Neuroptera: Chrysopidae) : Larvae of Ceraeochrysa from Mexico
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Systematics of Green Lacewings (Neuroptera: Chrysopidae) : Larvae of Ceraeochrysa 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 arthropod 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 Chrysopini 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