PROC. ENTOMOL. SOC. WASH. 110(2), 2008, pp. 417–438

GONZAGA NIGRICEPS (MCLACHLAN) (: ): DESCRIPTIONS OF LARVAE AND ADULTS, BIOLOGICAL NOTES, AND GENERIC AFFILIATION

CATHERINE A. TAUBER,GILBERTO S. ALBUQUERQUE, AND MAURICE J. TAUBER

(CAT, MJT) Department of Entomology, Comstock Hall, Cornell University, Ithaca, NY 14853, U.S.A. (e-mail: [email protected]); (GSA) Laborato´rio de Entomologia e Fitopatologia, CCTA, Universidade Estadual do Norte Fluminense, Campos dos Goytacazes, Rio de Janeiro, Brazil 28013-602 (e-mail: [email protected])

Abstract.—The larvae (three instars) of Gonzaga nigriceps (McLachlan, 1868) from Brazil are described; adults are redescribed (including male and female genitalia); and biological notes are provided. The morphology of both the larvae and adultsofG. nigriceps indicates that the species could fall within Gonzaga or ,a closely related leucochrysine genus. We retain its placement in Gonzaga, but we question the ultimate validity of the genus.

Resumo.—As larvas (treˆsı´nstares) de Gonzaga nigriceps (McLachlan, 1868) do Brasil sa˜o descritas, adultos sa˜o redescritos (incluindo a genita´lia do macho e da feˆmea) e informac¸o˜es biolo´gicas sa˜o fornecidas. A morfologia das larvas e adultos de G. nigriceps indica que esta espe´cie pode ser incluı´da tanto em Gonzaga como em Leucochrysa,geˆnero de proximamente relacionado. Sua inclusa˜o em Gonzaga e´ mantida, mas a validade deste geˆnero e´ questionada. Key Words: Leucochrysini, adult, larva, morphology, biology, Brazil, distribution, asymmetry

As currently constituted, the New there are no reports on the biological World, largely Neotropical, chrysopid characteristics of any species within the tribe Leucochrysini (Chrysopidae: sub- genus. Moreover, the genus is only family ) contains six genera weakly distinguished from other leuco- (Tauber 2007). Of these, Gonzaga Nava´s, chrysine genera (Penny 2001); as a con- with eight described species, is the second sequence questions persist concerning its largest. Gonzaga specimens are infre- systematic status. quently encountered in the field or in The lack of information on Gonzaga collections. And, except for its inclusion larvae is particularly regrettable. Recent in the generic review of the green studies encompassing a wide range of lacewings of the world (Brooks and chrysopid taxa reveal that larval mor- Barnard 1990), the genus has not re- phology often provides exceptionally ceived detailed, modern systematic treat- distinctive, stable characters for differ- ment (Table 1). The male genitalia of entiating chrysopid genera and for ana- only one Gonzaga species and the female lyzing their taxonomic relationships genitalia of another have been described; (e.g., Tauber 1974, 2003, 2004; Dı´az- Gonzaga larvae are undescribed; and Aranda and Monserrat 1995; Tauber Table 1. Catalogue of Gonzaga species and available information. 1 RCEIG FTEETMLGCLSCEYO WASHINGTON OF SOCIETY ENTOMOLOGICAL THE OF PROCEEDINGS 418

Gonzaga Species Original Description Subsequent Feferences Type Specimen* Larv ae Known Distribution amabilis Nava´s 1932: 32 Penny 1977: 22 prob. Univ. Torino, Unknown Ecuador (as Gonzaga ) Brooks and Barnard 1990: 276 Italy callipterus Banks 1944: 33 Penny 1977: 22 MCZ Unknown Brazil, Guyana (as Gonzaga ) Brooks and Barnard 1990: 276 nigriceps McLachlan 1867: (as Leucochrysa ): BMNH Described here Bolı ´via, Brazil, Ecuador, 251 (as Chrysopa ) Nava´s 1913a: 303 Guyana, Surinam, Nava´s 1913b: 96, 104 Venezuela (as Gonzaga ): Banks 1944: 34 Penny 1977: 22; Penny 2001: 13 Brooks and Barnard 1990: 247, 276, (female terminalia illustrated), this publication notatus Nava´s 1929: 861 Penny 1977: 22 Unknown** (Dresden? Unknown Peru (as Gonzaga ) Brooks and Barnard 1990: 276Sa˜o Paulo?) [Penny’s 1977 listing of Brazil is unconfirmed.] palliatus Nava´s 1929: 860 Penny 1977: 22 Unknown** (Dresden? Unknown Brazil (as Gonzaga ) Brooks and Barnard 1990: 276 Sa˜o Paulo?) palliceps McLachlan 1867: (as Leucochrysa ): BMNH Unknown Brazil, Costa Rica, 251 (as Chrysopa ) Nava´s 1913b: 149; Penny 2001: 13 Guyana Brooks and Barnard 1990: 277 (as Gonzaga ): Penny 2002: 191, 306 (wings, head, thorax illustrated) soroanus Alayo 1968: 60 Penny 1977: 22 IZAC Unknown Cuba (as Gonzaga ) Brooks and Barnard 1990: 276 torquatus Nava´s 1913a: 318 (as Allochrysa ): BMNH Unknown Brazil, Guatemala, Type species (as Gonzaga ) Banks 1914: 624 Panama of Gonzaga (as Gonzaga ): Banks 1944: 34, 1945: 172 Penny 1977: 22 Brooks and Barnard 1990: 246, 277 (wing, male terminalia illustrated) Penny 2002: 191, 307 (head, thorax, wings illustrated)

* Abbreviations: MCZ, Museum of Comparative Zoology, Harvard University, Cambridge, MA; BMNH, The Natural History Museum, London; IZAC, Instituto de Ecologı ´a y Sistema´tica, Ministerio de Ciencia, Technologı ´a y Medio Ambiente, La Habana, Cuba. ** Additional information may be available in Phillip A. Adams’ extensive no tes which were bequeathed to the California Academy of Sciences; our reque st to see the pertinent sections was refused. VOLUME 110, NUMBER 2 419 and de Leo´n 2001; Tsukaguchi 1995; body, primary setae are designated with Tauber et al. 2000, 2001, 2006). an ‘‘S’’ followed by a number; primary In this paper, we describe the larvae setae that are associated with thoracic (three instars) of Gonzaga nigriceps sclerites are designated with a setal num- (McLachlan, 1868); we also redescribe ber. Body segments are referred to as T1 the adult, with emphasis on the previously to T3 (thorax) and A1 to A10 (abdomen). undescribed male genitalia. To examine Thus, e.g., T1-S1Sc1 is the first seta possible generic relationships, we compare associated with sclerite 1 on the protho- G. nigriceps larvae and adults with those rax. Notations for other abbreviations are of other leucochrysine species that have on the captions to the pertinent figures. received modern attention. Also, to The number of setae in rows on the foster research on the genus, we provide thorax and abdomen are reported to the a catalogue of published work on the nearest even number; these figures are known Gonzaga species, including refer- approximate because the left and right ences to descriptions and taxonomic sides are often asymmetrical. In general, changes, location of the type specimen, G. nigriceps setae are smooth. Occasion- and reported distribution of each species. ally some of the long setae on the lateral tubercles of the third instar thorax appear MATERIALS AND METHODS to have very small, sparse thorns (barely Larvae.—We collected six living G. visible under 2003 magnification). We nigriceps adults (two females) over three categorize these setae as smooth. years, at forested sites in northern Rio de Adults.—We examined adult speci- Janeiro State, Brazil. These adults were mens from several localities in Brazil’s provided a diet of yeast autolysate, Atlantic forest and Amazonia (see below), fructose, and honey (1:1:1 volumetric and we confirmed our identification of mixture) (e.g., Tauber 2004), and one G. nigriceps by comparing our specimens female oviposited. The larvae were with McLachlan’s type, which is depos- reared individually in 40 ml glass vials ited in The Natural History Museum, on a diet of moth eggs [Anagasta London (BMNH). Methods follow those kuehniella (Zeller)]. Specimens were described earlier (Tauber 2007). killed in KAAD solution (Stehr 1987) All measurements (larvae and adults) ,2 days after hatching (first instar) or were made with NIH ImageJ software moulting (second and third instars) and (http://rsb.info.nih.gov/ij/). Unless noted transferred to 95% ethyl alcohol for otherwise, voucher specimens (larvae preservation. Subsequently, representa- and adults) from our study currently tive larvae were cleared, stained and reside in the collection at the mounted as described in Tauber (2004). Universidade Estadual do Norte Flumi- Measurements of head capsule width nense and the research collection of M. J. were made across the widest part of the and C. A. Tauber. head and include the eyes; head length was Images of representative G. nigriceps determined along the midline from clyp- specimens (adults and larvae) are avail- eus to cervix. Body length was measured able on the MorphoBank website: http:// along the curved midlateral line of dis- morphobank.geongrid.org/permalink/?P105. tended specimens (tip of clypeus to tip of abdomen). Mandible length was mea- Gonzaga nigriceps (McLachlan) suredalongthecenterofthemandibles’ Chrysopa nigriceps McLachlan 1868: 251 curved dorsal surface. Terminology for [description]. larval morphology follows previous usage Gonzaga nigriceps: Banks 1944: 34 [spe- (Tauber 2004). For both the head and cies transferred to Gonzaga, new ma- 420 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

terial]; Penny 1977: 22 [distribution]; with lateral section absent; mesal section Brooks and Barnard 1990: 247 [three paired, not fused mesally, mottled figures: apex of female abdomen, brown to dark brown, with irregular spermatheca, subgenitale]. margins, confluent with postfrontal marking; extending longitudinally from Diagnosis.—Gonzaga nigriceps adults near mid-dorsal base of cranium almost have the features that are typical of to midregion of head; together with leucochrysine chrysopids – e.g., very long postfrontal mark, extending anteriorly antennae with long flagellar segments from mesal base of head to mesal edge (length:width , 3:1), dark mark on base of antennal base; base extending lateral- of stigma, and pseudomedia that curves ly ,1/5 distance to lateral epicranial toward the costa before merging with the suture. Anterior (5 frontal + interman- outer gradates (Adams and Penny 1986, dibular) marking dark brown, with Brooks and Barnard 1990). They are fused narrow base, becoming broader, distinguished from other leucochrysines expanding anteriorly; distolateral mar- by a black to very dark brown head (frons, gins approaching but not reaching mesal clypeus, labrum, vertex), pronotum and margins of mandible bases; distomesal mesonotum, distinct markings on the ends connected by thin bridge. Mandi- wings, and unique genitalia (Figs. 7–9). ble, maxilla dark amber basally, becom- Gonzaga nigriceps larvae are trash carriers ing dark brown distally. Labial palp pale with typical leucochrysine characteristics basally becoming darker distally; basal (e.g., elongate thoracic tubercles bearing segment very light brown with brown very long, smooth setae, body with basal ring; second segment white basal- numerous, long, hooked setae). The larvae ly, becoming dark brown distally; termi- are distinguished from other described nal segment dark amber. Gena white leucochrysines by distinctive head, tho- with small brown spot posterodorsally, racic and abdominal markings, unraised irregularly shaped mark ventrally. An- spiracles, and a unique pattern of thoracic tenna with scape white dorsally, light and abdominal setae. brown laterally; pedicel white basally, Distribution.—We have seen speci- becoming brown distally, flagellum am- mens from Brazil (Amazonian and At- ber. Ventral margins of cranium brown; lantic forests), Colombia (Amazonia), cardo dark brown, stipes light brown; Ecuador, and Surinam. The species is mentum white mesally, with large brown also reported from Venezuela, eastern mesal spot posteriorly, small mark at Bolivia, and Guyana (Banks 1944). base of cardo; light brown spot at base of palpiger. Larvae Clypeus straight anteriorly, with right- Third instar.—Body mostly white; angled corners; surface of corners slight- 6.77–8.06 mm long. Integument with ly rugose ventrally; dorsum with three uniform coating of fine, pointed spinules. pairs of apical setae (2 lateral, 1 sub- Setae smooth throughout, mostly very mesal). Setae S1–S12, Vx present, most long and slender, especially distally, relatively short, thin; secondary seta next golden, except a few terminal setae on to S12. Labial palp: distal annulation of abdominal lateral tubercles dark brown. second segment with one long lateral, Head (Figs. 1 and 2): White to cream; two long mesal setae; remaining annula- 0.98–1.12 mm wide; 0.74–0.84 mm long. tions with , seven setae; basal segment Mandibles 1.32–1.74 mm long; ratio of with one lateral, one dorsal, one ventral mandible length to head width 5 1.45– seta. Palpiger with one dorsal seta, two 1.53:1. Eyes black. Epicranial marking ventral setae. Mandible with lateral seta VOLUME 110, NUMBER 2 421

Fig. 1. Gonzaga nigriceps third instar, head, dorsal. ant, anterior (5 frontal + intermandibular) marking; epi, epicranial marking; post, postfrontal marking; S1–7, S11, S12, primary setae; Vx, three short posterior setae surrounding a pore. 422 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Cervix white, with pair of purple (grape)-colored (subcutaneous) marks anterolaterally, pair of globular brown marks laterally, ,5 small setae antero- laterally, pair of V-shaped brown spots ventrolaterally. Thorax (Fig. 3): White to cream with white lateral tubercles (LTs) bearing golden to dark amber setae (LS); dorsum with mottled, subcutaneous markings as in figure, purple (grape)-colored; setae light amber. Prothoracic Sc1 brown, with whitish longitudinal stripe near mesal margin. Venter unmarked except meso- thorax with mesal patch of small, irregu- larly-shaped brown spots. Legs with tarsus, tibia brown; femur white with pair of distal brown spots (one anterior, one posterior) ,one fifth length of femur, another pair of anterior and posterior dark brown spots at base (where femur is narrow); trochanter with three ventral (basal) peaks (basal axis, two condyles) dark brown; coxa with dorsal base (at condyle) dark brown to black, with brown basal spot posterolaterally. Lateral tubercles (LTs) sclerotized, arising from sclerotized bases, with spinules on surface only at base. LS long, very slender distally; apical LS mostly pointed; sub- apical and lateral setae hooked. Prothorax (T1): Anterior margin con- vex, extending well over base of head. Pronotum with row of three small setae anteromesal to base of LT. Dorsal scler- ites (Sc1) large, globular, light brown; S1Sc1medium length, thin, straight; S2Sc1 mixed within secondary setae, not defin- itively identified. Sc2 transparent (visible only with transmitted light, not shown on Fig. 2. Gonzaga nigriceps third instar, head, figure), ovate, ,1/4 length of Sc1, without ventral. co, cardo; cr, ventral margin of cranium, associated setae. LTs long, narrow, ex- heavily sclerotized; l.p., labial palpus; md, mandi- tending beyond basal one-third of mandi- ble; mx, maxillary stylet; pg, palpiger; S8–S10, primary setae 8–10; stp, stipes. bles, with ,80 LS (,20 apical, ,60 lateral, dorsal). Primary setae S1, S3, S5 near base. Scape with single lateral seta; intermediate size; S4 short; all smooth, pedicel with small lateral thorn at apex; pointed; numerous additional setae, S2 flagellar seta more than one-half length not discerned. Fourteen to twenty small of flagellum. secondary (smooth, pointed) setae. VOLUME 110, NUMBER 2 423

six small, hooked secondary setae. Poste- rior subsegment with each LT bearing ,55 LS (,15 apical; ,40 lateral, dorsal); Sc2 very small, transparent, on anterior margin, with two associated setae (S1Sc2, S2Sc2); Sc3 large, posteromesal to LTs, brown; S1Sc3 small; S2Sc3 absent; ante- rior region with row of ,17 small, mostly hooked setae; posterior region with trans- verse row (R1) of ,60 small to medium- length, thin, mostly hooked setae, some on small chalazae. S1, S2 unidentified. Metathorax (T3): Anterior sclerites (Sc1) transparent, very small; S1Sc1, S2Sc1 very small; mesolateral sclerites (Sc2) transparent; S1Sc2 very small, mesal to Sc2; LTs long, narrow, each bearing ,50 LS (,15 apical; ,35 lateral, dorsal); anterior row of ,20–23 small, mostly hooked setae; posterior, trans- verse row (R1) of ,45 short to medium- length, thin, hooked setae, some on small chalazae. S1 unidentified. Abdomen (Fig. 4): Dorsum, venter white, unmarked except small brown spots below LT3–LT7. Integument transparent; all spiracles flush with surface, not enlarged, light amber. Dor- sal setae (submedian setae, SMS; setae on laterodorsal tubercles, LDS) long, slender, hooked (A1–A6), straight (A7– A10). LTs of segments 4–7 heavily sclerotized; LS (especially apical ones) on A2–A7 becoming more robust poste- riorly. LDTs present on A1–A7, small on Fig. 3. Gonzaga nigriceps third instar, thorax, A4–A5; each LDT bearing two slender, dorsal. A, Prothorax. B, Mesothorax and meta- hooked LDS, one minute microseta. thorax. LS, setae on lateral tubercle; LT, lateral tubercle; R1, row of setae; S1, S3–5, primary setae; A1: LTs absent; LDTs small, lightly Sc1–3, primary sclerites; SxScx, setae associated sclerotized, LDS mid-sized, without mi- with sclerites; Sp, spiracle. Mesothoracic and crosetae. Dorsum with two transverse metathoracic sclerites 1 and 2 are very small and rows of submedian setae (SMS): short difficult to discern within the intersegmental fold. anterior row with six (two mesal, four lateral, smaller); longer posterior row of Mesothorax (T2): Anterior subsegment 20 to 22 long SMS between LDTs, single with amber, circular, unraised spiracles; slender, hooked seta lateral to LDT. SSp spiracular chamber not enlarged. Sc1 very absent. small, transparent, on anterior margin, A2–A3: LTs papiliform, with 4–6 with two associated setae, one pair on medium-length, robust LS apically and each side (S1Sc1, S3Sc1); row of two to anteriorly, ,12 smaller LS laterally; 424 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Fig. 4. Gonzaga nigriceps third instar, abdomen, dorsal. A, Segments 1 and 2. B, Segments 4–6. C, Segments 7–10. A1–A10, abdominal segments; LT, lateral tubercle; LDT, laterodorsal tubercle; Sp, spiracle; SSp, seta associated with spiracle; R1, R2, R3, rows of submedian setae. Note: Abdominal segment 3 not shown; segments 2 and 3 are very similar. VOLUME 110, NUMBER 2 425 terminal LS on A2 slightly smaller than A10: Dorsum bearing microsetae on those on A3. Dorsum of each segment membranous tip; three short, straight with three transverse rows of long SMS, setae laterally. as follows: anterior row with ,14; Second instar.—Body 4.1–4.7 mm second row with approx. eight; posterior long. Similar to the third instar, with row with ,20 SMS between LDTs; the following exceptions. single slender, hooked seta lateral to Head. 0.62–0.66 mm wide; 0.41– LDT. SSp very small, straight. 0.44 mm long. Mandibles 0.79–0.92 mm A4–A5: LTs each with two very long, long. Second segment of labial palpus dark brown apical LS, two to three with seven or eight setae: distal annula- medium-length LS apically, approx 12 tion with one long, two or three short, shorter, slender LS dorsally. Dorsum remaining annulations with four short. with LDTs present, but difficult to Thorax. T1: LT bearing 42–45 LS distinguish; three transverse rows of (,15 apical, ,27–30 subapical, lateral); SMS as in A2–A3, anterior and posterior 14–20 small secondary setae; Sc1 light rows with slightly more SMS, sometimes brown throughout. T2: LTs bearing 29– with hooked seta(e) lateral to LDT. SSp 32 LS (10 apical, ,19–22 subapical, small, pointed. lateral); anterior subsegment with no A6: LTs with three long, robust, dark secondary setae; anterior section of brown apical LS, two long, robust, posterior subsegment with ,3 small, amber apical LS, four to five shorter, mostly hooked setae; R1 of posterior amber LS apically, subapically. Dorsum subsegment with ,13 small to medium with three transverse rows of dorsal length, hooked setae, some on small setae: anterior row of ,8 slender SMS, chalazae. T3: LTs bearing 29–31 LS (10 mesal row of ,8 SMS, posterior row of apical, ,19–22 subapical, lateral); R1 ,14 SMS between LDTs. LDT with with ,14 short to medium length, robust, hooked seta, one medium-length, hooked setae. pointed seta. SSp short, pointed. Abdomen. A1: anterior transverse row A7: LTs with five long, robust, dark having one to two hooked setae; poste- brown LS, four to five shorter, amber LS rior row with eight long, slender to apically, subapically. Dorsum with three robust SMS between LDTs; no setae transverse rows of setae: anterior row of lateral to LDT. A2–A5: LTs each bear- three to four slender SMS, second row of ing , eight to eleven LS; dorsum of each five to six small SMS, posterior row of segment with three transverse rows of ,10 SMS between pronounced LDTs, long, hooked SMS as follows: anterior three hooked setae anterolateral to each row with ,four to six SMS; mesal row LDT. LDTs each with one robust LDS with four SMS; posterior row with stemming from large chalaza, smaller ,eight SMS between LDTs; LDTs not LDS stemming from side of LDT. SSp difficult to distinguish on any segment; short, straight. no setae lateral to LDTs. A6, A7: LTs A8: LTs small, broad, with ,six LS (2 each bearing ,four to five long, robust medium length, 4 smaller). LDT not LS, four to five shorter LS. Dorsum of apparent. Dorsum with two transverse A6 with three transverse rows of four rows of four and 13 small SMS. SSp hooked SMS (posterior row with SMS short, straight. between LDTs); no setae lateral to A9: Dorsum with three transverse LDTs. Dorsum of A7 with two trans- rows of small, straight setae (2 anterior, verse rows of setae (anterior row with 4 mesal, 6–8 posteromesal). Venter with four slender, pointed SMS, second row ,six setae in two vertical rows. with 2–3 small SMS between LDTs). A8: 426 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

LTs bearing three LS (1 medium length, 2 smaller); dorsum with two transverse rows of four and five small SMS. First instar.—Body 2.3–2.7 mm long, predominantly white, with purple (grape)- colored markings confined to thorax, A6– A10. Setae smooth, straight (head), straight or hooked (thorax, abdomen). Setae on thoracic and abdominal LTs dark brown (except on A2, A3); remaining setae amber to cream. Head (Fig. 5): 0.41–0.43 mm wide; mandibles 0.47–0.51 mm long. White, with brown markings. Epicranial mark- ing paired, narrow, not fused mesally, mottled, with irregular margins, extend- ing from base of head anteriorly to level of eyes, not bending toward antennal base, with base extending laterally ,one fourth distance to lateral epicranial suture. Anterior marking dark brown, with fused broad base extending antero- laterally, becoming narrower; distal marking approaching, but not reaching mesal margins of mandible; distal ends not connected. Postfrontal marking not distinguishable; intermandibular mark- ing absent. Gena white, unmarked. Primary cephalic setae present; S2, S4, S5, S6, short; S12 very short; S1, S11 long. Labial palpi cream to very light brown basally, amber distally; distal annulation of second segment with one long, one short seta; remaining annula- tions without setae; basal segment with two mid-length setae; distal segment without setae. Mandible, maxilla amber throughout, with thin, dark brown me- sal, lateral margins. Mandible with single lateral seta. Maxilla with very small setae apically. Antenna with scape white to very light brown, pedicel white basally, brown distally, flagellum light brown. Fig. 5. Gonzaga nigriceps first instar, head, Venter with anterior region reddish dorsal. ant, anterior (probably 5 frontal + inter- brown; cardo light brown; stipes light mandibular) marking; epi, epicranial marking; brown; base of mandibles bulbous, post, postfrontal marking; S1–7, S11, S12, primary white. setae; Vx, three short posterior setae surrounding Thorax: White, with light brown a pore. patches posteromesal to prothoracic VOLUME 110, NUMBER 2 427

LTs, mesal to meso and metathoracic row, arising from chalazae between LTs; integument above pigmented areas LDTs. A6: LTs relatively large. Two with spinules; LTs elongate, white, scler- transverse rows of setae: four medium otized, with rough surface, but without length, hooked SMS mid-segment, two spinules. LS smooth, pointed, dark shorter, hooked SMS on chalazae be- brown. Prothorax (T1): LTs each with tween LDTs. A7: similar to A6, except two long LS, one very small, pointed setae smaller. LDT with one long, microseta mesally. Anterior region with robust, pointed LDS, one shorter point- row of three very small setae anteromesal ed LDS laterally on LDT. A8: LTs each to base of LT. Sc1 large, very light with two LS: one short, the other of brown; Sc2 not observed. S1Sc1 long, intermediate size, both straight, pointed. arising from chalaza. S1 long, S3, S4, S5 Dorsum with single pair of SMS. A9: 6 medium length, pointed; S2 absent. pairs of setae straight, pointed (four in Mesothorax (T2): Spiracles flush with anterior row short, two in posterior row integument, chamber unenlarged. LTs medium length). A10: three pairs of each bearing three long LS, no micro- straight, pointed setae, laterally. setae. Sc1, Sc2 small; associated setae Larval material examined.—Fifteen (S1Sc1, S1Sc2, sometimes S2Sc1, S2Sc2) specimens reared and preserved at the present, very small. Sc3 large, transpar- Universidade Estadual do Norte Flumi- ent; S1Sc3 short to mid-sized. Posterior nense, Campos dos Goytacazes, RJ, and row with four setae, lateral-most large, at the University of California, Davis [N hooked, arising from chalazae. Metatho- 5 7 (L3), 4 (L2) and 4 (L1)], from female rax (T3): LTs each with three long LS, collected in Brazil, Rio de Janeiro State, no microsetae. Sc1 elongate transversely, Desengano State Park, near Campos dos narrow, with associated setae (S1Sc1, Goytacazes (Fazenda Babiloˆnia), Oct 26, S2Sc1, S3Sc1) very small, in transverse 2003, G. S. Albuquerque, M. J. Tauber, row; Sc2 not discerned. Posterior row C. A. Tauber, Collectors. (R1) with four long, hooked setae on Species-specific characteristics of G. chalazae. Legs setose; mostly white; nigriceps larvae.—To date, the larvae of coxae brown basally, dark brown at five leucochrysine species have been coxopleural articulation; tibiae light described in detail: all are in the genus brown, with brown band basally and Leucochrysa McLachlan, four in subge- near distal margin; tarsi light brown with nus Nodita Nava´s and one in subgenus dark brown tip; claws amber; empodia Leucochrysa McLachlan (Tauber 2004, dark brown. Mantoanelli et al. 2006). Each of the five Abdomen: LTs present on A2 to A7, species has distinctive head markings each with two long, hooked LS, no that differ from those of G. nigriceps, microsetae; LS on A2, A3 amber, those and the numbers of thoracic and abdom- on A4–A7 dark brown. LTs absent from inal setae of each of the five species differ A1. LDTs present on A1–A6, each with from each other and from those of G. two long, hooked LDS, one short, nigriceps. In addition, unlike L. (L.) pointed microseta. Spiracles present on varia (Schneider), G. nigriceps (and the A1–A8, each with slender, pointed SSp four Nodita species) do not have raised, (except on A1). A1: Dorsum with single tuberculate spiracles (all instars). row of four long, hooked SMS arising Among the described leucochrysine from chalazae between LDTs. A2–A5: species, G. nigriceps most closely resem- Dorsum of each segment with two bles L. (N.) pavida (Hagen) and L. (N.) transverse rows of long, curved, hooked americana (Banks). All three species SMS: four in first row; two in second share a dark intermandibular marking 428 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Fig. 6. Gonzaga nigriceps adult, cranial appendages. A, Labium (ventral view; right palp and palpiger removed). B, Maxilla (left, ventral view). bg, basigalea; co, cardo; ga, galea; lc, lacinia; lg, ligula; l.p., labial palpus; m, mentum; m.p., maxillary palpus; pg, palpiger; sm, submentum; stp, stipes. that connects the frontal markings ante- determine if one side of the larva or the riorly, a single round, dark marking at other consistently had the higher num- the base of the mentum, pigmented spots ber. This observation of asymmetry in on the venter of the mesothorax and setal number is consistent with those metathorax, and darkly delineated distal from other chrysopid taxa (e.g., Man- femoral margins. In addition L. (N.) toanelli et al. 2006). However, it is pavida and G. nigriceps both have rela- noteworthy that unlike L. (L.) varia (in tively dark prothoracic sclerites (Sc1). which the right mandible is generally However, G. nigriceps larvae differ from longer than the left), the G. nigriceps L. (N.) pavida and L. (N.) americana larvae did not exhibit a consistent larvae in that there are 2, 2, 3 and 5 very pattern of asymmetry in mandible dark, long setae on the lateral tubercles length. The significance, if any, of the of segments A4, A5, A6, and A7 mandibular asymmetry in L (L.). varia respectively [in L. (N.) pavida and L. and its absence in G. nigriceps is un- (N.) americana, the long lateral setae are known. all amber]. In addition, the legs of L. (N.) americana larvae lack markings Adults [which are present on both L. (N.) Description.—Head (Fig. 6): 1.40– pavida and G. nigriceps], and the lateral 1.60 mm (male) – 1.59–1.82 mm (female) tubercles of the G. nigriceps abdomen are wide (including eyes); ratio head width to not elongated apically as they are on L. eye width 5 1.52–2.10:1. Vertex oval, (N.) pavida. raised, flat, with grainy surface, numer- Note on asymmetry in larvae.—We ous short, dark amber setae, posteriorly observed that most rows of submedian with pair of oval, grainy protuberances setae on the dorsum of the abdomen had surrounded in front with dense amber an uneven number of setae; we did not setae. Antenna almost twice length of VOLUME 110, NUMBER 2 429 wings (25.9–28.6 mm); scape large, lon- broad inner expansion, left one with ger than broad, (0.37–0.44 mm long, rounded, thumb-like lobe on inner mar- 0.34–0.35 mm wide), width 5 0.22–0.23 gin; both with row of short setae on crest times distance between antennae, with near inner margin. long pale setae dorsally, ventrally, some Head coloration: Antenna white to long (,0.2 mm); distal margin fairly cream, unmarked. Eyes black (holotype). straight; pedicel ,0.12 mm long, Vertex dark brown to almost black, with ,0.14 mm wide (at widest point); prox- cream, longitudinal mesal spot, cream imal flagellomeres long, narrow (seg- immediately mesal, posteromesal to eyes. ments 1, 2, 3: length 5 2.0–2.2 times Frons dark brown throughout. Torulus width), with four concentric rings of cream frontally, dark brown to black setae; middle and distal segments slightly near dorsal margin of antennae, dark longer (segments 6–8: length 5 2.5–2.9 brown distally on dorsal surface. Genal times width), also with four concentric area dark brown, slightly lighter ventral- rings of setae. Distance between anten- ly. Clypeus dark brown throughout. nae small (0.06–0.09 mm); distance be- Labrum dark brown, with small cream tween tentorial pits large (0.43– area distally. Maxilla with basal four 0.56 mm); length of frons (mid-antenna palpomeres light brown to brown basal- – midway between tentorial pits) large ly, cream distally; distal palpomere (0.36–0.49 mm). Torulus large (below cream. Labium, labial palp cream. scape, 0.10–0.14 mm deep). Frons raised Thorax: Cervix collar-like, quadrate, mesally, with prominent large, scalloped, concave anteriorly, dark brown to black outward fold above clypeus, below tor- dorsally, ventrally, with numerous short, uli; surface rough. Clypeus arcuate ba- dark setae. Prothorax 0.74–0.97 mm sally, 0.17–0.27 mm long; surface hori- long; 1.20–1.40 mm wide; ratio, length: zontally striated. Labrum with distal width 5 0.57–0.75:1; setae thin, long margin indented mesally, dorsal surface (longest 0.34–0.50 mm), dark brown to smooth, rounded; sides rounded. Gena golden; pronotum dark brown through- short; ratio, genal length to distance out, to dark brown with cream longitu- between tentorial pits 5 0.31–0.34:1. dinal stripe mesally, pair of small round, Maxilla with galea large, extending well cream spots sublaterally; pleuron, venter beyond lacinia, with small, narrow, white. Mesothorax, metathorax brown straight terminal papilla, distal one half dorsally, white laterally, ventrally. Legs evenly covered with short straight setae; white, unmarked; distal two tarsomeres lacinia approximately as wide as galea, dark brown. Claws simple, with elongate tapered distally, heavily sclerotized me- narrow dilation, small tooth at tip of sally, with patch of heavy anteriorly dilation; terminal setae approximately directed setae, distal ones curved and same length as claws. relatively long; palps tapered distally. Wings (Fig. 7): Forewing: 14.9– Labium with narrow submentum, sparse, 19.2 mm long, 5.4–7.0 mm wide. Costal thin setae; ligula short, broad-based, area not exceptionally broad; tallest with sparse setae ventrally; subapical costal cell (#9) 1.2–2.7 mm tall, 2.2–5.4 transverse depression below distal patch times width. First intramedian cell quad- of setae, distal patch of setae dense, rate, 0.92–0.96 width of third median short, thin, directed mesally, forming a cell. First radial crossvein distal to origin mesal peak; distal patch of setae contig- of radial sector (Rs); radial area (be- uous with dorsal patch (not shown in tween Radius and Rs) with single row of Fig. 6); palps tapered distally. Mandibles 14–16 cells; tallest cells 1.6–2.4 times narrow, asymmetrical, right one with longer than wide. First mediocubital 430 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Fig. 7. Gonzaga nigriceps forewing (top), hindwing (bottom). b1, b5, upper Banksian cells; b’1, b’5, lower Banksian cells; icu1–icu3, intracubital cells; im1, im2, intramedian cells; i.g., inner gradate veins; m3, third medial cell; o.g., outer gradate veins; Psm, pseudomedia; R, Radius; Rs, radial sector; r1, first radial vein; rx1, first radial crossvein; 2, 3, second and third cells bounded by gradates; 6, 7, sixth and seventh cells beneath Rs (5 first two cells beneath Rs, free of Psm). crossvein crassate (male and female); 4–5 median cell); last b’ cell 0.6–1.0 times b cells (5 cells beneath Rs, not including taller than wide. Three intracubital cells an inner gradate vein); distal b cell 0.7– (two closed); first cubital crossvein lo- 0.9 times wider than tall. First two cells cated near second mediocubital cross- beneath Rs, free of pseudomedia (5 vein; intracubital cells 1–3 of similar sixth and seventh cells beneath Rs) 2.1– lengths. Vein 1A forked. Membrane 2.6 times taller than wide. Two series of clear, with base of wing between sub- gradate veins; 7–8 inner gradates, 8–10 costa and anal vein dark brown; stigma, outer gradates; second and third cells radial area below, dark brown; brown bounded by gradates 1.7–3.3 times lon- mark extending distally along Rs ger than wide. Five to six b’ cells (cells through last radial cell and cells below; beneath pseudomedia after second intra- base of Rs with dark brown mark; VOLUME 110, NUMBER 2 431 second and third to last b cells with dark view) triangular in shape, with rounded brown spots basally; distal b cell mostly apices. Setae on S9 long, robust, stem- ringed with brown; distal one third of ming from heavy bases; terminus of S9 pseudocubitus with brown mark extend- with field of gonocristae. Gonarcus V- ing through several outer gradates. Veins shaped with apodemes rounded apically, pale; setae pale to amber. broad, attached distally, extending be- Hindwing: 13.1–17.6 mm long, 3.8– yond gonarcal bridge; apex with blunt 5.1 mm wide. Two series of gradate mesal extension abutting dorsal edge of veins; 6–7 inner, 5–8 outer; 13–14 radial mediuncus. Gonocornua absent. Mediun- cells (counted from origin of Radius, not cus cylindrical; dorsal surface sclerotized, false origin). Four to five b cells (includ- with longitudinal striations; tip beak-like, ing small ‘‘b1’’ cell); four-five b’ cells pointed; ventral surface rounded, with beyond second intramedian cell; three subapical, lip-like extension. Entoproces- intracubital cells (two closed). Mem- sus arm-like, extending dorsolaterally brane clear, with stigma, area below, from side of gonarcal bridge, below base dark brown; distal two radial cells of mediuncus. Membranous skirt below margined with dark brown; distal vein mediuncus flat, lightly striated horizon- of last b’ cell sometimes with dark brown tally, with two gonosetae on chalazae. mark. Veins pale; setae pale to amber. Area below gonarcal complex membra- Abdomen: Segments with pleural re- nous, bearing elongate, rounded, internal gion ,one half depth of sternites; distal pouch with heavy base; pouch attached to segments expanded at least slightly. distal margin of S9. Tignum, gonapsis, Integument mostly white; tergites 5–8 pseudopenis, spinellae absent. Hypan- dark brown; tergite 3 with small brown drium internum V-shaped, lightly sclero- mark posteriorly; venter with tinge of tized, attached distally to membrane brown on segments 5–8; gonapophyses beneath gonarcal complex; comes cres- laterales, callus cerci cream; setae pale. cent-shaped. Sternites , same height as length, with Female (Fig. 9): Callus cerci almost microsetae relatively sparse, small. Ter- circular, 0.17–0.19 mm maximum diam- gites narrow, caplike, with heavier, eter, with 32–37 relatively short tricho- denser setae and microsetae than ster- bothria (0.04–0.11 mm long). Tergite 8 nites. Spiracles oval externally; atria not quadrate (lateral view), deeper ventrally enlarged. than T7 or T6. Tergite 9+ectoproct Male (Fig. 8): Callus cerci oval, 0.17– elongate, distal (dorsal) margin angled 0.23 mm maximum diameter, with 24–34 ventrally; margin quadrate ventrally, not relatively long, thin trichobothria (0.075– extending below gonapophyses laterales. 0.31 mm long). Sternites 3–8 with micro- Sternite 7 with distal half tapering tholi (perhaps S1 and S2 also). Dorsum of abruptly, broad and rounded distally; T9+ectoproct truncate distally, fused me- terminus unmodified, with dense, long sally, midline without deep cleft, distal setae. Gonapophysis lateralis narrow, setae robust; ventral section of T9+ecto- rounded. Colleterial gland large, extend- proct with rounded proximal extension ing to end of abdominal segment 6, with reaching less than half length of T8; bulbous reservoir entirely within T9+ec- proximal margin well sclerotized. Dorsal toproct; duct between gland and reser- apodeme faint, extending below and voir relatively thick. Transverse sclerifi- around proximal margin of callus cerci. cation well sclerotized, broadly U- S8+9 fused, although clearly distinct, shaped (posterior view), tapering lateral- without suture; S9 much smaller than ly, with long, fine longitudinal ridges. S8, without microtholi; S8+9 (lateral Bursa copulatrix membranous, loosely 432 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Fig. 8. Gonzaga nigriceps male. A, Terminal abdominal segments, lateral. B, Gonarcal complex, lateral, withdrawn. C, Gonarcal complex, lateral, extended. D, Gonarcal complex, dorsal, extended. c, comes; c.c., callus cerci; ent, entoprocessus; gcr, gonocristae; gs, gonarcus (lateral apodeme); gse, gonosetae; g.b., gonarcal bridge (extension); h.i., hypandrium internum; mu, mediuncus; pouch, pouch below gonarcal complex, above tip of S8+9; S8+9, fused eighth and ninth sternites; sr.m., subrectal membrane; T9+ect, fused ninth tergite and ectoproct (dorsal apodeme faint, not shown). folded, extending posteriorly over sper- with mesal invagination extending entire matheca almost to gonapophyses later- depth. Spermathecal velum sail-shaped, ales, connected ventrally to spermatheca expanded and truncate distally. Sper- via very long slit along dorsal surface of mathecal duct attached to dorsal margin spermathecal velum. No bursal glands of spermatheca, heavily sclerotized, long found. Spermatheca round, doughnut (over twice length of spermatheca), shaped (,three-fourths length of T9+ect), curved before U-shaped, distal bend VOLUME 110, NUMBER 2 433

Fig. 9. Gonzaga nigriceps female. A, Terminal abdominal segments, external, lateral. B, Same, internal, lateral. b.c., bursa copulatrix; col.g., colleterial gland; col.r., colleterial reservoir; c.c., callus cerci; g.l., gonapophysis lateralis; inv, spermathecal invagination; sg, subgenitale; sp, spermatheca; s.d., spermathecal duct; S7, seventh sternite; t.s., transverse sclerification; T8, eighth tergite; T9+ect, fused ninth tergite and ectoproct; vel, velum. within subgenitale; terminal ca. one tenth terminus with sclerotized, bilobed pro- enlarged, setose. Subgenitale relatively cess, with lobes separated by flat trough, narrow, tapering and extending well extending distally, nestled between gona- beyond lower margin of T9+ectoproct, pophyses laterales; membrane above containing small part of spermathecal lobes with crescent-shaped, very lightly duct; small ventral fold without setae; sclerotized lamellae. 434 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Notes on type.—McLachlan (1868) cliff’; macaos (5 araras) nest in such identified ‘‘Ega, Brasilia’’ as the type places], R. Solimo˜es, 10/11/74 in forest’’; locality and ‘‘Bates’’ as the collector; he second label with a relatively lengthy did not indicate how many specimens he description of the specimen. (2) A male worked with. One specimen with this with one label citing an undeciphered information remains in BMNH; we locality in Ecuador. The nine other regard it as a syntype. It is a male with specimens are from Brazil, Colombia, one flagellum missing, the other broken; and Surinam. Brazil, RJ, Desengano right forewing missing; abdomen cleared, State Park, Fazenda Babiloˆnia (21u519S, unstained, in glycerin, segment 4 onward 41u489W), Mar 21, 2001 (M; M. J. Tauber, present, genitalia separated from abdo- C. A. Tauber, G. S. Albuquerque), Oct 26, men; hypandrium internum missing. 2003(F;G.S.Albuquerque,M.J.Tauber, Labels on the specimen read: 1. ‘‘Type’’ C. A. Tauber; lab-reared M, Tauber Lot (within red circle). 2. ‘‘Ega, Bates’’ (light 2003:40); Desengano State Park, Sta. blue). 3. Small green label with no M. Madalena, Terras Frias (21u559S, writing. 4. ‘‘Chrysopa nigriceps McL. 41u559W), Oct 28, 2003 (M; G. S. Albu- (type)’’ (light blue). 5. McLachlan Coll. querque, M. J. Tauber, C. A. Tauber); Sta. B.M.1938-674 (light blue). The locality M. Madalena, Fazenda Santo Antoˆnio de ‘‘Ega’’ (currently Tefe´) was Mr. Bates’ Imbe´(22u009S, 41u529W), Oct 28, 2003 (F; headquarters during part of his extended G. S. Albuquerque, M. J. Tauber, C. A. (1848–1859) excursion to the Amazon Tauber); Conceic¸a˜o de Macabu, Fazenda region; it is located on the Tefe´ River at Carrapeta (22u109S, 41u529W), Feb 28, its confluence with the Upper Amazon 2002 (F, M; G. S. Albuquerque). Colom- (R. Solimo˜es). bia, Amaz. Letı´cia, Feb. 1–3, 1969 (F; W. The unstained syntype male and the D. Duckworth; United States National male specimens that we studied from Museum). Surinam, Feb 1982 (teneral northern Rio de Janeiro State resemble female reared from larva collected by J. each other very closely. However, there Carpenter, D. Smith; Cornell University was some noteworthy variation. The Insect Collection). most obvious involved the number of gonosetae on the stiff membranous Biology ‘‘skirt’’ below the mediuncus. The syn- Eggs were globular (not elongate) in type bears nine gonosetae, whereas the shape, and they were deposited at the specimen from Terras Frias that we end of very long, flexible stalks. When dissected has only two. Also, the gonar- oviposited, they were dark bluish green; cus of the syntype is considerably nar- they became darker and hatched within rower than that on any of our specimens; 6–8 days (N 5 4) (ambient room tem- we suspect that it remained flattened perature). Larval developmental times after being held beneath a coverslip for a were L1: 3–5 days (N 5 4), L2: 5 days long period. (N 5 2), L3: 5–6 days (N 5 2). Two Adult material studied.—Syntype (data adults (both males) emerged in good above), two additional specimens from condition and survived for several weeks. the McLachlan Collection (BMNH), and The larvae carried trash – pieces of plant nine more recently collected specimens. and prey material that they encountered The two McLachlan specimens are from in the vial; they did not construct dense the Upper Amazon, Brazil, and Ecuador: packets (a pattern that is typical of lab- (1) A female with one label reading reared leucochrysine larvae). In general, ‘‘Barr(–?) das Araras (?) [probably ‘Bar- their behavior in the laboratory resem- ranco’ or ‘Barranca,’ which mean ‘small bled that of the Leucochrysa larvae and VOLUME 110, NUMBER 2 435 adults that we have reared and main- proposed a set of eleven features that tained under similar conditions. may distinguish Leucochrysa larvae from other trash-carrying chrysopids in the Comparison of Gonzaga nigriceps with New World. Our study here indicates Leucochrysa species that G. nigriceps larvae share the entire Within a year after Nava´s (1913a) set of eleven character states, with only proposed the genus Gonzaga, Banks minor variation (as noted below): (1914) stated his reluctance to accept (a) Thorax: Each segment with pair of the genus as valid, and he synonymized very long, sclerotized lateral tuber- the name with Allochrysa Banks (now 5 cles [i.e., the prothoracic lateral Leucochrysa). Later, he reversed his tubercles extend anteriorly at least opposition, reinstated the generic desig- to the midregion of the head; the nation, and used it in several papers (e.g., meso- and metathoracic tubercles Banks 1944, 1945). By the time of are longer than half the width of Brooks and Barnard’s (1990) review of the respective segment] (Fig. 3). All the world’s chrysopids, the genus name three instars. was commonly accepted (e.g., Penny (b) Thorax: Lateral tubercles with very 1977; Adams and Penny 1986). The worldwide review and later work (e.g., long apical and lateral setae (all Penny 2002) reaffirmed the opinion that instars), apically arranged in a fan- Gonzaga is distinct from (although close- shape (second and third instars) ly related to) Leucochrysa; but doubts (Fig. 3). that had been raised earlier (e.g., Penny (c) Body: Anterior abdominal segments 2001) were not addressed. (A1–A5) thicker (dorsum to venter) Here, we again re-open the question of than thoracic segments, but without Gonzaga’s validity as a genus, and we an abrupt increase in height as is suggest that the currently available larval typical of Chrysopodes Nava´s (meta- and adult morphological data do not lend thorax and A1) and Ceraeochrysa strong support for maintaining Gonzaga Adams (A1 and A2) (Tauber et al. as a genus distinct from Leucochrysa. 2000; Tauber 2003). All three instars. Larvae.—Like a large number of (d) Abdomen: Segments A2 and A3 with taxonomically diverse chrysopid species lateral tubercles papiliform, bearing with larvae that carry large packets of long setae on anterior, dorsal and/or trash on their dorsa, G. nigriceps has apical surfaces (Fig. 4). In the four larvae with gibbous, setose bodies, elon- Leucochrysa (Nodita) species de- gate thoracic tubercles, well-developed scribed from the U.S.A., the long abdominal tubercles, and elongate, ser- setae are on the anterior surface; in L. rated and/or hooked setae. In addition, (L.) varia and G. nigriceps,mostof G. nigriceps larvae also express all of the the long setae are dorsal and apical. traits that have been shown to typify the Second and third instars. trash-carrying Leucochrysa larvae (Smith (e) Abdomen: Segments A4–A7 with 1926; Skorepa and Sharp 1971; Adams lateral tubercles slightly elongated, 1987; Tauber 2004; Mantoanelli et al. bearing long setae arising mainly 2006). from apex. All three instars. In all of Based on comparative study of the the species the lateral tubercles on larvae of five species of Leucochrysa A2 are the most broadly based and (four in the subgenus Nodita and one in rounded, and the lateral tubercles the subgenus Leucochrysa), Tauber become more elongate on subse- (2004) and Mantoanelli et al. (2006) quent segments (Fig. 4). 436 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

(f) Abdomen: Segments A7–A10 small, tubercles is similar to that in the curved ventrally and partially with- Leucochrysa species that have been drawn into each other. All three described to date, except that it instars. contains several more setae and, in (g) Antenna: Terminal seta at least K the L3, a single, smooth, hooked length of flagellum (Figs. 1, 5). All seta slightly lateral to each LDT. three instars. (k) Abdomen: Segments A2–A5 each (h) Thorax: Primary prothoracic setae with three (L3, L2) or two (L1) S1, S3, S4, S5 present; S2 absent dorsal rows of smooth, hooked (Fig. 3). All three instars. This submedian setae (SMS), as follows. character state holds for the five In each case the posterior row of species of Leucochrysa (both sub- SMS lies between the two laterodor- genera). However it is difficult to sal tubercles. interpret in the third instar of G. nigriceps; the pronotum of the G. N L3: Row 1 with 6–14 SMS; Row 2 nigriceps L3 bears several very small with 4–8 SMS; Row 3 with 6–26 SMS setae; whether all of these setae are (Fig. 4). secondary or one of them is homol- N L2: Row 1 with 2–6 SMS; Row 2 with ogous with S2 is unknown. 4 SMS; Row 3 with 2–10 SMS. (i) Abdomen: Segments A1–A5 each N L1: Row 1 with 4 SMS; Row 2 with 2 with pair of laterodorsal tubercles SMS. bearing two long, smooth, hooked setae (LDS) and a single small, Thus, our detailed study of the larval pointed microseta (Fig. 4). All three morphology of G. nigriceps did not offer instars. Note: In G. nigriceps, the any characters that would support the microsetae are very small, but they separation of Gonzaga from the genus are present. Also, in the third instar, Leucochrysa. the laterodorsal tubercles of seg- Adult.—A similar, but not identical, ments A4 and A5 are relatively flat situation appears to pertain for the adults. and difficult to see except under Several features of the adult (a heavily relatively high magnification (. marked head, thorax, and wing tips, 803); however, under magnification rectangular intramedian cell, lack of a they are noticeably sclerotized and suture between S8 and S9, narrow med- each has the two LDS and a iuncus) that occur in G. nigriceps and that microseta. have been used to distinguish Gonzaga (j) Abdomen: Leucochrysa (both sub- (e.g., Brooks and Barnard 1990), also genera): Segment A1 with single row occur in species of Leucochrysa (e.g., see of 8–16 (L3), 3–6 (L2), or 4 (L1) Brooks and Barnard 1990; Mantoanelli et smooth hooked SMS between the al. 2006; Tauber 2007). Moreover, the two laterodorsal tubercles (Fig. 4). short lobe at the apex of the male Note: In G. nigriceps (L3 and L2), ectoprocts, which occurs in Gonzaga segment A1 has two rows of setae – torquatus Nava´s (the type species of the the anterior row with one pair of genus) and that has been proposed as a medium length, smooth, hooked distinguishing character state for Gonzaga setae (sometimes a single seta) near (Brooks and Barnard 1990), is absent the midline and four short, slender, from G. nigriceps. pointed setae submesally. The pos- Thus, at this time, the only currently terior row of 20–24 smooth, hooked identified character that may separate SMS between the two laterodorsal Gonzaga from Leucochrysa appears in VOLUME 110, NUMBER 2 437 the male – a narrow mediuncus with tamu.edu/research/neuropterida/neuroweb. dorsal striations. This character state is html), maintained by J. D. Oswald was shown here for G. nigriceps, and it was helpful during our study. reported for G. torquatus (Brooks and Barnard 1990). Whether these striations LITERATURE CITED are unique to Gonzaga and whether they Adams, P. A. 1987. Studies in neotropical Chryso- signify phylogenetic relatedness above pidae (Neuroptera) III. Notes on Nodita amazonica Nava´s and N. oenops, n.sp. Neu- the species level are presently unknown. roptera International 4: 287–294. Also, the distribution of this character Adams, P. A. and N. D. Penny. 1986. Faunal state within the eight species that cur- relations of Amazonian Chrysopidae, pp. 119– rently comprise Gonzaga is unreported. 124. In J. Gepp, H. Aspo¨ck, and H. Holzel, Therefore, while the preponderance of eds. Recent research in Neuropterology. Druc- khaus Thalerhof, Graz, Austria. the evidence from the larval and adult Alayo, D. P. 1968. Los Neuro´pteros de Cuba. morphology does not support keeping G. Poeyana Instituto de Biologia (B) 2: 1–127. nigriceps separate from Leucochrysa,we Banks, N. 1914. New neuropteroid , native will refrain from making taxonomic and exotic. Proceedings of the Academy of changes until other species within Gon- Natural Sciences of Philadelphia 66: 608–632. ———. 1944. Neuroptera of northern South zaga and Leucochrysa have been exam- America, Part III. Boletin de Entomologia ined in detail. Venezolana 3: 1–34. ———. 1945. Review of the Chrysopidae (Notho- ACKNOWLEDGMENTS chrysidae) of Central America. Psyche 52: Our work was supported by the 139–174. Brooks, S. J. and P. C. Barnard. 1990. The green National Science Foundation (Grants lacewings of the world: A generic review INT 9817231, INT-9912449, DEB (Neuroptera: Chrysopidae). Bulletin of the 0542373), the National Geographic So- British Museum of Natural History (Entomol- ciety, the ‘‘Conselho Nacional de Desen- ogy) 59: 117–286. ´ volvimento Cientı´fico e Tecnolo´gico’’ Dıaz-Aranda, L. M. and V. J. Monserrat. 1995. Aphidophagous predator diagnosis: Key to (CNPq, Brazil – Grants 303287/2005-5 genera of European chrysopid larvae (Neur.: and 475848/2004-7), Regional Project W- Chrysopidae). Entomophaga 40: 169–181. 1185, Cornell University, and Universi- Mantoanelli, E., G. S. Albuquerque, C. A. Tauber, dade Estadual do Norte Fluminense. We and M. J. Tauber. 2006. Leucochrysa (Leuco- chrysa) varia (Neuroptera: Chrysopidae): Lar- (C.A.T. and M.J.T.) thank Les E. Ehler, val descriptions, developmental rates, and Robert E. Page, Diane E. Ullman, adult color variation. Annals of the Entomo- Walter Leal, Sandra Kelley, Lynn S. logical Society of America 99: 7–18. Kimsey, Michael Parrella, and the De- McLachlan, R. 1868 (1867). New genera and partment of Entomology, University of species, etc. of neuropterous insects; and a revision of Mr. F. Walker’s British Museum California, Davis, for their help and Catalogue of Neuroptera, part ii. (1853), as far cooperation in a variety of ways. We as the end of the genus Myrmeleon. Journal of also thank David Goodger, The Natural the Linnean Society of London, Zoology 9: History Museum, London, and O. S. 230–281. Flint, Smithsonian Institution, Washing- Nava´s, L. 1913a. Les Chrysopides (Ins. Ne´vr.) du Muse´e de Londres. Annales de la Socie´te´ ton, D.C., for facilitating the loan of Scientifique de Bruxelles 37(pt. 2): 292–330. specimens. The following people collect- ———. 1913b. Criso´pidos sudamericanos. Brote´ria ed specimens or assisted with our field (Serie Zoolo´gica) 11: 73–104, 149–168. work: E. A. Silva, Paul J. Tauber, ———. 1929. Insectos del Brasil. Revista do Agatha J. Tauber, Michael J. Tauber, Museu Paulista, Sa˜o Paulo 16: 857–864. ———. 1932. Alcuni insetti del Museo di Zoologia James M. Carpenter, Deborah Smith; it della R. Universita` di Torino. Bollettino dei is a pleasure to acknowledge their help. Musei di Zoologia e di Anatomia comparata The ‘‘Neuroweb’’ website (http://insects. della R. Universita` di Torino 42: 1–38. 438 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

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