas, La Amistad National Park (8º57’N / 82º50’W, 1500 m), 18.ii.1991, 2♂, Norman D. Penny Leg (MZC); Same, 18.ii.1991, exemplar without abdomen; Same, 18.ii.1991, 1 exemplar without abdomen, clear head; Same, 19.ii.1991; 1♂ not dissected; Same, 1600 m, 21.ii.1991, 1♀, not dissected, Same, 1.iii.1991, 1♂, not dissected. Remarks. Penny (1998) described symmetrical mandibles; however on the paratype we examined the mandibles are clearly asymmetrical, with a basal tooth on the left mandible and without indication of a basal tooth on the right mandible. Species relationships. Titanochrysa pseudovaricosa comb. nov. is similar to Titanochrysa ferreirai sp. nov. Both species share similar forewing venation, S8+9 shape, male gonarcal complex, and gonapsis. However, the Titanochrysa pseudovaricosa comb. nov. forewing has a network of short, dark, reticulate veins with shadows in the middle of the inner gradate series, whereas in Titanochrysa ferreirai sp. nov. all of the inner gradates are shaded, but no reticulate veins are present. Sometimes, the Titanochrysa ferreirai forewing has a third series of gradates, which is not present on Titanochrysa pseudovaricosa comb. nov. The most important difference is observed

A NEW GENUS OF NEOTROPICAL CHRYSOPINI Zootaxa 3351 © 2012 Magnolia Press · 9 in the female terminalia: S7 in Titanochrysa pseudovaricosa comb. nov. is almost 3.4 times longer than wide, and it is covered with intermediate-sized setae; in Titanochrysa ferreirai sp. nov. S7 is 2.5 times longer than wide and densely covered by long, thickened setae. Geographical distribution. Costa Rica.

Titanochrysa ferreirai Sosa & Freitas sp. nov.

Holotype male: Brazil. Minas Gerais. Tres Pontas [21o25’S / 45o30’W, 900 m], 8.viii.2009, Ferreira C. S Leg. Deposited in the Museum of Zoology, Universidade de São Paulo (MZUSP), São Paulo, Brazil. Measurements. Head: width 1.1 mm. Pronotum: length 0.5 mm, width 0.8 mm. Forewing: length 11.4 mm, width 4.1 mm, length/ width ratio = 2.8:1. Five inner, six outer gradates. Hindwing: length 9.9 mm, width 3.3 mm, length/width ratio = 3: 1. Four inner, five outer gradates. Diagnosis. Adults olive green, with longitudinal dorsal yellowish green band. Protonum with dark red, longitudinal stripe laterally. Forewing inner gradates margined with round black spots, outer gradates veins lightly shaded. Microtholi present on S2–8. Mandibles asymmetrical. Description. Head. Vertex striated transversally; occiput with red lateral spots (Fig. 9A); scape and pedicel yellowish green with dark red longitudinal stripe laterally, extending onto antennal base (Fig. 9B); flagellum pale, slightly shorter than forewing; frons creamy with dark red spots beneath scapes; clypeus white, marked with dark red laterally; gena dark red; maxillary and labial palpi black (Fig. 9C); mandibles with prominent basal tooth on left mandible (Fig. 17C). Thorax. Green, with yellowish green longitudinal band dorsally. Pronotum wider than long, green, with broad, longitudinal, dark red stripe laterally (Fig. 9A); meso- and metanota pale green without markings; pleura, sternal areas, legs pale green. Wings (Fig. 10): forewing with longitudinal veins pale green; crossveins pale green, dark at intersections with longitudinal veins; costal crossveins with black extremities, radial crossveins with anterior extremity shaded black; inner and outer gradates black, arranged in parallel series; inner gradates margined by round, black spots, outer gradates lightly shaded. Hindwing with longitudinal veins, crossveins pale green, except terminal extremities of forked apical veins marked with dark spots. Abdomen. Green, with yellowish green band dorsally, posterior extremity of each segment, tinged with dark red dorsally. Male terminalia (Fig. 11A): T9+ect elongate, basally tapering to acute terminus, covered anteriorly with scattered medium-sized setae; dorsal apodeme simple, straight reaching callus cerci ventrally; callus cerci ovate bearing 21–23 trichobothria; S8+9 tapering; S8 slightly longer than wide, densely covered with medium- sized setae and microtholi, dorsal margin dome-like; ventral apodeme of medium length. Male genitalia: Gonarcus truncated in dorsal view (Fig. 11B); lateral apodemes narrow with inverted comma-shape, anterior extremity tapering to acute tip projected laterally (Fig. 11B–E); arcessus short, with dorsal rods broad, decurved apically, dorsal surface lightly striated, flanked by lateral lobes (Fig. 11D); gonosaccus simple, with few, thin, scattered gonosetae, densely covered by microsetae (Fig. 11C, E). Gonapsis short, expanded anteriorly, spoon-shaped, with subapical teeth (Fig. 11F). Hypandrium internum broadly U-shaped (Fig. 11G). Female terminalia (Fig. 12A): S7 trapezoi- dal, ca. 2.5 times longer than wide; slightly tapering distally, densely covered with long, stalked setae, dorsal margin sinuous; T9+ect with dorsal margin straight, forming ca. 90o angle at posterodorsal margin; posterior margin with ventral cleft beneath callus cerci; callus cerci ovate, with ca. 30 trichobothria (Fig. 12A); spermatheca pillbox- shaped, ventral impression deep, spermathecal duct elongate (Fig. 12B–D). Subgenitale cordate, lobate dorsally (Fig. 12E–F). Measurements. Male (n=2): Head: width 1.1–1.2 mm. Pronotum: length 0.5–0.5 mm, width 0.8–0.9 mm. Forewing: length 11.4–11.8 mm, width 4.0–4.2 mm, length/width ratio = 2.8–2.9. Four inner, five outer gradates. Hindwing: length 10.2–10.4 mm, width 3.0–3.3 mm, length/width ratio = 3.2–3.4:1. Three inner, four outer gradates. Females (n=2): Head: width 1.2–1.3 mm. Pronotum: length 0.5–0.6 mm, width 0.9–0.9 mm. Forewing: length 12.6–13.8 mm, width 4.5–5.0 mm, length/width ratio. Four to five inner, five to six outer gradates = 2.8–2.8. Hindwing: length 11.2–11.9 mm, width 3.7–3.9 mm, length/width ratio = 3.0–3.1: 1. Three to four inner four to five outer gradates.

10 · Zootaxa 3351 © 2012 Magnolia Press SOSA FIGURE 9. Titanochrysa ferreirai Sosa & Freitas sp. nov. (Male, Três Pontas, Minas Gerais, Brazil). A) Head and pronotum, dorsal. Note the greenish-yellow dorsal band; B) Head and pronotum, lateral. C) Head, frontal.

FIGURE 10. Titanochrysa ferreirai Sosa & Freitas sp. nov. Wing venation. bsx = basal subcostal crossvein; c.a = costal area; cx = costal crossvein; ig = inner gradates; im = intramedian cell; og = outer gradates; r-m1 = first radial crossvein; rx = radial crossvein; Rs = radial sector

A NEW GENUS OF NEOTROPICAL CHRYSOPINI Zootaxa 3351 © 2012 Magnolia Press · 11 FIGURE 11. Titanochrysa ferreirai Sosa & Freitas sp. nov., (Male, Três Pontas, Minas Gerais, Brazil). A) Terminalia, lateral; B) gonarcal complex, dorsal; C) gonarcal complex, lateral; D) Gonarcal complex, frontodorsal; E) Gonarcal complex, ventral. Note the prominent papillae; F) Gonapsis, lateral; G) Hypandrium internum, frontal. ap-gs = lateral apodemes of the gonarcus; cc = callus cerci; d.ap = apodeme of T9+ect; gp = gonapsis; gs = gonarcus; gsac = gonosacus; gst = gonosetae; l.lb = lateral lobe; mst = microsetae; S8 = sternite 8; S9 = sternite 9; v.ap = apodeme of S8+9.

Material examined. Allotype: Brazil. Minas Gerais. Tres Pontas [21o25’S / 45o30’W, 900 m], 6.vi.2009, Fer- reira C. S Leg (SFCC). Paratypes: Brazil. Minas Gerais. Tres Pontas [21o25’S / 45o30’W, 900 m], 15.viii.2008, 1♂, Ferreira C. S Leg (SFCC); Same, 14.iii.2009, 1♂, Ferreira C. S Leg (SFCC); Same, 6.vi.2009, 1♀, Ferreira C. S Leg (SFCC); Same, 13.vi.2009, 1♂, Ferreira C. S Leg (SFCC); Same, 20.vi.2009, 3♂♂, Ferreira C. S Leg (SFCC); Same, 4.vii.2009, 1♂, 2♀♀, Ferreira C. S Leg (SFCC); Same, 11.vii.2009, 4♂, 4♀, Ferreira C. S Leg (SFCC); Same, 25.vii.2009, 1♂, 2♀♀, Ferreira C. S Leg (SFCC); Same, 8.viii.2009, 1♂, 2♀♀, Ferreira C. S Leg (SFCC); Same, 5.ix.2009, 1♂, Ferreira C. S Leg (SFCC); Same, 6.vi.2009, 2♂♂, Ferreira C. S Leg (UCOB); Same, 8.viii.2009, 1♂, Ferreira C. S Leg (MIZA). Remarks. In some specimens, the left and right wings show different venation, and sometimes two inner gradate veins are surrounded by a single spot. Etymology. The species is named in honor of the agricultural engineer Cleidson Soares Ferreira who collected many chrysopids from the southern part of Minas Gerais state, Brazil. Species relationships. See T. pseudovaricosa comb. nov. Externally, Titanochrysa ferreirai sp. nov. resembles Ungla laufferi (Navás, 1922) based on descriptions provided by Freitas et al. (2009) and Tauber & Flint (2010). Both species have the frons tinged with red laterally; genae, lateral clypeus, maxillary and labial palpi black, and spermatheca pillbox-shaped. However, Titanochrysa ferreirai sp. nov. differs from U. laufferi in that it has a wide costal area, shading on the membrane around the inner gradates, and a cordate subgenitale with elongate distal section.

FIGURE 12. Titanochrysa ferreirai Sosa & Freitas sp. nov. (Female, Três Pontas, Minas Gerais, Brazil). A) Terminalia, lateral; B–C) Spermatheca, lateral; D) Spermatheca, dorsal E) Terminalia, ventral; F) Subgenitale, ventral. cc = callus cerci; g.l = gonapophysis lateralis; S7 = sternite 7; sg = subgenitale; sp = spermatheca; sp.d = spermathecal duct; T9+ect = Tergite 9+ectoproct; v = velum; v.i = ventral impression.

Titanochrysa trespuntensis Sosa & Freitas nov. sp

Holotype male, Minas Gerais. Tres Pontas [21o25’S / 45o30’W, 900 m], 10.i.2009, Ferreira C. S Leg. Deposited in the Museum of Zoology, Universidade de São Paulo (MZUSP) São Paulo, Brazil. Measurements. Head: width 1.1 mm. Pronotum: length 0.5 mm, width 0.8 mm. Forewing: length 10.1 mm, width 3.6 mm, length/width ratio = 2.8:1. Four inner, six outer gradates. Hindwing: length 9.3 mm, width 2.8 mm, length/width ratio = 3.3:1. Four inner, four outer gradates. Diagnosis. Adults yellowish green, with broad, dark red, longitudinal stripe laterally. Fore- and hindwings without shading on surrounding membrane, gradate veins in parallel series. S8+9 with a protuberance on the apical margin. Microtholi present on S2–8. Mandibles symmetrical. Description. Head. Vertex smooth, yellowish; occiput without marks (Fig. 13A); scape pale green, with dark red, longitudinal line laterally not reaching antennal base; pedicel pale green with dorsal black spot laterally; flagellum pale, covered with back bristles, slightly shorter than forewing (Fig. 13B); frons white; gena black; clypeus white, tinged with black laterally; maxillary, labial palpi black (Fig. 13C). Mandibles both with prominent basal tooth (Fig. 17D).

A NEW GENUS OF NEOTROPICAL CHRYSOPINI Zootaxa 3351 © 2012 Magnolia Press · 13 FIGURE 13. Titanochrysa trespuntensis Sosa & Freitas sp. nov. (Male, Tres pontas, Minas Gerais, Brazil). A) Head and pronotum, dorsal. Note the greenish yellow, dorsal band; the arrow marks the black spot at the base of the antenna; B) Head and pronotum, lateral. Note the black marks on the lateral scape (arrow), pedicel (arrow), gena, lateral clypeus, and maxillary and labial palpi. Also, note the black bristles on the flagellum (arrow); C) Head, frontal.

FIGURE 14. Titanochrysa trespuntensis Sosa & Freitas sp. nov. Wings. A = anal veins; bsx = basal subcostal crossvein; c.a = costal area; cx = costal crossvein; ig = inner gradates; im = intramedian cell; og = outer gradates; m-cu2 = second medial- cubital crossvein; r-m1 = first radial crossvein; Rs = radial sector.

14 · Zootaxa 3351 © 2012 Magnolia Press SOSA FIGURE 15. Titanochrysa trespuntensis Sosa & Freitas sp. nov. (Male, Três Pontas, Minas Gerais, Brazil). A) Terminalia, lateral; B) Terminalia, lateral. Note: the apical protuberance on S8+9 (arrow); C) Gonarcal complex, dorsal; D) Gonarcal complex, lateral; E) Gonarcal complex, frontodorsal. Note: the dotted orange line indicates the location of the setae beneath the arcessus. F) Gonapsis, lateral; G) Hypandrium internum, frontal. ap-gs = lateral apodemes of gonarcus; ar = arcessus; d.ap = apodeme of T9+ect; l.lb = lateral lobe; gp = gonapsis; gs = gonarcus; gst = gonosetae; S8+9 = fused sternites 8 and 9; v.ap = apodeme of S8+9; T9+ect = tergite 9+ectoproct.

Thorax. Pale green, with yellow, longitudinal band dorsally; pronotum wider than long, with dark red, longitudinal stripe laterally. Meso- and metanota with red-wine line dorsolaterally (Fig. 13A). Pleura, sternal areas, legs pale green. Wings (Fig. 14): forewing, hindwing densely covered with black microtrichia apically. Forewing with longitudinal veins dark green; crossveins dark at intersections with longitudinal veins, not infuscate; costal crossveins, radial crossveins, basal subcostal crossvein, inner gradates, outer gradates, m–cu2, 1A, 2A black. Outer, inner gradates in parallel series, slightly convergent apically. First radial crossvein originating after origin of radial sector, extending approximately to apex of intramedian cell; intramedian cell ovate. Hindwing with longitudinal veins green, black intersections with crossveins; costal crossveins, inner and outer gradates black. Abdomen. Yellowish green dorsally, green laterally, with dark red spots laterally on each tergite. Male terminalia (Fig. 15A): T9+ect elongate basally, tapering at basal margin, covered by stalked setae; dorsal apodeme simple, reaching callus cerci. Sternite S8+9 with a protuberance on the apical margin (Fig. 15B), numerous setae stemming from thickened bases, ventral apodeme elongate. Male genitalia: gonarcus truncated in dorsal view (Fig. 15C), slender in lateral view; lateral apodemes shaped like inverted comma, anterior extremity with truncated apex (Fig. 15D); arcessus short, broad, decurved, trifurcate apically, with field of short setae beneath, dorsal rods parallel, (Fig. 15C, D); gonosaccus with sparse, thin, scattered gonosetae (Fig. 15D). Gonapsis expanded anteriorly, with short, acute projection laterally, wide with round, expanded margin apically (Fig. 15F). Hypandrium internum V-shaped (Fig. 15G). Female terminalia: S7 ca. 2.0 times longer than wide, densely covered with medium-sized setae; T9+ect with dorsal, ventral margins round; callus cerci round, with ca. 24 trichobothria (Fig. 16A). Female genitalia: spermatheca (Fig. 16B–D) pillbox-shaped, spermathecal duct elongate; velum curved laterally; ventral impression shallow. Subgenitale cordate, with elongate medial notch (Fig. 16E–F).

A NEW GENUS OF NEOTROPICAL CHRYSOPINI Zootaxa 3351 © 2012 Magnolia Press · 15 FIGURE 16. Titanochrysa trespuntensis Sosa & Freitas sp. nov. (Female, Três Pontas, Minas Gerais, Brazil). A) Terminalia, lateral; B–C) Spermatheca, lateral; D) Spermatheca, dorsal E) Terminalia, ventral; F) Subgenitale, frontal and lateral. gl = gonapophysis lateralis; S7 = sternite 7; sg = subgenitale; sp = spermatheca; sp.d = spermathecal duct; T9+ect = Tergite 9+ectoproct; v = velum; v.i = ventral impression.

Measurements. Male (n=2): Head: width 1.2–1.2 mm. Pronotum: length 0.5–0.5 mm, width 0.9–0.9 mm. Forewing: length 10.7–10.8 mm, width 3.7–3.8 mm, length/width ratio = 2.8–2.9:1. Five inner, five to six outer gradates. Hindwing: length 9.6–9.7 mm, width 2.9–3.0 mm, length/width ratio = 3.2–3.3:1. Four inner, four to five outer gradates. Female (n=2): Head: width 1.–1.1 mm. Pronotum: length 0.5–0.5 mm, width 0.9–0.9 mm. Fore- wing: length 11.0–11.3 mm, width 3.9–3.9 mm, length/width ratio = 2.8–2.9:1. Five inner, five outer gradates. Hindwing: length 9.6 –10.3 mm, width 2.8–3.1 mm, length/width ratio = 3.3–3.4:1. Four to five inner, four to five outer gradates. Material examined. Allotype ♀: Brazil. Minas Gerais. Tres Pontas [21o25’S / 45o30’W, 900 m], 15.xi.2008, Ferreira C. S Leg (MZUSP); Paratypes: Brazil. Minas Gerais. Tres pontas [21o25’S / 45o30’W, 900 m], 13.ix.2008, 1♂, 1♀, Ferreira C. S Leg (SFCC); Same, 13.ix.2008, 1♂, 1♀, Ferreira C. S Leg (SFCC), head clari- fied, without abdomen; Same, 20.ix.2008, 1♂, Ferreira C. S Leg (SFCC); Same, 4.x.2008, 8♂♂, 5♀♀, Ferreira C. S Leg (SFCC); Same, 18.x.2008, 1♂, Ferreira C. S Leg (SFCC); Same, 25.x.2008, 2♂♂, 1♀, Ferreira C. S Leg (SFCC); Same, 10.i.2009, 4♂♂, 3♀♀, Ferreira C. S Leg (SFCC); Same, 13.vi.2009, 1♀, Ferreira C. S Leg (SFCC); Same, 11.iv.2009, 1♂, Ferreira C. S Leg (SFCC); Same, 25.iv.2009, 1♂, Ferreira C. S Leg (SFCC); Same, 8.viii.2009, 1♂, 1♀ (UCOB); Same, 6.vii.2009, 1♂, Ferreira C. S Leg (MIZA); Same, 13.ix.2008, 1♀, Ferreira C. S Leg (MIZA). Etymology. The name refers to the type locality “Tres Pontas, Minas Gerais, Brazil”

16 · Zootaxa 3351 © 2012 Magnolia Press SOSA Species relationships. Titanochrysa trespuntensis sp. nov. is the only species in the genus with an protuberance on the apical margin of S8+9, wings without black markings on membrane, and wide gonapsis with acute lateral projection. On the male, there are setae beneath the arcessus as in Titanochrysa circumfusa (Burmeister) comb. nov., but T. trespuntensis males do not have X-shaped rods on the arcessus, or striations on the apical surface of the arcessus. Externally, T. trespuntensis resembles some Ungla species, but it can be distinguished from those species because its inner gradates meet the Psm vein and the males have a gonapsis. Geographical distribution. Brazil.

FIGURE 17. Mandibles. A) Titanochrysa circumfusa (Burmeister) comb. nov.; B) Titanochrysa pseudovaricosa (Penny) comb. nov.; C) Titanochrysa ferreirai Sosa & Freitas sp. nov.; D) Titanochrysa trespuntensis Sosa & Freitas sp. nov.

Key to species of Titanochrysa Sosa & Freitas gen. nov.

1 Microtholi absent from S2–8; forewing with membrane surrounding crossveins (except costal crossveins) infuscate; mandibles symmetrical; male: dorsal rods of arcessus X-shaped, membrane beneath arcessus with two lines of setae mesally ...... Titanochrysa circumfusa (Burmeister, 1839) comb. nov. (Figs. 1–4) 1’ Microtholi present on S2–8; membrane around crossveins with or without infuscate markings; mandibles symmetrical or asymmetrical; males: dorsal rods of arcessus parallel, membrane beneath arcessus with or without two lines of setae mesally2 2 Forewing membrane entirely without shading; mandibles symmetrical; male: membrane beneath arcessus with two lines of small setae mesally ...... Titanochrysa trespuntensis Sosa & Freitas sp. nov (Figs. 13–16) 2’ Forewing membrane surrounding inner gradates with infuscation; mandibles asymmetrical; male: membrane below arcessus without small setae mesally...... 3 3 Forewing with network of short, dark, reticulate veins covered by a black spot on medial area; male: arcessus with dorsal rods short, merging apically; female: seventh sternite 3.5 times longer than wide ...... Titanochrysa pseudovaricosa (Penny, 1998) comb. nov. (Figs. 5–8) 3’ Forewing with membrane around inner gradates infuscate, but without reticulate network of veins; male: arcessus with sclero- tized dorsal rods long, merging basally; female: seventh sternite 2.4 times longer than wide ...... Titanochrysa ferreirai Sosa & Freitas sp. nov. (Figs 9–12)

Acknowledgments

The authors thank: Dr. Norman D. Penny for making the type series of Ceraeochrysa pseudovaricosa Penny, 1998 available for study; Dra. Catherine Tauber for her valuable collaboration and suggestions, as well as for providing the identification of Titanochrysa circumfusa and unpublished information on the species type; Dr. James Becnel for review of the English and his useful comments and suggestions; Dra. Ariadne Chloe Furnival for review of the English; Dr. Nelson Wanderley Perioto, Dra. Rogéria Inês Rosa Lara (APTA, Ribeirão Preto, Brazil), Dr. Francisco Díaz (UCOB), M.Sc. Daniell R. R. Fernandes (FCAV-UNESP) for valuable collaboration; and the Observatorio Nacional de Ciencia, Tecnología e Investigación (ONCTI, the Bolivarian Republic of Venezuela) for financial support.

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