Systematics of Green Lacewings (: ) : Larvae of from Mexico

CATHERINE A. TAUBER' AND TERESA DE LEON

Department of Entomology, Cornstock Hall, Corriell University, Ithaca, NY 14853-0901

Ann. Entomol. Soc. Am. 94(2): 197-209 (2001) ABSTRACT Currently, 16 species in the green lacewing genus Ceraeochqsa are known from Mexico. Many of these species occur in agricultural situations, and their trash-carrying larvae are attractive candidates for mass-rearing and use in biological control. To facilitate the use of Ceraeo- chqscr in biological control and to contribute to a better understanding of the phylogenetics of the group, emphasis is placed on systematics. Previously, we described the larvae of seven of the 16 species; here we describe the larvae of five more and provide keys for identifying the larvae of all known Mexican species. We also summarize the little information that is available on the biology of each species.

RESUMEN Actualmente, dieciseis especies de cris6pidos del g6nero Cerneochnjsa se conocen de MBxico. Muchas de Cstas se encuentran bajo diferentes condiciones agncolas; sus larvas carga-basura son candidates atractivos para la cria masiva y utilizacidn en programas de control biol6gico. Para facilitar el uso prictico de Cei-neochrysn, y contribuir a un nlejor entendinliento de las relaciones 6logen6ticas del grupo, nosotros enfatizamos el desarrollo de la sistemiitica del g6nero. Previamente describimos siete larvas de las dieciseis especies; en el presente documento describimos cinco m5s y proporcionamos claves para identificar todos 10s instares de las especies Mexicanas conocidas. Tambier1 resumimos la escasa informaci6n que existe acerca de la biologia de cada especie.

KEY WORDS larval n~orphology,trash-carriers, Chrysopidae, systematics, Mexico

SIXTEENOF THE 46 species in the New World lacewing tion on Cermochnjsa species has become available either genus Ceraeochnrsa are known to occur in Mexico (Ad- to help realize their potential in biological control or to ams 1982, Adams and Penny 1987, Penny 1998; unpub- offer a phylogenetic perspective on the group. The few lished data). The larvae of seven of these species, those recent biological and behavioral studies that deal with that also occur in the United States, were treated sys- Cernaochrysa highlight the immediate need for descrip- tematically in earlier articles (Tauber et al. 1998,2000). tive systematics and keys to the larvae of this group In this article we describe the larvae of five additional (Eisner and Silberglied 1988; Mason et al. 1991; Venzon species, including a new one, and we provide a key for and Carvalho 1993; Eisner et al. 1996; Tauber et al. 1998; identifjnrig the known Ceraeochnjsa larvae of Mexico. To Mpez Arroyo et al. 1999% 1999b, 1999~;Albuquerque et aid in identifying adults of Ceraaochnjsa, two regional al. 2001). keys are available (Amazonia: Adams and Penny 1987; the United States: Tauber et al. 2000), and a compre- hensive review of the genus is being prepared (C.A.T. and N. D. Penny, unpublished data). Materials and Methods Cwmochnjsa species are frequently associated with Larvae were reared from field-collected females agricultural crops, including citrus, vegetables, and and preserved as described by Tauber et al. (1998, maize, where larvae prey upon pests. Many 2000). Larval descriptions are provided for each of species are amenable to mass-production and appear well protected from their own natural enemies. Conse- the two systematically meaningful periods: the first quently, they have considerable potential as classical or instar (L1 = Semaphoront A), and the second and augmentative biological control agents, especially when third instars (L2 and L3 = Semaphoront B) as well intra~uildwedation is a consideration (N6iiez 1988b: as for the egg. Terminology follows that used earlier ~6pe; ~rrb~oet a]. 1999% 1999c; ~auberet al. 20001 (Tauber et al. 2000), and for convenience we in- Albuquerque et al. 2001). Partly because of the poor clude Figs. 1, 2 and 3 (reprinted from Tauber et al. status of their systematics, very little biological informa- 2000) that illustrate essential structures and setae. Codes for acronyms used in the descriptions are provided in the figure legends, except for cranial setae 8-10 (S8-SlO), which are ventral setae and ' E-mail: [email protected] illustrated by Rousset (1966).

0013-874610110397-0209$02.0010 0 2001 Entomological Society of America ANNALSOF THE ENTOMOLOGICALSOCIETY OF AMERICA Vol. 94, no. 2

Fig. 1. Primary thoracic structures of Ceraeochrysa Semaphoront B (second and third instars). T1, T2, T3 = thoracic segment, Sp = spiracle, LT = lateral tubercle, LS = seta on lateral tubercle, Sc = sclerite, Sx = seta, SxScx = seta associated with sclerite, R = row of setae (reprinted from Tauber et al. 2000).

Ceraeochrysa Larvae from Mexico Like most trash-carrying chrysopid larvae, Ceraeochrysa larvae have gibbous (humped) bodies, elongate thoracic tubercles and papilliform abdom- inal tubercles that bear long serrated or hooked

Fig. 3. Thorax and abdomen of typical Crmeochrysu semaphoront A (first instar). T = thoracic segments, A = abdominal segments, Sc = sclerite, LT = lateral tubercle, LDT = laterodorsal tubercle, LS = seta on lateral tubercle, LDS = seta on laterodorsal tubercle, Sx = seta, SxScx = seta associated with sclerite, SSp = seta associated with spiracle, SMS = submedian seta, R = row of setae (reprinted from Tauber et al. 2000).

setae, and rows of hooked abdominal setae. In a previous article we provided a detailed description of the morphological and setal characteristics that are shared by all Ceraeochrysa species that occur in the United States (Tauber et al. 2000). The five Mexican species described here also express these shared traits, with one exception. Some primary setae on the head of Ceraeochrysa sanchezi (Navks) first instars are thorny (Sl, S11) or blunt (S2, S5). These character states differentiate the larvae of C. Fig. 2. Primary abdominal stuctures of Ceraeochry.sa sanchezi from the described larvae of other Ceraeo- Semaphoront B (second and third instars). A = abdominal chr.sya as well as other species, all of segment, Sp = spiracle, LT = lateral tubercle, LDT = lat- erodorsal tubercle, LS = seta on lateral tubercle, LDS = seta which have smooth, pointed cephalic setae (Diaz- on laterodorsal tubercle, Sx = seta; SMS = submedian seta, Aranda and Monserrat 1995). The cephalic setae of SSp = seta associated with spiracle (reprinted from Tauber C. sanchezi third instars are typical of chrysopine et al. 2000). A2 to A4 are similar to A5. larvae-smooth and pointed. March 2001 TAUBERAND DE LE~N:Ceraeochrysa OF Mwco 199

Keys to Mexican Ceraeochrysa Larvae Semaphoront 11. Epicranial and frontal marks linear; postfrontal B (Second and Third Instars, L2 and L3) mark very small and beneath antenna (L3) 1. Intermandibular mark present (Fig. 4 B-E) . 2 (Fig. 4J) or absent (L2);prothoracic sclerites 1'. Intermandibular mark absent (Fig. 4A and uncolored (white) to light amber...... F-L) ...... 5 ...... C. sanchezi 2. Frontal marks distinct (Figs. 4 D and E) . . 3 11'. Epicranial and frontal marks narrow, but not 2'. Frontal marks absent or indistinct from post- linear; postfrontal mark distinct, extending frontal marks (Fig. 4 B and C) ...... 4 from midregion of head to external margin of 3. Frontal marks irregularly shaped (Fig. 4D); pro- antenna (Fig. 4L); prothoracic sclerites brown ...... C. ualida thorax with three lareeL, sclerlites. no second- ary sclerites; abdomen without marks ...... C. sp, near cincta (Mexico) 3'. Frontal mark oblong (Fig. 4E); prothorax with three large sclerites and five to six pairs of small secondary sclerites; lateral Semaphoront A (First Instar, L1) Ceraeochrysa tubercles on abdominal segments 2-6 lineaticornis not available marked with brown ...... C. claoeri 1. Intermandibular mark present (Fig. 5 B-D) . 2 4. Postfrontal marks present, confluent medially 1'. Intermandibular mark absent (Fig. 5A and (Fig. 4C); prothoracic sclerites light brown to E-K) ...... 4 dark brown .....C. sp. near cincta (U.S.A.) 2. Cervix with small, light brown mark laterally 4'. Postfrontal marks absent (Fig. 4B) ;prothoracic (Fig. 5 C and D) ...... 3 sclerites uncolored ...... C. caligata 2'. Cervix without lateral marks (Fig. 5B). .... 5. Epicranial mark globular, extending laterally to ...... C. caligata edge of cervical margin (Fig. 4A, F-H, and 3. Head with S-12 absent or greatly reduced; in- K) ...... termandibular mark very thin (Fig. 5C) ... C. sp. near cincta (U.S.A.) 5'. Epicranial mark narrow, not extending later- ...... ally to edge of cervical margin (Fig. 4 I and 3'. Head with S-12 present; intermandibular mark J,L) ...... 10 wide (Fig. 5D) . C. sp. near cincta (Mexico) 6. Lateral tubercles on abdominal segments 2-7 4. Head with frontal marks only (if other marks marked with brown ...... 7 are present, they are very light) (Fig. 5A, E, 6'. Lateral tubercles on abdominal segments 2-7 andG) ...... 5 without marks...... 8 4'. Head with 2-3 pairs of dorsal marks (Fig. 5F 7. Frontal marks broad, joined by light brown area and H-K) ...... 7 medially (Fig. 4K); genal mark extending 5. Head with frontal marks extending posteriorly ventrally to base of maxillae; distal segment of at least to midregion of head ...... 6 labial palp brown; pronotal plates (TI-Scl) 5'. Head with frontal marks not extending poste- brown throughout ...... C. .smithi riorly (Fig. 5A) ...... C. berlandi 7'. Frontal marks narrow, separate (Fig. 4F) ;genal 6. Frontal marks joined medially, forming a Y that mark not extending ventrally; distal segment extends posteriorly beyond midregion of of labial palp cream to light brown; pronotal head (Fig. 5E) ...... C. claoeri plates (TI-Scl) with small clear area interi- 6'. Frontal marks not joined medially, extending orly ...... C. cubana posteriorly to midregion of head, but not 8. Pronotal sclerites amber or brown; venter of beyond (Fig. 5G) ...... C. eoeres head marked with brown ...... 9 7. Postfrontal (submedian) mark present (Fig. 5 F 8'. Pronotal sclerites uncolored; venter of head andH) ...... 8 without marks ...... C. berlandi 7'. Postfrontal mark absent (Fig. 5 ILK) ....9 9. Meso- and metanotal sclerites uncolored; head 8. Epicranial (lateral) mark extending anteriorly as in 4 H C. linenticornis ...... to external margin of mandibles; frontal 9'. Meso- and metanotal sclerites amber or brown; marks not joined posteriorly, extending only head as in 4 G ...... C. eueres to midregion of head (Fig. 5H) C. placita 10. Head with submedian mark elongate, extending ... from antenna posteriorly to cervical margin 8'. Epicranial mark not reaching mandibles; fron- (Fig. 41); epicranial mark extending from ex- tal marks connected posteriorly by subcuta- ternal margin of jaws posteriorly to cervical neous pigment that extends into epicranial margin; mesothoracic spiracles borne on mark (Fig. 5F) ...... C. cuhana prominent tubercles ...... C. placita 9. Frontal mark appearing V-shaped, with subcu- 10'. Head with submedian (=postfrontal) mark taneous pigment extending posteriorly from short, at most extending from midregion of midregion of head, joining near base of head; head to antenna (Fig. 4 J and L); spiracles epicranial mark narrow throughout (Fig. 5 I not on raised tubercles ...... 11 and K) ...... lo 200 ANNALSOF THE ENTOMOLOGICALSOCIETY OF AMERICA VO~.94. no. 2

Fig. 4. Dorsum of head, third instar. (A) Ceraeochrysa berlandi. (B) Ceraeochrysacaligata. (C) Ceraeochr!/saspecies near cincta (U.S.A.). (D) Ceraeochrysa species near cincta (Mexico). (E) Ceraeochrysa claveri. (F) Ceraeochrysa cubana. (G) Ceraeochrysa eoeres. (H) Ceraeochrysa lineaticmis. (I) Ceraeochrysa placita. (J) Ceraeochnj.~asanchczi. (K) Cerncochrysa smithi. (L) Ceraeochrysa valida. Marks: epi = epicranial, fr = frontal, post = postfrontal, int = intermandibular, sub = submedian, lat = lateral; setae: S1-7, Sll-12, Vx.

9'. Frontal marks separate posteriorly, not extend- Ceraeochrysa berlandi (Banks) ing beyond midregion of head; epicranial (Figs. 4A and 5A) mark wider mesally than basally or distally Larvae. In its larval body shape, setation, and head (Fig. 5J) ...... C. srnithi 10. Epicranial mark wider basally than distally (Fig. markings (lack of an intermandibular mark and 5K); prothorax with pair of brown sclerites. presence of narrow, amber frontal and postfrontal ...... C. oalida marks), C. berlandi most closely resembles C. lin- 10'. Epicranial mark linear throughout (Fig. 51); eaticornis. It can be distinguished from C. lineati- prothoracic sclerites uncolored . C. sanchezi cornis by its entirely white body; in C. berlandi all sclerites, including the episterna and epimera, are uncolored; only the dorsal head marks, tarsal claws, Ceraeochrysa arioles (Banks) and empodia are amber. The larvae described below Larvae, Egg, Habitat, Biology. Unknown. were reared from adults collected 10 miles north of Recorded Distribution. Mexico, Central America Caracol, Cayo District, Belize (January); although (Banks 1945,1948, Adams 1982). MEXICO. Chiapas, the species occurs in Mexico, we have not seen Nuevo Le6n, Sinaloa (Banks 1948, Adams 1982). larvae from there. March 2001

Fig. 4. Continued.

Semaphoront B (L2 and L3). Body. White. Integu- setae (distal three pairs long); distal segment amber. ment with spinules sparse, small, rounded to pointed. Antenna white. Venter white. Cervix white, with six Thorny setae with thorns elongate, thin, raised off short setae laterally. surface of seta, arranged linearly along length of seta. Thorax. White, with sclerites uncolored; LTs Head (Fig. 4A). Cream to white; 0.74-0.78 mm wide elongate, slender, slightly enlarged apically, un- (L3), with two to three pairs of dorsal marks. Epicra- marked basally. Prothorax with SI to S5 smooth- nial mark absent to very light amber; when present, pointed, S4 posterior to Scl, four to five pairs of globular, extending laterally to tip of cervical margin. small secondary setae mesa1 to Scl. Pleuron, venter Postfrontal mark amber, extending from anteriome- white, with one pair small setae posteromesally. dian margin of epicranial mark almost to base of an- Episternum white. Mesothorax with anterior sub- tenna; frontal mark amber, extending from midregion segment approximately one-fourth length of poste- of head to inner margin of mandible. Genal mark rior subsegment; spiracles uncolored, unraised. Pos- absent. S1 to S12 present; S1, S11, S12 long. Mandible terior subsegment with all setae smooth-pointed, S1 light amber basally, darker distally, with five small short, two to five pairs of small secondary setae apical teeth; dorsal surface with sparse microsetae. anteromesal to Sc3. Pleuron, venter white. Epister- Maxilla with five small avical setae: ventral surface num, epimeron generally uncolored; episternum mostly smooth. Labium Ghite to light amber; second sometimes light brown. Metathorax with S1 small, segment of palpus with approximately seven pairs of three to six pairs of small secondary setae anterome- ANNALSOF THE ENTOMOLOGICALSOCIETY OF AMERICA Vol. 94, no. 2

Fig. 5. Dorsum of head, first instar. (A) Ceraeochrysa berlandi. (B) Cmeochrysa caliguta. (C)C~m~ochrysa species near cincta (U.S.A.). (D) Cwaeochrrtsa species near cincta (Mexico). (E) Cwaeochrysa claoeri. (F) Cwaeochrysa czrbana. (G) Cerueochrysa eoeres. (H) Ceraeochrysa placita. (I) Ceraeochrysa sanchezi. (J) Ceraeochlrysa smithi. (K) Cerueochrysa oulida. Marks: epi = epicranial, fr = frontal, post = postfrontal, int = intermandibular, sub = submedian, lat = lateral; setae: S1-7, Sll-12, vx.

sal to LTs; R1 with two pairs of long, thorny-pointed extending from white chalazae, one to two pairs of setae extending from uncolored chalazae, two to smooth-hooked Ssp; LTs papilliform apically, en- three pairs of short smooth-pointed setae. Pleuron, larged basally, extending laterally. LTs on A2, A3 venter white. Episternum, epimeron generally un- with apical and subapical setae smooth-hooked, me- colored; sometimes episternum light brown. Legs dium-length. LTs of A4, A5 bearing two to three long setose, white; claws amber, empodia amber distally. thorny-pointed and one to two long smooth hooked Abdomen. White, without marks; setae, chalazae setae apically, shorter smooth-hooked setae subapi- white. Al: LDT bearing two long, smooth-hooked cally. LTs on A6, A7 bearing long thorny-pointed setae, one microseta; transverse band of approxi- setae apically, short smooth-pointed setae subapi- mately four to five pairs of long, thin, smooth- cally. A6 with two pairs of smooth-pointed setae in hooked setae stemming from chalazae between anterior row, five pairs in mesal row, approximately LDTs, one pair of shorter smooth-pointed setae four pairs between LDTs, one to two pairs of thin lateral to LDTs; Ssp relatively long, thin; S1, S2 thin, Ssp. A7: LT cylindrical; two pairs of short thin setae moderately long. A2-A5: each bearing approxi- in anterior row, one pair in mesal row, two pairs mately three dorsal bands of smooth-hooked setae between LDTs; one pair of thin, medium length Ssp. March 2001 TAUBERAND DE LE~N:Ceraeochrysa OF MEXICO 203

Fig. 5. Continued.

Variation.L2 and L3 very similar except L2 has only hooked; one pair of smooth-hooked setae anterior to one pair of Ssp per abdominal segment; head width submedian setae; Ssp small, thin. 0.52-0.55 mm. Egg. Unknown. Semaphoront A (Ll). Body. White. Thorny setae Habitat. Collected in moist tropical forest. with thorns slightly elongate, raised off surface of seta, Biology. Duration of developmental stages (21 + arranged linearly along length of seta. 1°C): egg: 5-7 d, first instar: 6-7 d; second instar: 5-6 Head (Fig. 5A). White, with light amber frontal d; third instar 4-6 d; cocoon (from spinning to emer- mark anteriorly; 0.36-0.37 mm wide. Genal mark ab- gence): 17-19 d. sent. S12 present, medium-length; eyes black. Cervix Recorded Distribution. Mexico, Central America white, with lateral sclerites uncolored. (Banks 1948). MEXICO: Chiapas (Banks 1948). Thorax. White, with sclerites uncolored; all setae (except LS, T3-R1) smooth. T1-S2 small; T2-S1 small; Ceraeochrysa caligata (Banks) T2-S2 long; T2-S1, T2-S2 arranged vertically; epimera, (Figs. 4B and 5B) episterna white. Abdomen. Al: Ssp long, smooth-hooked; LDT with Larvae. C. caligata larvae resemble those of C. sp. two smooth-hooked setae and one microseta. A2: LS near cincta (U.S.A. population) in shape, coloration smooth-hooked; A3-A4: one LS long, thorny-pointed, and setation (absence of cranial seta 12). However, one LS medium-length, smooth-hooked. A5-A7: LS they can be distinguished by the absence of all head thorny-pointed or thorny-hooked. A6: S2 absent; one markings except the intermandibular (all instars) and pair of submedian setae between LDTs, smooth- genal (L1 only) marks. The larvae described below

March 2001 TAUBERAND DE LE~N:Ceraeochrysa OF MEXICO 205 frontal marks in Semaphoront B. Although we have Sc2, Sc3 uncolored to light amber, all setae smooth- not seen specimens from there, it is likely that this pointed, five to eight short setae anterior to Sc2 (S1 not form occurs in Mexico, so we include it here. distinguishable among them). Pleuron, venter cream. Larvae, Egg, Habitat, Biology. Described in Tauber Episternum, epimeron light brown. Metathorax with et al. 2000. Scl, Sc2 uncolored to light brown; S1 short, few or no Recorded Distribution. Florida, prob. Texas secondary setae, R1 with two pairs of long, thorny- (Tauber et al. 2000). pointed setae extending from uncolored chalazae, one to two pairs of smaller smooth-pointed setae lateral to Ceraeochrysa species near cincta (Mexico) long setae. Pleuron, venter cream. Episternum, (Figs. 4D and 5D) epimeron light brown. Legs unmarked, setose, with -50 -80 setae. Larvae. Similar to C. sp. near cincta from the U.S.A., Abdomen. White, unmarked except light brown except cranial seta 12 is present (Semaphoronts A, B), the frontal mark is present (Semaphoront B), and the sclerites on A9. A10: setae. chalazae white. Al: LDT postfrontal mark is paired, not confluent medially small, bearing three long, smooth-pointed setae; trans- (Semaphoront B). Although the head marks of the verse band of approximately eight to nine pairs of long, Mexican sp. near C. cincta resemble those of C. clazjeri smooth-hooked setae on chalazae between LDTs; Ssp (L3), the Mexican sp, near C. cincta does not have long, thin, slightly removed mesally from spiracle; S1, enlarged thoracic sclerites or brown marks on the S2 thin. A2-A5: each bearing approximately three dor- abdomen. The larvae described below were field-col- sal bands of smooth-hooked setae. LT on A2 with lected in Mexico (Ciudad Victoria, Tamaulipas apical, subapical setae long, smooth-hooked. LTs of A3 [March, October, November] ) , and reared from to A5 with one to several thorny-pointed setae api- adults collected in Mexico (Tecomkn, Colima [Sep- cally, numerous long, smooth-hooked setae apically, tember, October] ) . subapically; venter white to cream. A6: anterior trans- Semaphoront B (L2 and L3). Body. Generally white; verse row with three to four vairs of smooth-hooked integument with spiuules dense, pointed, uncolored to setae, mesa1 row with two to three pairs of long light brown. Thorny setae with thorns slightly elon- smooth-hooked setae; approximately four pairs of gate, raised off surface of seta, arranged linearly along smooth-hooked submedian setae between LDTs. A7: length of seta. LT cylindrical, slightly more pronounced than LT on Head (Fig. 4D). White, 0.81-0.86 mm wide; with A6. AS: venter with posterior patch of -20 setae, three pairs of dorsal marks. Epicranial marks, conflu- posterior ones long. A9: light brown mark posteriorly, ent or near confluent at anteromedian tips, tan, large, sometimes laterally. A10: venter with pair of light extending posterolaterally to edge of cervical margin. brown sclerites. Postfrontal mark dark brown. narrow. extendine" from above anteromedian margiri of epicranial mark to- Variation. L2 and L3 similar except L2 with epicra- ward, but not reaching lateral base of antenna. Frontal nial and frontal marks light to absent, postfrontal mark mark short, brown, between postfrontal marks in mid- amber, genal mark brown, head width 0.54-0.60 mm, region of head, not reaching clypeus. Intermandibular T1-Scl amber, other thoracic sclerites uncolored, and mark dark brown, unpaired, contacting inner base of A1 with LDT bearing three long smooth-hooked setae, each jaw, slightly enlarged medially. Genal mark transverse row of three pairs of long, smooth-hooked brown. S1-S12 present. Mandible brown basolaterally, setae. dark distally, with six apical teeth; dorsal surface SemaphorontA (Ll).Very similar to C. cincta. Body. slightly scaley distally, relatively smooth basally, with White to cream dorsallv. Thornv setae with thorns sparse microsetae. Maxilla with three small apical se- slightly elongate, raised off surface of seta, arranged tae; ventral surface slightly scaley distally, relatively linearly along length of seta. smooth basally. Labium cream to light amber; annu- Head (Fig. 5D).White, without dorsal marks; 0.36- lations of middle segment light amber laterally; second 0.42 mm wide. Intermandibular mark amber. Genal segment of palpus bearing ten setae (distal six long). mark brown. Eyes black. S1 to S12 present. Cervix Antenna cream. Venter white, with very light brown white. mark at base of labium. Cervix white dorsally, with pair Thorax. White, with Scl, Sc2 uncolored; all thoracic of large light brown sclerites laterally, five pairs of setae (except LS, T3-R1) smooth. TI42very small; T2-S1 setae ventrolaterally. short, T2-S2 long; epimera, episterna amber. Thorax. LTs elongate, tubular, slightly enlarged api- Abdomen. Al: SSp medium-length, smooth-pointed. cally, unmarked basally. Prothorax with Scl tan to LS smooth-hooked; A3-A7: LS long, thorny-pointed. brown, darker medially than laterally; Sc2 light brown. A2: lighter posteriorly than anteriorly. S1 to S5 smooth- Egg. Undescribed. pointed; S4 posterior to Scl. Pleuron, venter white to Habitat. Collected from Citrus spp. (citrus) and cream, without setation. Episternum dark brown. Me- Psidium guajazju (guava). sothorax with anterior subsegment approximately Biology. Unknown. one-fourth length of posterior subsegment; spiracles Distribution. MEXICO: Colima (Tecomiin-Sep- uncolored to light brown, unraised; Scl light brown tember, October); Tamaulipas (Ciudad Victoria- (sometimes uncolored). Posterior subsegnient with March, October, November) (new records). ANNALSOF THE ENTOMOLOGICALSOCIETY OF AMERICA Vol. 94, no. 2

Cerueochrysa claveri (Navas) almost to edge of cervical margin, distally to level of (Figs. 4E and 5E) eyes. Postfrontal mark dark brown basally, light brown distally, extending from anteromedian margin of epi- Larvae, Egg, Habitat, Biology. Described in Tauber et al. 2000. cranial mark to base of antenna. Frontal mark amber Recorded Distribution and New Records. Mexico, to brown. extending" from midregion" of head to inner Central America, Caribbean, South America, United margin of mandible. Genal mark amber to brown, States (Florida) (Banks 1945, Adams 1982, Adams and extending from cervical margin to eye. S1 to S12 Penny 1987). MEXICO: Colima (TecomBn-Septem- present; S1, Sll, S12 long. Mandible amber basally, ber, October); Oaxaca (Tuxtepec-January); Tabasco darker distally, with five small apical teeth; dorsal (Huimanguillo-June); Tamaulipas (Ciudad Victoria- surface with sparse microsetae. Maxilla with five small October, November); Yucatan (MCrida-January) (all apical setae; ventral surface scaley distally. Labium new records). amber to light brown; second segment of palpus with 11-12 setae (distal three long), distal annulations brown laterally. Antenna light to dark amber. Venter Ceraeochrysa cubana (Hagen) cream, with amber to light brown mark at base of (Figs. 4F and SF) labium. Cervix cream to white dorsally, with large, Larvae, Egg, Habitat, Biology. Described in Tauber amber to brown sclerotized mark laterally, with six to et al. 2000; also see Lbpez-Arroyo et al. 1999a, 1999c, seven short setae. 2000. Thorax. White; LTs elongate, slender, slightly en- Recorded Distribution and New Records. Mexico, larged apically, unmarked basally. Prothorax with Scl Central America, Caribbean, South America, United large, amber to brown; Sc2 uncolored. S1 to S5 States (Banks 1948, Adams 1982, Adams and Penny smooth-pointed; S1, SlScl long; S4 posterolateral to 1987, Penny et al. 1997). MEXICO: Chiapas, Colima, Scl; four pairs of secondary setae mesal to Scl. Pleu- Oaxaca, Sonora, Tabasco, Tamaulipas, Veracruz ron, venter white, without setation. Episternum amber (Banks 1945, 1948, Adams 1982, Penny et al. 1997); to brown. Mesothorax with anterior subsegment ap- Nuevo Le6n (San Nicolas de 10s Garza, nr. Monter- proximately one-third length of posterior subsegment; rey-June) (new record). spiracles white to amber, unraised; Scl amber to brown. Posterior subsegment with Sc2 uncolored; Sc3 light to dark brown, large; all setae smooth-pointed; six Ceraeochrysa efisa (Navis) pairs of secondary setae anterior to Sc3, one pair mesal Larvae, Egg, Habitat, Biology. Unknown. to S2. Pleuron, venter white. Episternum, epimeron Recorded Distribution. Mexico (Navis 1911). amber to brown. Metathorax with Scl uncolored. Sc2 amber to brown; R1 with two pairs of long, thorny- pointed setae extending from cream chalazae, one pair Ceraeochrysa elegans Penny short smooth-vointed setae lateral to thornv setae. one Larvae, Egg, Habitat, Biology. Unknown. pair betweeniateral and mesal thorny seLe ani one Recorded Distribution. Mexico, Cental America pair mesally. Pleuron, venter white. Episternum, (Penny 1998). MEXICO: Tamaulipas (Penny 1998). epimeron amber to brown. Legs cream, setose; coxae brown proximally; claws amber, empodia brown. Abdomen. White, without marks; setae, chalazae Ceraeochrysa everes (Banks) white. Al: LDT bearing three long, smooth-hooked (Figs. 4G and 5G) setae. one microseta: transverse band of anvroxi-L L Larvae. The head markings of C. meres second and mately eight to nine'pirs of long, smooth-hooked third instars resemble those of C. cuhana. However, setae stemming from chalazae between LDTs; one the C. eueres body is white to cream; only the primary pair of small-pointed setae lateral to LDTs; Ssp long, thoracic sclerites are amber or brown, and the abdo- thin; S1, S2 long. A2-AS: each bearing approximately men is unmarked. The C. everes first instar is notable three dorsal bands of smooth-hooked setae extending because of its bulbous abdominal lateral and lat- from white chalazae, one to three pairs of smooth- erodorsal tubercles and its head markings. The larvae hooked Ssp; LTs papilliform apically, enlarged basally, described below were reared from adults collected at extending laterally; spiracles each with two narrow Guanacaste National Park (-5 miles E. Cuajiniquil subdermal wings. LTs on A2, A3 with apical, subapical, turnoff), Guanacaste Province, Costa Rica (Decem- and dorsal setae long.u, smooth-hooked. LTs of A4. A5 ber) and from eggs that were collected at Ciudad with thorny-pointed setae apically, smooth-hooked Victoria, Tamaulipas, Mexico (December). setae subapically and dorsally; venter white. A6: two Semaphoront B (L2 and L3). Body. White. Integu- rows of small setae anteriorly, row of short setae pos- ment with spinules rounded, knobby. Thorny setae teromesal to spiracles, two pairs of small Ssp, approx- with thorns relatively sparse, elongate, raised off sur- imately two pairs of short smooth-hooked submedian face of seta, arranged linearly along length of seta. setae between LDTs. A7: LT cylindrical; three pairs of Head (Fig. 4G). White to light cream; 0.91-0.94 mm short setae anteromesal to spiracles, one pair Ssp, two wide (L3),with three pairs of dorsal marks. Epicranial pairs of short setae between LDTs. A9: with numerous mark amber to brown, globular, extending laterally dorsal setae. March 2001 TAUBEKAND DE LE~N:Ceraeochrrrsa OF MEXICO

Variation. L2 and L3 very similar except L2 has pale Ceraeochrysa placita (Banks) euicranial mark:. -aenal mark light- brown: head width (Figs.. - 41 and 5H) 0.62-0.63 mm; T3-Scl, T3-Sc2 light tan to uncolored; Larvae, Egg, Habitat, Biology. Described in Tauber thorax with fewer secondary setae; T3-R1 without et al. 1998, 2000. short smooth-pointed setae; A1 with LDT bearing Recorded Distribution. Mexico, United States, Can- three setae, with transverse row of five pairs of long, ada (Banks 1948, Adams 1982, Adams and Penny 1987, smooth-hooked setae; A2-A7 with one Ssp per spir- NavBs 1914, Penny et al. 1997). MEXICO: MichoacBn, acle; A6 with three pairs of setae in anterior transverse Distrito Federal (as Chrysopa forreri) (Navris 1914). row, one pair of long smooth-hooked setae and one pair of short setae in mesa1 row, one pair of long smooth-hooked setae between LDTs. Ceraeochrysa sanchezi (Navk) Semaphoront A (Ll).Body. White dorsally. Thorny (Figs. 4J and 51) setae with thorns sparse, elongate, raised off surface of seta, arranged linearly along length of seta. Larvae. C. sanchezi third instars generally resemble Head (Fig. 5G). White, 0.44-0.45 mm wide; frontal those of C. lineaticornis in their setation, light colora- mark light amber, paired, reaching from medial base of tion, absence of intermandibular marks, and elongate, mandible to midregion of head. Postfrontal and epi- slender thoracic tubercles. C. .sanchezi can be distin- cranial marks absent or very light. Genal mark amber. guished by the shape and narrowness of the epicranial Eyes black. S1 to S12 present. Cervix white; lateral marks, small postfrontal marks, and the presence of sclerite uncolored to light amber. four long setae on the laterodorsal tubercles of the first Thorax. White dorsally, light cream laterally; scle- abdominal segment. First instars are unique among the rites uncolored; all thoracic setae (except LS, T3-R1) species that have been studied in that they have mod- smooth. T1-S1, T1-S2Sc1, T2-S2 long; epimera, epis- ified cephalic setae (thorny S1 and S11; blunt S2 and terna white. S5). The descriptions below are based on specimens Abdomen. White; LTs, LDTs, and chalazae at bases (one of each instar) that were reared from adults of SMS large, bulbous. Al: Ssp long, sn~ooth-hooked. collected at Caiias, Guanacaste Province, Costa Rica A2: LS smooth-hooked; A3-A4: one LS smooth- (December); we have not seen larvae from Mexico. hooked, one thorny pointed; A5-A7: LS long, thorny- Semnpbront B (L.3 and L2). Body. Cream. Integu- pointed. A2-A5: pair of LDTs, two pairs of SMS on ment with spinules dense, pointed, cream to light chalazae anteriorly and one pair of SMS posteriorly amber. Thorny setae with thorns slightly elongate, between LDTs; Ssp long, smooth-hooked; spiracles raised off surface of seta, usually arranged linearly and Ssp arising from fleshy chalazae. A6: two pairs of along length of seta. long, smooth-hooked setae mesa1 and anterolnesal to Head (Fig. 45). Cream, with three pairs of dorsal LDTs. A7: SMS between LDTs long. A8: LT bulbous. marks. Epicranial mark brown, elongate, linear Egg. Green with distinct white micropyle; ovoid, throughout, extending from near lateral edge of cer- 0.91-1.0 nlm long, 0.41-0.47 mm wide; stalk 2.9-3.6 vical margin, anteriorly to level of eye. Postfrontal mm long, with sticky droplets; usually laid in circular mark small, linear, light amber, near lateral margin of cluster of 10-20 eggs. antenna1 socket. Frontal mark linear, dark amber, ex- Habitat. Collected in dry tropical forest. tending anteriorly from midregion of head to inner Recorded Distribution and New Records. Mexico, base of mandible, with posterior arms connected by Central America (Banks 1945,1948). MEXICO: Chia- light amber, V-shaped pigmentation. Anterior margin pas (Banks 1948), Tamaulipas (Ciudad Victoria-De- of clypeus marked with amber. Genal mark light am- cember) (new record). ber, linear, extending from cervix to eye. S1 to S12 present. Mandible amber distally, laterally, light am- ber mesally, with apical teeth very small, with row of Ceraeochrysa graduta (Navb) approximately ten small teeth subapically and medi- Larvae, Egg, Habitat, Biology. Unknown. ally; dorsal surface smooth, with sparse microsetae. Recorded Distribution. Mexico, Central America Maxilla with five relatively long apical setae; ventral (Banks 1948, Adams 1982). MEXICO: Chiapas (Banks surface smooth, with sparce microsetae. Labium 1948). cream to light amber; middle segment of palpus with annulations unmarked laterally, with approximately eight setae, distal three relatively long. Antenna light Ceraeochrysa lineaticornis (Fitch) amber. Venter, cervix cream to white. (Fig. 4H) Thorax. LTs elongate, slender, slightly enlarged api- Although this species has not been reported from cally, unmarked basally; all setae, except LS and T3- Mexico, we include it here in the likelil~oodthat it R1, smooth-pointed. Prothorax with Scl, Sc2 uncol- occurs there. ored; S1 to S5, SlScl long; in addition to S1 to S5, Larvae, Egg, Habitat, Biology. Described and sum- several (approximately five) pairs of short setae be- marized in Tauber et al. 2000. tween paired Scl. Venter unmarked. Episternum light Recorded Distribution. United States, Canada, pos- amber. Mesothorax with anterior subsegment approx- sibly Mexico (Banks 1948, Adams 1982, Adarns and imately one-fourth length of posterior subsegment; Penny 1987, Penny et al. 1997). spiracles uncolored, unraised; all sclerites uncolored. 208 ANNALSOF THE ENTOMOLOGICALSOCIETY OF AMEIIICA Vol. 94, no. 2

Posterior subsegment cream throughout, sclerites un- (Navh) colored. S1, S2, and setae associated with sclerites (Figs. 4K and 55) (Scl, Sc2, Sc3) relatively long; in addition to S1 and S2, Larvae, Egg, Habitat, Biology. Described in Tauber posterior subsegment with eight to nine pairs of short et al. 2000. setae behind Sc2. Pleuron, venter cream. Episternum, Recorded Distribution and New Records. Mexico, epimeron light amber. Metathorax with sclerites (Scl, United States, Central America, Caribbean (Adams Sc2) uncolored; S1 relatively long, no other setae 1982, Penny et al. 1997). MEXICO: Jalisco (Navb anterior to large sclerite; R1 with two pairs long 1929, Penny et al. 1997); Colima (TecomBn-Septem- thorny-pointed setae extending from white chalazae; ber) (new record). one pair very small setae between two thorny setae. Pleuron, venter cream. Episternum, epimeron light amber. Legs cream, setose; coxae amber proximally; Ceraeochrysa valida (Banks) claw amber; empodiurn dark distally. (Figs. 4L and 5K) Abdomen. White, without marks. Al: Ssp damaged Larvae, Egg, Habitat, Biology. Described in Tauber on our specimen; S1, S2 medium-length, mesa1 to spi- et d.2000. racles; LDTs each bearing three long, smooth-hooked Recorded Distribution and New Records. Mexico, setae apically, one long smooth hooked seta subapi- United States, Central America, Caribbean, South cally on distal side; LDTs separated by two rows of America (Banks 1895, NavBs 1914, Adams 1982, Penny setae, each row containing eight pairs of long smooth- et al. 1997). MEXICO: Baja California Sur (Banks hooked setae stemming from large chalazae. A2-A5: 1945); Tamaulipas (Tampico-December) , Yucatjn each segment bearing three dorsal rows of smooth- (ChichCn Itzli- January) (new records). hooked setae, one to three smooth-pointed setae near each spiracle; LTs enlarged basally; those on A2, A3 bearing smooth-hooked setae apically, dorsally; those Acknowledgments on A4, A5 bearing thorny-pointed, smooth-pointed We thank Maurice J. Tauber, Paul J. Tauber, Michael J. setae apically, smooth-hooked setae dorsally. A6: Tauber, Agatha J. Tauber, Jose I. L6pez-Arroyo, Robert G. three transverse rows each with =three to six pairs of Helgesen, Diane M. Helgesen, Elisaveta Chouvakhina, Hugo smooth-hooked setae, posterior row with three pairs C. Arredondo, Victor W. Gonzdez, Mario Rodriguez, and of setae between LDTs; LDT with long seta smooth- Sim6n Hernkndez for help in obtaining specimens; Phillip A. hooked, without thorny setae. A7: S1, S2, Ssp short, Adams (deceased) and N. D. Penny for help and discussions; and Frances Fawcett for preparing the figures. The research stun~py;LDT with long seta thorny-pointed; two pairs was supported in part by USDAINRI Competitive Grant of setae between LDTs. A8: S1, S2 short, five pairs of (9802447), National Science Foundation Grant INT-9817231, short setae posteriorly. A9: sclerites uncolored, setose. Regional Project W-185, and the Grace Griswold Fund, Cor- A10: sclerites uncolored, patch of short, medium- nell University (M.J.T., C.A.T.).We also appreciate the co- length setae ventrally. operation of Rincon-Vitova Insectaries, California, and Ben- Variation.L2 coloration and gross structures similar eficial Insectary, California. NBCI (USDA, APHIS) provided to L3; head width 0.55 mm. The specimen was not partial support for publication costs. cleared so the setation was not studied. Semaplzoront A (Ll) Body. White dorsally. Thorny References Cited setae with thorns short, arranged linearly along length of seta. Adams, P. A. 1982. Crraeod~rysa,a new genus of Chrysopi- Head (Fig. 51). White, 0.39 mm wide; with paired nae (Neuroptera) . (Studies in New World Chrysopidae, part 11). Neuroptera Int. 2: 69-75. light amber epicranial and frontal cuticular marks, Adams, P. A., and N. D. Penny. 1987. Neuroptera of the Y-shaped, subcutaneous pigmented mark extending Amazon Basin, Part lla. Introduction and Chrysopini. posteriorly from frontal mark. Genal mark amber. Acta Amazonica. 15: 413-479. Eyes black. S1-S12 present; S1, S11 long, thorny; S2, S5 Albuquerque, 6. S., C. A. Tauber, and M. J. Tauber. 2001. blunt. Cervix white. Chrysoperlo rxterna and Ceraeochrysa spp.: potential use Thorax. White, with sclerites uncolored; T1- S1, in biological control in the New World tropics and sub- T2-S2 (in addition to LS, T3-R1) thorny; T1-S2 very tropics. In P. McEwen ancl T. New [eds.],Lacewings in the crop environment. Cambridge University Press, Cam- short, T1-S4 short; T2-S1 short; epimera, episterna bridge (in press). white. Banks, N. 1895. Some Mexican Neuroptera. Proc. Calif. Abdomen. White. Al: Ssp long, thorny-pointed. A2, Acad. Sci. 5: 515-522. A3: LT with two smooth-hooked LS; A4: LT with one Banks, N. 1945. A review of the Chrysopidae (Nothochry- smooth-hooked, one thorny-pointed seta; A5-A7: LT sidae) of Central America. Psyche 52: 139-174. with two long thorny-pointed LS. Banks, N. 1948. Chrysopidae (Nothochrysidae) collected in Egg. Unknown. Mexico by Dr. A. Dampf (Neuroptera). Psyche 55: 151- 177. Recorded Distribution and New Records. Mexico, Diaz-hranda, L. %I.,and V. J. hlonserrat. 1995. Aphidopha- Central America, Caribbean, South America (Adams gous predator diagnosis: key to genera of European chry- and Penny 1987).MEXICO: Tamaulipas (Ciudad Vic- sopid larvae (Neur.: Chrysopidae). Entomophaga 40: toria-October) (new record). 169-181. March 2001 TAUBERAND DE h6~:Ceraeochntsa OF MEXICO 209

Eisner,T., andR. E. Silberglied. 1988. A chrysopidlawa that Nuiiez, E. Z. 1988a. Chrysopidae (Neuroptera) del Per6 y cloaks itself in mealybug wax. Psyche 95: 15-19. sus especies mis comunes. Rev. Per. Entomol. 31: 69-75. Eisner, T., A. B. Attygalle, W. E. Conner, M. Eisner, and E. Nuiiez, E. Z. 1988b. Ciclo biol6gico y crianzade Clirysop~rla MacLeod. 1996. Chemical egg defense in a green lace- extrrna y Ct-raeochnj.~acincta (Neuroptera, Chrysopi- wing (C:ercirochrysu smitlli). Proc. Natl. Acad. Sci. U.S.A. dae). Rev. Per. Entomol. 31: 76-82. 93: 3280 -3283. Penny, N. D. 1998. New from Costa Rica L6pez-Arroyo, J. I., C. A. Tauber, and M. J. Tauber. 1999a. (Neuroptera: Chrysopidae). J. Neuropterol. 1: 55-78. Comparative life-histories of the predators Cernc,ochrtjsa Penny, N. D., P. A. Adams, and L. A. Stange. 1997. Species cincta, C. cubancr, and C. smitiii (Neuroptera: Chrysopi- catalog of the Neuroptera, Megaloptera, and Raphid- dae). Ann. Entomol. Soc. Am. 92: 208-217. ioptera of America north of Mexico. Proc. Calif. Acad. Sci. Lbpez-Arroyo, J. I., C. A. Tauber, and M. J. Tauber. 1999b. 50: 39-114. Intermittent oviposition and re-mating in Crrc~eochrtlsci Rousset, A. 1966. Morphologie ckphalique des larves de Pla- cinda (Neuroptera: Chrysopidae). Ann. Entomol. Soc. nipennes (Insectes NkvroptCroi'des). Mem. Mus. Nat. Am. 92: 587-593. Hist. Ser. A Zool. 42: 1-199. Lbpez-Arroyo, J. I., C. A. Tauber, and M. J. Tauber. 199Yc. Tauber, C. A., T. de Le611, J. I. Lbpez-Arroyo, and M. J. Effects of prey on survival, development, and reproduc- Tauber. 1998. Crru~ochr~saplacita (Neuroptera: Chry- tion of trash-carrying chrysopids (Neuroptera: Crrc~ro- sopidae): generic characteristics of larvae, larval de- chrYsa).Environ. Entomol. 28: 1183-1 188. scription, and life cycle. Ann. Entomol. Soc. Am. 91: Lbpez-Arroyo, J. I., C. A. Tauber, and M. J. Tauber. 2000. 608-618. Survival of stored lacewing eggs and quality of subse- Tauber, C. A,, T. de Lebn, J. I. L6pez-Arroyo, and M. J. quent stages (Neuroptera: Crra~oclir~~sa).Biol. Control Tauber. 2000. The genus C~~rcieochr~scr(Neuroptera: 18: 165-171. Chrysopidae) of America North of Mexico: larvae, adults Mason, R. T., H. M. Falles, M. Eisner, and T. Eisner. 1991. and corrlparative biology. Ann. Entomol. Soc. Am. 93: Wax of a whitefly and its utilization by a chrysopid larva. 1195-1221. Naturwissenschaften 78: 28-30. Venzon, M., and C. F. Carvalho. 1993. Desenvolvinlento Navis, L. 1911. Chrysopides nouveaux. Ann. Soc. Sci. Brux- larval, pr6-pupal e pupal de Crraeoc.hry,sa cubann (Hagen) elles 35: 266-282. (Neuroptera: Chrysopidae) em diferentes dietas e tem- Navis, L. 1914. Les Chrysopides (Ins.,'Nevr.) du Muske de peraturas. Ann. Soc. Entomol. Braz. 22: 477-483. Londres. [lB].Ann. Soc. Sci. Bruxelles 38: 73-114. NavL, L. 1929. Insectos Neur6pteros del Museo de Ham- burgo. Mem. Real Soc. Espan. Hist. Nat. 15: 315-322. Bibliography of the Neuropterida

Bibliography of the Neuropterida Reference number (r#): 9793

Reference Citation: Tauber, C. A.; de León, T. 2001 [2001.??.??]. Systematics of green lacewings (Neuroptera: Chrysopidae): larvae of Ceraeochrysa from Mexico. Annals of the Entomological Society of America 94:197-209.

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Notes:

File: File produced for the Bibliography of the Neuropterida (BotN) component of the Global Lacewing Digital Library (GLDL) Project, 2006.