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Generic Characteristics of Chrysopodes (Neuroptera: Chrysopidae), with New Larval Descriptions and a Review of Species from the United States and Canada

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Generic Characteristics of Chrysopodes (Neuroptera: Chrysopidae), with New Larval Descriptions and a Review of Species from the United States and Canada SYSTEMATICS Generic Characteristics of Chrysopodes (Neuroptera: Chrysopidae), with New Larval Descriptions and a Review of Species from the United States and Canada CATHERINE A. TAUBER1 Department of Entomology, Comstock Hall, Cornell University, Ithaca, NY 14853Ð0901 Ann. Entomol. Soc. Am. 96(4): 472Ð490 (2003) ABSTRACT Many of the adult features that are considered diagnostic for Chrysopodes occur in other New World chrysopid genera; a stable and phylogenetically signiÞcant generic classiÞcation requires additional deÞnitive characters. The current study demonstrates a number of morphological char- acters in the larvae that provide strong evidence for the validity of the genus and also provide a foundation for broadening and reÞning the deÞnition of the genus. For example, the absence of a gonapsis no longer serves as a diagnostic character for Chrysopodes. Moreover, based primarily on larval characteristics but also on adult features, the widely distributed Nearctic species Ceraeochrysa placita (Banks) is now transferred to Chrysopodes. Finally, the manuscript reviews the systematics of the two Chrysopodes species reported from the United States; the adults are characterized and illustrated, the larvae are described, and biological data are presented. RESUMEN Muchas de las caracterõ´sticas de los adultos que son consideradas para el diagno´stico de Chrysopodes, ocurren en otros ge´neros de criso´pidos del Nuevo Mundo. Una clasiÞcacio´n gene´rica estable y Þlogene´ticamente signiÞcante requiere de caracteres diÞnitivos adicionales. Este estudio muestra un nu´ mero de caracterõ´sticas morfolo´gicas en la larva que provee evidencias so´lidas para la validez del ge´nero y tambie´n proporciona un fundamento para ampliar y reÞnar la deÞnicio´ndee´ste. Por ejemplo, la ausencia de gonapsis ya no sirve como un cara´cter determinante para Chrysopodes. Adema´s, basado primariamente en caracterõ´sticas larvales asõ´ como tambie´n en rasgos del adulto, la especie Nea´rtica distribuida ampliamente, Ceraeochrysa placita (Banks), es ahora transferida a Chry- sopodes. Finalmente, en el manuscrito se reexamina la sistema´tica de las dos especies de Chrysopodes citadas para los Estados Unidos de Ame´rica; los adultos son caracterizados e ilustrados, las larvas son descritas, y los datos biolo´gicos son presentados. KEY WORDS Chrysopodes, Ceraeochrysa, green lacewings, larvae, Chrysopidae THE NEOTROPICAL GREEN LACEWING genus Chrysopodes in misidentiÞcations. First, sclerotized internal struc- Nava´s is a large, widespread, and complex assemblage tures, especially genitalic structures, can be difÞcult to of Ϸ40 described and many undescribed species differentiate on young specimens (Adams 1982, Ad- (Brooks and Barnard 1990). The taxon was erected on ams and Penny 1987), even those that are cleared and the basis of external adult features (Nava´s 1913). Sub- stained. This limitation poses a signiÞcant problem sequently, Adams and Penny (1987) described the because specimens in museums and those that agri- internal and external genitalic characteristics of the culturalists send to systematists for identiÞcation are genus and subdivided the group into two subgenera: often teneral. Second, many of the traits in the generic Chrysopodes Nava´s, with sickle-shaped mandibles, and diagnosis of Chrysopodes are shared by other genera of Neosuarius Adams and Penny, with broadly tipped Chrysopini, especially Ceraeochrysa Adams, which is mandibles. More recently, in their worldwide review another large, widely distributed New World genus. of chrysopid genera, Brooks and Barnard (1990) pro- Thus, the array of genitalic and other features that vided keys and descriptions for identifying the adults characterize Chrysopodes, vis a´ vis other chrysopine of this and the other known genera of chrysopids. genera, needs to be reÞned and expanded. SpeciÞ- For various reasons given below, some of the geni- cally, new characters are needed to stabilize the ge- talic traits that have been proposed to differentiate the neric classiÞcation of the family and to promote a genus Chrysopodes are problematic and have resulted phylogenetic analysis. Recent work on the chrysopid larvae of Europe, 1 E-mail: [email protected]. Japan, and the New World demonstrates the useful- 0013-8746/03/0472Ð0490$04.00/0 ᭧ 2003 Entomological Society of America July 2003 GENETIC CHARACTERISTICS OF Chrysopodes 473 ness of larval morphology in chrysopid systematics, mens (adults and larvae) are being used for on-going especially at generic and higher levels (e.g., Dõ´az- studies and are in the Tauber collection. Aranda and Monserrat 1995; Tsukaguchi 1995; Tauber Procedures and Terminology. The rearing, preser- et al. 2000, 2001; Dõ´az-Aranda et al. 2001). The current vation, and descriptive procedures are presented by study reinforces this point. Although a very distinctive Tauber et al. (1998, 2000). Interpretations of male and set of structural, setal, and color traits distinguishes the female genital structures coincide with those of pre- larvae of most species that are currently included in vious authors (Tjeder 1970, Principi 1977, Adams and Ceraeochrysa, that of one North American species Penny 1987), and the morphological terms are gen- [Ceraeochrysa placita (Banks)] differs signiÞcantly erally consistent with those in current use (e.g., from all others. Consequently, it was proposed that Brooks and Barnard 1990, Tauber et al. 2000, de Freitas this species might fall into another genus (Tauber et al. and Penny 2001, Aspo¨ck 2002). The only exception is 2000). R. C. Smith and P. A. Adams reared Chrysopodes that I retain the older usage of “mediuncus” (rather larvae and reported that they are trash-carriers (Smith than “arcessus”) to refer to the median process of the 1931, Adams and Penny 1987). However, until now upper part of the gonarcus. The median process of the systematically important morphological traits of some species of Ceraeochrysa and Chrysopodes appears Chrysopodes larvae have remained undescribed and, to bear a distinct terminal sclerite, which here is called thus, unavailable for comparative evaluation. During the arcessus. This terminology is consistent with the course of the current study, it became apparent Tjeder (1970), Adams (1985), Adams and Penny that Ce. placita, does indeed, belong within Chrysopo- (1987), and de Freitas and Penny (2001), but differs des and that the scope of the generic traits that deÞne from that of Brooks and Barnard (1990) and Aspo¨ck the genus Chrysopodes needs considerable modiÞca- (2002). tion. Terms for larval structures follow those of Rousset This paper (1) evaluates the adult characteristics (1966) for the head and Tauber et al. (2000) for the currently used to deÞne the genus Chrysopodes, es- thorax and abdomen (summarized in Tauber et al. pecially in relation to Ceraeochrysa and speciÞcally 2000). Unless stated otherwise, all larval setae are Ce. placita; (2) describes the character states that smooth, pointed, and relatively straight; setae desig- distinguish Chrysopodes larvae and compares them nated as “hooked” are smooth, blunt tipped, and with those in Ceraeochrysa; (3) transfers Ce. placita to hooked apically. “Thorny” setae (ϭ“serrated setae” of Chrysopodes (Neosuarius); and (4) reviews the sys- Tauber et al. 1998) are usually relatively straight and tematics of the two Chrysopodes species known from pointed, but sometimes blunt; those that appear oth- the United States [Ch. placita and Chrysopodes collaris erwise are noted in the descriptions. The “thorns” on (Schneider)]. Adults and larvae of both species are thorny setae can be large (visible under Ͻ200ϫ mag- characterized, the larvae of Ch. collaris are described niÞcation) or small (only visible under Ͼ200ϫ mag- for the Þrst time, and biological data are given. niÞcation). The term “semaphoront” refers to a de- Þned period or life-stage in the development of individuals within a species that is systematically meaningful (see Wheeler 1990). In the discussion Materials and Methods here, semaphoront A refers to Þrst instars, which, in Species Examined. To evaluate the adult character- chrysopids, have a distinct set of interspeciÞcally vari- istics of Chrysopodes, I examined specimens of nine able traits, and semaphoront B refers to second and species: Ch. collaris, Ch. copia de Freitas and Penny, third instars, which share a set of interspeciÞcally Ch. divisa (Walker), Ch. n. sp. near Ch. lineafrons variable traits. Adams and Penny, Ch. pulchella (Banks), and four unidentiÞed or new species. These nine species in- clude representatives of both Chrysopodes subgenera Characteristics of Chrysopodes (Chrysopodes and Neosuarius). I also examined adult Adult specimens of all Ceraeochrysa species from the United States (see Tauber et al. 2000) and four addi- Chrysopodes adults are medium-sized, pale green to tional species from Latin America. In addition, I stud- dark green lacewings with a suite of genitalic (male ied the drawings of males and females in all recent and female) characters (Adams 1985, Adams and publications on the two genera (Adams 1985; Adams Penny 1987, Brooks and Barnard 1990, de Freitas and and Penny 1987; Penny 1997, 1998, 2001; de Freitas and Penny 2001). Although these characters are believed Penny 2001). to be consistent for Chrysopodes, the same character Descriptions of generic-level larval characteristics states (e.g., sternites 8 ϩ 9 fused, absence of tignum, are based on specimens from the following species parameres, pseudopenis, spinellae, and microtholi,
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