SYSTEMATICS Generic Characteristics of Chrysopodes (: ), 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 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 , 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, (Tauber lot numbers): Ch. collaris (lots 63:086, 96:072, presence or absence of gonocristae) are also consis- 99:051), Ch. divisa (lots 96:017, 96:018, 96:019, 99:020, tently expressed by Cereaeochrysa (Adams 1982). 99:043), Ch. n. sp. near Ch. lineafrons Adams and Penny Thus, they are of little value in differentiating the two (lot 96:006), Ch. pulchella (lots 2001:003), and four genera and do not receive further discussion here. unidentiÞed or new species (lots 99:037, 2001:007, Other character states that are considered to be 2002:21, 2002:26). All of the larvae were reared from distinctive for Chrysopodes either vary within the ge- eggs laid by Þeld-collected females. Voucher speci- nus, or overlap with Ceraeochrysa; the traits below are 474 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA Vol. 96, no. 4

Barnard 1990, Fig. 336) it is elongate and up-curved. Thus, the occurrence of an elongate gonapsis in Ce. placita (Figs. 1and 2C) does not exclude this species from Chrysopodes. Furthermore, it should be noted that the gonapsis is believed to have multiple, independent origins within the Chrysopini (Principi 1977, Adams 1982). Its occurrence in Chrysopodes could represent another independent derivation or the retention of an ancestral condition. 2. Gonarcus complex, exclusive of mediuncus (see below) (as described by Brooks and Barnard 1990): Chrysopodes—gonarcus usually short, broadly ex- panded laterally, sometimes with narrow lateral horn Fig. 1. Chrysopodes placita male terminalia; gonarcus (probably ϭ gonocornu); entoprocessus absent; complex expanded. Abbreviations: arc ϭ arcessus; dapo ϭ gonosaccus short, gonosetae absent. Ceraeochrysa— dorsal apodeme of 9Tϩect; gp ϭ gonapsis; gs ϭ gonarcus; gonarcus long, narrow; often with well developed gsac ϭ gonosaccus; vapo ϭ ventral apodeme of 9TϩectÕ gonocornua; entoprocessus absent, but lateral horns ϩ ϭ 8S 9S eighth and ninth sternites (fused), 8T eighth ter- often well developed; gonossaccus short, gonosetae gite; 9Tϩect ϭ ninth tergite and ectoproct (fused). absent, few or numerous. Despite the description above, the gonarcus of particularly pertinent to Ce. placita, whose generic Chrysopodes, like that of Cereaochrysa, varies in shape placement has been questioned (Tauber et al. 2000). from very broadly to very narrowly arched [compare 1. Gonapsis (as described by Brooks and Barnard Ch. pulchella (Adams and Penny 1987) with Chryso- 1990): Chrysopodes Ð absent. Ceraeochrysa Ð elongate, podes polygonica Adams and Penny (de Freitas and narrow, bifurcate apically. Penny 2001)]. Moreover, the arms (apodemes) may All Ceraeochrysa species whose male terminalia vary from broad and expanded, to small or almost have been described have an elongate gonapsis absent [compare Ch. pulchella with Chrysopodes (Adams and Penny 1987, Penny 1997, de Freitas and delicata de Freitas and Penny (2001)]. Thus, the Penny 2001). Although both Brooks and Barnard broad transverse bridge and expanded apodemes of (1990) and de Freitas and Penny (2001) list the ab- Ce. placitaÕs gonarcus could Þt either genus. sence of a gonapsis as a distinguishing characteristic of As stated above, the Chrysopodes gonarcus is typi- Chrysopodes, each set of authors cites an example of cally less elaborate than that of Ceraeochrysa. That is, Chrysopodes in which the gonapsis is present. In Chry- it lacks a dorsal hood and usually lacks entoprocessus sopodes mediocris Adams and Penny, the gonapsis is (unarticulated, paired ventral projections) and gono- short and Y-shaped (bifurcate apically) (Adams and cornua (unarticulated, paired lateral projections) [for Penny 1987), but in Chrysopodes ?sp. (Brooks and a notable exception, see Chrysopodes victoriae Penny

Fig. 2. Chrysopodes placita male genital structures. (A) Gonarcus complex (expanded), dorsal view. (B) Gonarcus complex (expanded), lateral view. (C) Gonapsis, lateral view, modiÞed from Tauber et al. (2000). arc ϭ arcessus; gsac ϭ gonosaccus; la ϭ lateral arm of gonarcus. July 2003 GENETIC CHARACTERISTICS OF Chrysopodes 475

(1998)]. In Ceraeochrysa, dorsal hoods and entopro- gonarcal area in Ce. placita lacks the Þeld of microsetae cessus are variable (absent, to large or elongate), and that is associated with the arcessus in many Chrysopo- gonocornua range from very small and simple to elon- des spp. gate and elaborate (Adams 1982, Adams and Penny 4. Apodeme of tergite 9: Brooks and Barnard (1990) 1987, de Freitas and Penny 2001). state that in Chrysopodes the apodeme in tergite nine Like all Chrysopodes spp. and some species of Cere- extends apicoventrally and protrudes beyond the apex aeochrysa, Ce. placita lacks a dorsal hood and ento- of the segment. processus (Fig. 1). However, the presence or absence This character state occurs in many, but not all of gonocornua in Ce. placita is open to interpretation. Chrysopodes spp., and also in several species of Ceraeo- In an earlier study, the paired structures arising me- chrysa (Adams and Penny 1987). Thus, its absence dially from the bridge of the gonarcus (Fig. 2A and B, (Fig. 1) does not exclude Ce. placita from either genus. arc) were believed to be unarticulated; thus they were It is noteworthy that in Ce. placita the heavily scle- identiÞed as gonocornua (Tauber et al. 2000). Study of rotized apodeme along the dorsolateral margin of additional specimens indicates that these structures sternite 8 ϩ 9 extends anteriorly into segment seven are articulated with the gonarcus, and that they more and posteriorly almost to the tip of sternite 8 ϩ 9 (Fig. closely resemble the paired sclerotized rods that are 1). This situation appears unusual for both genera. often associated with the arcessus (see below). In this The female genital systems of Chrysopodes and case, the Ce. placita gonarcus would lack gonocornua Ceraeochrysa are similar in several ways (e.g., prae- as is typical for most Chrysopodes. genitale absent, subgenitale bilobed apically), and Both Ceraeochrysa and Chrysopodes are considered they generally differ in others (Brooks and Barnard to have a “short” gonosaccus. This character-state ap- 1990). Analysis of all available descriptions and illus- pears to be fairly consistent for Chrysopodes, but vari- trations, as well as examination of female specimens able for Ceraeochrysa. Ce. placitaÕs large gonosaccus (see above), indicates that the differences hold for with its trough-like structure on the surface, would be most species, that signiÞcant variation occurs, and somewhat unusual for Chrysopodes. However, a large that this variation is consistent with the transfer of eversible sac-like structure does occur in Chrysopodes Ce. placita to Chrysopodes as follows. nigripilosa (Banks) (C. A.Tauber, unpublished data). 5. Bursa and associated structures (as described by 3. Mediuncus [mesal process attached to gonarcus, Adams and Penny 1987): ChrysopodesÑspermatheca ϭ “median plate” and “arcessus” of Brooks and Bar- connected apically to the bursa by a slender, more nard (1990)], as described by Brooks and Barnard or less elongate bursal duct; bursal glands saccate, (1990): ChrysopodesÑmedian plate (ϭbasal section dorsolateral when present. CeraeochrysaÑsperma- of mediuncus here) absent; arcessus weakly sclero- theca opening into the bursa by a dorsal longitudinal tized, triangular, often domed, with apical hook, pair slit; elaborately developed bursal ducts absent. of mediolateral sclerotized rods, and dorsal micro- setae. CeraeochrysaÑmedian plate large, with median The consistency and distinctiveness of these char- horns; arcessus long or short, narrow tapering apically, acteristics are difÞcult to interpret. The structures are often with median hook and lateral lobes. obscure in most specimens and also in published de- The mediuncus of the known Chrysopodes species scriptions and illustrations; thus the basis for compar- generally follows Brooks and BarnardÕs (1990) de- ison is very weak. In Ce. placita, the spermatheca scription above. In contrast, the mediuncus of Ceraeo- connects to the bursa via a short, twisted duct; the chrysa is variable and often differs from their descrip- bursa is large, saccate, and very delicate (Fig. 3). tion. It can be triangular, domed, and with an apical 6. Spermatheca (as described by Brooks and Bar- hook as in Chrysopodes, or it can be elongate, forked nard 1990): ChrysopodesÑtall, cylindrical, with very or unforked, and with or without lobes or apical barbs. deep ventral invagination; CeraeochrysaÑvery small, In an earlier paper (Tauber et al. 2000), the paired narrow, with a deep ventral invagination. medial structures attached to the bridge of the gonar- In many species of Chrysopodes, the spermatheca, cus were identiÞed as gonocornua, and the arcessus including the velum is an elongate, convoluted or was believed to consist of a very weakly sclerotized, coiled tube-like structure with a trumpet-shaped elongate, decurved structure resting on the surface of opening [e.g., Chrysopodes duckei Adams and Penny, the gonosaccus. A reinterpretation indicates that the Ch. pulchella (Adams and Penny 1987)]. In other spe- paired structures are probably internal rods associated cies, the structure is U-shaped (Ch. lineafrons), or with the arcessus, not gonocornua; they are articu- rounded (Ch. polygonica)asinCeraeochrysa (Adams lated with the gonarcus and they are enclosed within and Penny 1987). The ventral invagination on the a membranous covering. Thus, the Ce. placita medi- spermatheca can be very deep (as in Ch. duckei and uncus appears to lack a median section (ϭmedian Chrysopodes nebulosa Adams and Penny) to shallow plate) (as in Chrysopodes), and the arcessus has the (as in Ch. lineafrons) (Adams and Penny 1987). By pair of mediolateral sclerotized rods that are typical of comparison, the spermatheca of Ce. placita, is rounded Chrysopodes (Fig. 2). The weakly sclerotized, elon- and has a relatively shallow ventral invagination (Fig. gate, decurved thickening of the surface of the 4A). It is, therefore, more typical of Ceraeochrysa than gonosaccus, which originally was interpreted as the Chrysopodes, but it would not be unusual for Chry- arcessus, appears to be part of a trough-like enlarge- sopodes. For example, in many respects it resembles ment of the gonosaccus. It should be noted that the the “inßated, bean-shaped” spermatheca of Ch. poly- 476 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA Vol. 96, no. 4

Fig. 3. Chrysopodes placita female terminalia. b.d.ϭ bursal duct; coll. g. ϭ colleterial gland; cop.b. ϭ copulatory bursa; g.l. ϭ gonopophysis lateralis; ovd ϭ oviduct; sg ϭ subgenitale; sp ϭ spermatheca; sp.d. ϭ spermathecal duct; 7S ϭ seventh sternite; 8T ϭ eighth tergite. gonica (as described and Þgured by Adams and Penny single, simple, terminal seta; and thorax without rows 1987, de Freitas and Penny 2001). of hooked setae (Dõ´az-Aranda and Monserrat 1995). 7. Spermathecal duct (as described by Brooks and The one character in which most Chrysopodes lar- Barnard 1990): ChrysopodesÑlong and sinuous. vae differ from those of other previously described CeraeochrysaÑlong and highly coiled. chrysopine larvae is the presence of thorny cephalic According to the species-speciÞc descriptions of setae. The larvae of all non-Chrysopodes chrysopine Adams and Penny (1987) and de Freitas and Penny species that have been studied [with the exception of (2001), the twisitng of the spermathecal duct ranges Ceraeochrysa sanchezi (Nava´s)] have smooth (not from sinuous, to convoluted, to coiled in both genera. thorny or serrated) cephalic setae (Tauber and de However, the degree of twisting generally appears to Leo´n 2001). In contrast, most of the Chrysopodes lar- be less in Chrysopodes than in Ceraeochrysa. Thus, the vae studied here have some thorny cephalic setae, transfer of Ce. placita (with its relatively short, sinous usually S1, S4, and/or S11. spermathecal duct; Fig. 4A) from Ceraeochrysa to Semaphoront B (Second and Third Instars, L2 and Chrysopodes is clearly consistent with the known vari- L3). Body. Dorsal and ventral integument white to ation within Chrysopodes (c.f. Chrysopodes breviata cream with areas of dense, tan to brown spinules; Adams and Penny, Ch. polygonica) (as shown in Ad- spinules irregularly shaped, tips generally pointed, ams and Penny 1987). without distinct microsetae. Only primary sclerites, no secondary sclerites. Spiracles cylindrical, with dis- tinct, amber to brown sides. Setal bases mainly brown. Larvae Setae of several types: smooth and hooked, smooth Chrysopodes larvae are typical trash-carrying larvae: and pointed, thorny and pointed, or thorny and blunt. they have gibbous, setose bodies, elongate thoracic Thorny setae with thorns dense, raised off surface of tubercles, large abdominal tubercles and long serrated seta, arranged irregularly along length of seta. [In the setae. The eight species that were examined in this following descriptions, unless stated otherwise, all se- study express all but one of the morphological features tae are smooth and pointed]. that typify larvae of the green lacewing tribe Chry- Head (Fig. 5). Cream to white; epicranial mark sopini. That is, they all have mouthparts as long as, or either divided into two separated longitudinal bands longer than the cephalic capsule; ßagellum with a [submesal epicranial mark (epi-m), lateral epicranial

Fig. 4. Chrysopodes placita female genital structures (redrawn from Tauber et al. 2000). (A) Spermatheca, lateral view. (B) Subgenitale, lateral view. (C) Subgenitale, ventral view. sp.d. ϭ spermathecal duct; v ϭ velum; v.i. ϭ ventral invagination. July 2003 GENETIC CHARACTERISTICS OF Chrysopodes 477

Fig. 5. Dorsum of third instar head. (A) Chrysopodes placita (redrawn from Tauber et al. 1998). (B) Chrysopodes collaris. epi-l ϭ lateral epicranial mark; epi-m ϭ mesal epicranial mark; fr ϭ frontal mark; post ϭ postfrontal mark; S1ÐS12 ϭ primary setae; Vx ϭ three small posterior setae. mark (epi-l), as in Fig. 5A and B], or a single mark with illa with two to several small apical setae, one basal seta two conßuent sections: a lighter, submesal one ventrolaterally; ventral surface smooth or with sparse (epi-m) and a darker, lateral one (epi-l) (in some striations. Labium pale to light brown; palpiger with species, the distinction between epi-m and epi-l can be two to three setae ventrally; palpus with basal segment subtle). Lateral epicranial mark (epi-l) extending an- bearing three to four distal setae (one long, ventral; teriorly from posterolateral cervical margin to area two to three short to medium-length; one less in L2); between antennal base and eye. Submesal epicranial second segment annulated, usually marked distolat- mark (epi-m) extending from posterior cervical mar- erally with amber or light brown, with approximately gin to midregion of head, conßuent with postfrontal eight long setae in L3 (three to six setae in L2); distal mark (post) that extends toward antennal base. Fron- segment without setae, slightly enlarged medially. An- tal marks separated or conßuent mesally, extending tenna white to light tan or light brown; pedicel with anteriorly from midregion of head. Intermandibular one spiney microseta near apex; scape with one long mark absent; clypeolabral region unmarked or with lateral seta apically, one microseta in L3 (absent in one or two brown triangular marks. Genal mark light L2). Cervix pale or marked with subcutaneous pig- brown, extending from cervical margin to, or almost to ment and/or spinules; pleuron with or without light eye, bifurcated mesally; prementum with or without brown marks. marks. Eye black. Primary cranial setae S1to S12,Vx Thorax (Figs. 6 and 7). Anterior segments dorso- present; S11, S1 (usually) long; S1, S4, and/or S11 ventrally ßattened; metathorax (T3) with posterior usually thorny. Labrum truncate, with three pairs dis- subsegment raised above anterior subsegment, bear- tal setae (mesal pair long; two lateral pairs shorter). ing prominent row of long, thick, pointed, thorny setae Mandible longer or shorter than head width; amber, arising from large setal bases (R1). Lateral tubercles with three to four very small apical teeth, base with (LT) elongate, robust, enlarged apically, bearing long, dorsal row of four oval structures (placoid organs of pointed, thorny setae (LS) apically, and fewer, some- Rousset 1966), one lateral basal seta (L3 only); dorsal times shorter, pointed, thorny LS subapically on dorsal surface smooth or with sparse vertical striations. Max- and lateral surfaces. Prothoracic LTs tubular through- 478 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA Vol. 96, no. 4

Fig. 6. Primary thoracic structures of Chrysopodes semaphoront B (second and third instars), dorsal view. T1, T2, T3 ϭ thoracic segment; Sp ϭ spiracle; LT ϭ lateral tubercle; LS ϭ seta on lateral tubercle; Sc ϭ sclerite; Sx ϭ seta; SxScx ϭ seta associated with sclerite; R1 ϭ row of setae. out; mesothoracic and metathoracic LTs enlarged ba- medium-sized, oval sclerites (Sc1) on anterior margin sally. Legs mostly white or cream-colored, setose; (sometimes distinguishable only on cleared speci- coxal bases marked with brown; femora each with mens), each with one to three small setae (S1Sc1 to light brown mesal band or unmarked; tarsi white to S3Sc1)(in L2, Sc1cream colored, S3Sc1absent). Pos- light brown; claws amber to dark brown; empodia and terior subsegment with two pairs of sclerites: Sc2 very episterna dark distally; empodial-coxal apodemes very small, oval, bearing two very small setae (S1Sc2 and dark. S2Sc2); Sc3 posteromesal to LTs, large, round, with Prothorax (T1). Paired notal sclerite (Sc1) rhom- one small seta (S1Sc3) attached (S2Sc3 absent). S1, S2 boidal, brown to light brown, usually with interior very small, anteromesal to Sc3, posterior row consist- white to cream, bearing relatively long thorny seta ing of one pair of small smooth setae (S4), two pairs (S1Sc1) medially, with additional small, smooth seta of long, pointed, thorny setae, arising from pale cha- (S2Sc1); median sclerite (Sc2) usually small (very lazae with dark setal bases (S3, S5). Venter with one small or absent in L2); primary setae S1to S4 present, to two pairs of setae mesally. Episternum large, trap- S5 absent; S1, S2, S3 thorny, arising from brown setal ezoidal, light to dark brown, distally with one prom- bases, S4 short, smooth; R1small or absent; venter with inent seta, extending from brown setal base; epimeron approximately two pairs of setae mesally. Episternum small, triangular or irregularly shaped, light brown to large, triangular, light brown to dark brown, with two dark brown, with prominent seta distally. pairs of short setae distally, one pair proximally; Metathorax (T3). Segment distinctly folded, with epimeron unmarked or dark brown. posterior fold conspicuously raised above dorsum of Mesothorax (T2). Anterior subsegment with spi- T2 and anterior of T3. Primary sclerite (Sc1) and racles prominent, borne apically on large ßeshy associated seta (S1Sc1) very small; Sc2 usually absent, (sometimes retracted) protuberances; pair of small to or small and with associated short setae very small; S1, July 2003 GENETIC CHARACTERISTICS OF Chrysopodes 479

A2ÐA5: Subequal in size and shape; LTs well developed, thick basally, rounded distally, extending ventrolaterally, with brown marks anteriorly and posteriorly (marks sometimes light on all segments, sometimes missing from A2), with long, pointed, thorny LS apically, patch of hooked, smooth LS sub- apically and dorsally. LDTs absent. S1, S2 absent; patch of hooked, smooth setae mesal to spiracles (SSp); SMS long, hooked, smooth, generally curving posteriorly, extending from brown setal bases in transverse bands; total number of SMS/segment ϭ 140Ð230 (L3), 44Ð100 (L2). Venter of A3-A5 with pair of large chalazae laterally, each bearing long seta stemming from brown to light brown setal base. A6: LTs prominent, hemispherical, unmarked or marked with brown dorsally and ventrally, with nine to ten (L3) or eight (L2), thorny LS; LS restricted to LT, not extending mesally onto body of segment. LDTs marked with brown basally, each bearing one to two long, thorny setae, sometimes a long, smooth, hooked seta, three to four shorter, smooth or thorny setae (L3) (two long, smooth or thorny, hooked or pointed setae on L2). S1, S2 absent; SSp small or indistinguishable from SMS; one to three pairs of Fig. 7. Metathorax of Chrysopodes semaphoront B (sec- ond and third instars), lateral view. LT ϭ lateral tubercle; thorny or smooth SMS between or slightly above R1 ϭ row of setae. Note that R1extends from the uppermost LDTs; two to three transverse rows of hooked, smooth edge of the segment and that the setae in R1arise from large SMS anteriorly; total number of SMS in all rows ϭ chalazae that are marked with brown anteriorly. The mark- Ϸ20Ð80 (L3) or 10Ð20 (L2). Venter with pair of lat- ings on the chalazae extend anteriorly onto the surface of the eroventral tubercles (LVT) each bearing two setae segment. stemming from brown setal bases, with small brown marks mesally. A7: LTs hemispherical, well-formed, unmarked or marked with brown dorsally, each bearing eight or S2 very small; transverse row of setae (R1) on crest of nine long, pointed, thorny LS apically and subapically, elevated subsegment, with six or more pairs of large, two to three pointed, smooth LS basally; LS restricted pointed thorny setae (Þve to seven pairs in L2), stem- to LT, not extending mesally onto body of segment. ming from large, light brown to brown chalazae, with LDTs prominent, each bearing one long, thorny seta, brown marks extending anteroventrally beyond cha- four to Þve shorter, smooth LDS (L3) (one in L2). S1, lazae. Venter sometimes with white to brown sclerite S2 small, if present; SSp absent; one to two pairs of mesal to coxa, bearing two short setae. Episternum small, pointed, smooth SMS between LDTs. Venter large, trapezoidal, amber to dark brown, prominent with pair of ventrolateral tubercles each bearing one seta on distal margin, setal base brown; epimeron to two setae; one to two pairs of mesal setae stemming small, triangular, amber to dark brown, with promi- from brown setal bases. nent seta distally. A8: Shorter and narrower than A7. Dorsum with Abdomen (Fig. 8). Cream, unmarked to densely pair of prominent brown marks. LTs small, bearing covered with dark marks, moderately gibbous, broad- est at A2ÐA5, tapering posteriorly. Lateral tubercles two to Þve short LS (one on L2). SSp, S1small if (LT) on A2 to A5 of moderate size, so abdomen does present; S2 absent; one to Þve pairs of small to very not appear signiÞcantly broader than mesothorax or small setae posteromedially. Venter with small to metathorax. medium-sized setae mesally and posteriorly. A1: Shorter and narrower than A2 through A7, A9: Tubular. Dorsum with one to four brown marks rising slightly above metathorax. LTs and latero- mesally, two transverse rows of sparse setae. Venter dorsal tubercles (LDTs) absent. Anterior setae with one to six pairs of long setae. (S1, S2) absent; spiracular setae (SSp) indistinguish- A10: Dorsum cream-colored, with brown triangular able from submedian setae (SMS); SMS/SSp long, mark apically; with one to two pairs of setae mesally, smooth-hooked, generally curving posteriorly, ex- three to fours pairs of short setae in membranous tending from brown or amber setal bases in two trans- triangular area around anus. Pleuron cream-colored, verse bandsÑone anterior, one posterior; total num- with one to two pairs of long setae. Venter brown to ber of SMS/SSp in both bands ϭ 150Ð250 (L3), 48Ð75 cream-colored; posterior with transverse row of short (L2). setae. 480 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA Vol. 96, no. 4

Fig. 8. Dorsum of third-instar Chrysopodes collaris abdomen, dorsal view (illustrating primary structures of Chrysopodes semaphoront B). A1ÐA10 ϭ abdominal segment; LT ϭ lateral tubercle; LS ϭ seta on lateral tubercle; LDT ϭ laterodorsal tubercle; LDS ϭ seta on laterodorsal tubercle; Sp ϭ spiracle; SSp ϭ seta associated with Sp; SMS ϭ submedian seta.

Semaphoront A (L1). Body. Cream-colored to with lateral seta. Labium cream-colored to light am- white, with amber to brown marks; most setal bases ber; palpiger with two pairs of setae ventrally; palpus light brown to brown. Integument largely smooth; with basal segment usually bearing one long lateral spinules absent. Thorny setae with thorns dense, seta distally, one long or short seta mesally; second raised off surface of seta, arranged irregularly along segment annulated, with amber markings distolater- length of seta. ally, two long setae distally (one dorsal, one ventral); Head (Fig. 9). Cream-colored to white, with amber distal segment tapered apically, without setae. An- to dark brown marks similar to those on L2, L3. Genal tenna with ßagellum bearing one long apical seta, one marks pronounced, bifurcated anteriorly. All primary to two microsetae subapically; pedicel with one short, cranial setae (S1ÐS12, Vx) present; S1, S4, S11 usually thick apical seta, sometimes one to two medial setae; thorny, (sometimes S6). Eyes dark brown to black. scape without setae. Cervix white, without setation or Labrum with three pairs of distal setae (mesal pair with very small setae; pleural region with large, light long, lateral pairs shorter). Mandible amber, with four brown to brown mark. to Þve apical teeth, dorsal surface generally smooth, Thorax (Fig. 10). LTs well developed, with two base with dorsal row of four oval structures (placoid (T1) or three (T2, T3) long, robust, crooked or organs of Rousset 1966). Maxilla with three to six small straight, pointed, thorny LS and sometimes very small subapical setae, apical brush of microsetae; venter seta apically. Epimera and episterna large, robust, July 2003 GENETIC CHARACTERISTICS OF Chrysopodes 481

two very small setae on margin (S1Sc1, S2Sc1); Sc2 mesal to LT, with one very small seta attached mesally (S1Sc2); posterior row of two to three pairs of long, robust, thorny setae (R1) stemming from large, light brown to brown chalazae at crest of fold; chalazae with elongate brown marks anteriorly. Abdomen (Fig. 10). A1: LTs absent. S1, S2 absent; SSp long, smooth, hooked, extending from chalaza mesal to spiracle; anterior with row of two to six pairs of long smooth, hooked SMS stemming from large chalazae; posterior row with four to Ϸ18 pairs of smooth, hooked SMS, stemming from large chalazae; lateral-most two pairs of SMS sometimes located very close to each other, comprising an LDT. A2ÐA7: LTs well developed, each bearing two long, thorny LS, each with brown basal mark extending ventrolaterally. S1, S2 absent. A2ÐA5: SSp long, smooth, hooked; anterior row of two to three pairs of long, smooth, hooked SMS stemming from chalazae; posterior row of four to ten pairs of long, smooth, hooked SMS stemming from chalazae; sometimes two lateral-most SMS located very close to each other, forming an LDT. Pleuron sometimes with long smooth, hooked seta stemming from chalaza ventral to LT (seta small on A2, A3, larger on A4, A5). Venter with pair of thorny setae stemming from small cha- lazae. A6ÐA7: LTs each with two long, pointed, thorny setae. LDTs each with one long, thorny, hooked or pointed seta, one shorter, smooth or thorny, straight or hooked seta. S1, S2 absent; SSp small to very small, pointed, smooth. A6: dorsum with two pairs of long, Fig. 9. Dorsum of Þrst instar Chrysopodes collaris head. smooth, hooked SMS; venter with zero to one pair of epi-l ϭ lateral epicranial mark; epi-m ϭ mesal epicranial long lateral setae, two pairs of short lateral setae. A7: mark; fr ϭ frontal mark; post ϭ postfrontal mark. dorsum with two pairs of short SMS; venter sometimes with pair of short, thorny setae. A8: With acute ventral bend. LTs each with one medium-length, thorny LS apically, one short hooked, brown to dark brown. Legs white to amber, except smooth LS subapically. S1, S2, SSp absent or very small; base of coxa marked with light brown; apex of coxa zero to one pair of SMS (medium length); venter with white; femur sometimes with brown band mesally; approximately two to six short setae. A9: two trans- empodium light brown to brown. verse rows of medium length, smooth or thorny setae. T1: Sc1 rhomboidal, with white or cream-colored A10: two to seven pairs of setae laterally. area mesally (sometimes pale, indistinguishable); S1, S1Sc1 thorny, stemming from brown chalazae; S3 Biology thorny or smooth; S2, S4 absent or very small; R1, S5 absent. Very little is known about the biology of Chrysopo- T2: Anterior subsegment bearing spiracles laterally des. Adults are usually collected in open areas or forest on prominent tubercles, sclerites on anterior margin edges, rather than deep in the forest. Some species (Sc1) white to light brown, each bearing two very (e.g., Ch. collaris, Ch. divisa) can be abundant in ag- small setae (S1Sc1, S2Sc1), one on each side. Posterior ricultural situations (Adams and Penny 1987, de Frei- subsegment with sclerites on anterior margin (Sc2) tas and Penny 2001) and may have a role in biological white to light brown, each bearing two very small control. setae (S1Sc2 and S2Sc2) laterally; pair of larger, round Chrysopodes adults are probably not carnivorous. sclerites (Sc3) mesal to LTs, each with one to two Oviposition occurs in the laboratory when adults do setae (S1Sc3 and S2Sc3) attached mesally; S1Sc3 not receive prey, and the guts of Þeld-collected spec- small, S2Sc3 long, thorny, extending from chalaza; one imens were found not to contain remains. The pair of setae (S1) mesal to Sc3 long, thorny or smooth; guts of one species were full of pollen grains (Brooks S2 absent. and Barnard 1990). T3: Segment folded, with posterior fold conspicu- Eggs of the species that we (C.A.T. and M. J. ously raised above dorsum of T2 and anterior subseg- Tauber) have reared are stalked and laid singly. Two ment of T3. Sc1uncolored, sometimes bearing one to types of defensive behavior are apparent. First, fe- 482 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA Vol. 96, no. 4

Fig. 10. Primary thoracic and abdominal structures of Chrysopodes semaphoront A (Þrst instar), dorsal view. T1, T2, T3 ϭ thoracic segment; A1ÐA10 ϭ abdominal segment; LT ϭ lateral tubercle; LS ϭ seta on lateral tubercle; Sp ϭ spiracle; SSp ϭ seta associated with Sp; Sc ϭ sclerite; Sx ϭ seta; SxScx ϭ seta associated with sclerite; R1 ϭ row of setae; LDT ϭ laterodorsal tubercle; LDS ϭ seta on laterodorsal tubercle; SMS ϭ submedian seta.

males of at least some Chrysopodes species place drop- Comparison between Chrysopodes and lets of an oily secretion on the egg stalks. Similar Ceraeochrysa Larvae types of droplets have been shown to protect Ceraeo- chrysa eggs from natural enemies (Eisner et al. 1996). Chrysopodes and Ceraeochrysa larvae (all instars) Second, the larvae carry camoußaging plant and/or are distinguished by numerous qualitative and quan- material on the dorsal surfaces of their titative morphological differences (Tables 1and 2) thorax and abdomen. Such packets of trash have been that are clearly consistent with generic-level differ- shown to protect chrysopid larvae from attack by ences found among European, Japanese, and other predators, especially ants (Eisner and Silberglied 1988, New World genera (c.f., Dõ´az-Aranda and Monserrat Milbrath et al. 1993). 1995, Tsukaguchi 1995, Tauber et al. 2000). Indeed, at July 2003 GENETIC CHARACTERISTICS OF Chrysopodes 483

Table 1. Differences in larval characteristics between Ceraeochrysa and Chrysopodes (Semaphoront B, second and third instars) and the characteristics that support placing C. placita in Chrysopodes

Ceraeochrysa C. placita Chrysopodes Head C-S12 present/absent present present Epicranial mark entire divided divided Genal mark unforked bifurcated anteriorly bifurcated anteriorly Spiracles circular cylindrical, with distinct sides cylindrical, with distinct sides Setal bases white-cream tan-brown tan-brown Thorax Spiracular tubercle (T2) absent present present (sometimes retracted) T1-S1 smooth/thorny thorny* thorny T1-S2&S3 smooth thorny* thorny T1-S5 present absent absent T2-S2 smooth/thorny thorny* thorny T3-posterior subsegment unraised raised above anterior subsegment raised above anterior subsegment T3-R1(L3, No. pairs thorny setae) 2Ð4 8 (eastern U.S.A.) 6Ð10 12 (western U.S.A.) T3-R3 chalazae brown anteriorly no yes yes T3-S2 present absent absent Abdomen A1-LDTs present (small) absent absent A1(No. pairs sm-h SSp/SMS) L3 6Ð12 ϳ75 (eastern U.S.A.) 75Ð125 ϳ110 (western U.S.A.) L2 ϳ5Ð7 ϳ29 24Ð40 A2-A5 (No. pairs sm-h SMS) L3 ϳ20Ð40 ϳ75Ð120 70Ð120 L2 ϳ15Ð30 10Ð15 20Ð50 A6-LT papilliform spherical spherical A6 (No. pairs sm-h SMS) L3 ϳ7 ϳ20Ð25 10Ð40 L2 ϳ1Ð6 5 5Ð10

* Thorniness visible with Ͼ200ϫ. C-S12 ϭ cervical seta 12; T1, T2, and T3 ϭ pro-, meso-, and metathorax; A1 ϭ abdominal segment 1; LDT ϭ laterodorsal tubercle; LDS ϭ seta on LDT; SMS ϭ submedian seta (dorsal seta between LDTs); SSp ϭ spiracular seta; sm-h ϭ smooth, hooked. this time, the generic differences between the larvae given that both the male and female terminalia may of Chrysopodes and Ceraeochrysa appear to be clearer more closely resemble those of Chrysopodes than and more consistent than those found in the adult Ceraeochrysa (see above), Ce. placita is herein moved stage. to the genus Chrysopodes. The red marks on the genae Given that Ce. placita larvae exhibit all of the char- and vertex of Ce. placita, as well as the slightly widened acter states that distinguish known Chrysopodes larvae costal cells are also more consistent with Chrysopodes from those of Ceraeochrysa (Tables 1and 2), and also than Ceraeochrysa.

Table 2. Differences in larval characteristics between Ceraeochrysa and Chrysopodes (Semaphoront A, first instar) and the char- acteristics that support placing C. placita in Chrysopodes

Ceraeochrysa C. placita Chrysopodes Head C-S12 present/absent present present Epicranial mark entire divided divided Genal mark unforked bifurcated anteriorly bifurcated anteriorly Thorax T2 - Spiracular tubercle absent present present T1-S1 smooth/thorny thorny* thorny T1-S2 & S3 smooth thorny* thorny T1-S5 present absent absent T2-S2 present absent absent T3-R1(No. pairs thorny setae) 2 3 2Ð3 Abdomen LDTs (A1) present absent absent SMS ϩ LDS (A1) No. pairs sm-h 4 ϳ15 6Ð24 LDTs (A2-A5) present absent absent or present SMS (A2-A5), No. pairs sm-h 3 7 6Ð13

* Thorniness visible with Ͼ200ϫ. S12 ϭ cervical seta 12; T1, T2, T3 ϭ pro-, meso-, metathorax; A1-A7 ϭ abdominal segments 1Ð7; LDT ϭ laterodorsal tubercle; LDS ϭ seta on LDT; SMS ϭ submedian seta (dorsal seta between LDTs); sm-h ϭ smooth, hooked. 484 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA Vol. 96, no. 4

Fig. 11. Adult head and prothorax, dorsal view; mandibles. (A) Chrysopodes placita. (B) Chrysopodes collaris.

Chrysopodes Species from the United States for Ch. placita and Ch. divisa reßects the absence of clear and consistent differences between the subgen- Chrysopodes (Neosuarius) placita (Banks) (New era; the validity of the subgenera will be considered Combination) later, in a revision of the genus. . Described as Chrysopa placita Banks Descriptions of Ch. placita adults, larvae, and bio- 1908: 259. Male syntypes: United States: Colorado: logical characteristics can be found in Tauber et al. Chimney Gulch, 20.I.1907; Clear Creek (Museum of (1998, 2000) [under the name Ceraeochrysa placita Comparative Zoology) (Penny et al. 1997) (Banks)]. Below is a summary of the information for Moved to Ceraeochrysa (Adams 1982: 73) comparative purposes and for convenience in identi- Chrysopa forreri Nava´s ϭ Ce. placita (Adams 1982: Þcation. 73) Adult. Although Chrysopodes adults can be identi- Chrysopa intacta Nava´s ϭ Ce. placita (Brooks and Þed to species reliably only by examining the male or Barnard 1990: 269) female terminalia, some external characteristics are Chrysopodes (Neosuarius) placita (Banks) (New distinctive. Both Ch. collaris and Ch. placita adults lack combination) distinct stripes on the scape and the pronotum, but It was difÞcult to assign Ch. placita to either of the each species (most specimens) has distinctive red two Chrysopodes subgenera (Chrysopodes or Neosu- markings on the vertex and near the eyes (Fig. 11). In arius), but based on the speciesÕ overall greater afÞnity Ch. placita, the prothorax is elongate and the dorsum with Neosuarius, it is tentatively assigned to this sub- of the thorax usually has a medial yellow stripe, as well genus. The adult mandibles are relatively broad and as submedial red marks; the costal area is slightly blunt-tipped as in Neosuarius, but they are largely widened. symmetrical and only slightly toothed, as in the sub- Ch. placita male terminalia are distinctive (Figs. 1 genus Chrysopodes (Fig. 11A). The vertex is unpitted, and 2): the gonarcus is broad, and the gonarcal and the palps are broad and tapering as in Neosuarius; apodemes are expanded. The transverse bridge of the but the forewings are broad and setose and the costal gonarcus bears a short arcessus, which consists of a area is widened as in Chrysopodes. The species also pair of articulated rods enclosed within a membrane does not resemble Ch. (Neosuarius) divisa, which (these rods are not fused with the gonarcus and are no Brooks and Barnard (1990) consider to be interme- longer considered gonocornua, as originally proposed diate between the two subgenera. Unlike Ch. divisa, by Tauber et al. 2000). Entoprocessus are absent. The Ch. placita does not have narrow, asymmetrical man- gonosaccus is bulbous and has large lateral lobes and dibles. Moreover, neither the palpus nor the galea is a dorsal trough; it lacks gonosetae. The gonapsis is an narrow, and the papilla on the galea is small. However, elongate rod, bowed at midlength, with serrations at as in Ch. divisa, the costal area is slightly broadened. the tip. The female terminalia (Figs. 3 and 4) are I suspect that the difÞculty in subgeneric assignments typiÞed by a round spermatheca with a shallow ventral July 2003 GENETIC CHARACTERISTICS OF Chrysopodes 485

Fig. 12. First instar thoracic and abdominal segments. (A) Chrysopodes placita. (B) Chrysopodes collaris.T3ϭ metathorax; A1ÐA5 ϭ abdominal segment; LT ϭ lateral tubercle; LS ϭ seta on lateral tubercle; Sp ϭ spiracle; SSp ϭ seta associated with Sp; R1 ϭ row of setae; LDT ϭ laterodorsal tubercle; LDS ϭ seta on laterodorsal tubercle; SMS ϭ submedian seta. invagination and a simple, elongate velum with a cranial setae, and their head markings are narrow and single fold. The spermathecal duct is short and narrow; have straight (not jagged) margins; the lateral epicra- the terminus is brushy. The bursal duct is short and nial markings are contiguous (not broken); and the coiled. prothoracic and mesothoracic setae and setal bases Larva. The three instars of Ch. placita were de- are usually dark brown (not pale). [Populations of scribed in detail by Tauber et al. (1998, as Ceraeochrysa Ch. placita from the western United States may have placita), and thus are not redescribed here. Ch. placita light amber thoracic setae and setal bases]. Head larvae resemble those of Ch. collaris, but a suite of widths range from 0.75 to 0.83 mm (L3), 0.55Ð0.57 mm characters distinguishes the two species. Ch. placita (L2), and 0.39Ð0.42 mm (L1). The ratios of mandible second and third instars are relatively small in size and length to head width are relatively small: 0.83Ð0.94: 1.0 they have short mandibles (Fig. 5A). They lack thorny (L3), 0.76Ð0.88 : 1.0 (L2), and 0.74Ð0.87 : 1.0 (L1). 486 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA Vol. 96, no. 4

Fig. 13. Chrysopodes collaris male terminalia; gonarcus complex expanded. arc ϭ arcessus; dapo ϭ dorsal apodeme of 9Tϩect; gs ϭ gonarcus; gsac ϭ gonosaccus; p.t. ϭ terminal projection of the dorsal apodeme of 9T; 8Sϩ9S ϭ eighth and ninth sternites (fused); 8T ϭ eighth tergite; 9Tϩect ϭ ninth tergite and ectoproct (fused).

First instars of C. placita are reliably distinguished A list of synonyms is given by Penny et al. (1997) and from those of Ch. collaris by their setal pattern (Fig. Adams and Penny (1987) described the adults; and 12A and B). The posterior row of setae on the met- Smith (1931) provided a small illustration of the third athorax (T3-R1) of Ch. placita has three (not two) instar (as Chrysopa thoracica Walker). It is believed pairs of thorny setae, and abdominal segments 1Ð4 that this species has a potential role in biological con- each have Ϸ15 pairs (not seven pairs) of long, hooked, trol (e.g., Adams and Penny 1987), especially in or- smooth setae (SMS and SSp). chards or ornamental plantings. Egg. Described by Tauber et al. (1998). Ovoid, Adult. Adult Ch. collaris are distinguished from stalked. Stalk length geographically variable (Eastern other green lacewings in the United States by their United States: 2.1Ð3.3 mm; Western United States: conspicuous red or pink head markings (Fig. 11B): 1.4Ð2.3 mm). faint oblique marks along the upper border of the Distribution. Southern Canada, eastern and west- antennal sockets, stripe along the margin of the eyes, ern United States, northern Mexico. Records include: crescent-shaped marks below the antennae, trans- United States: Arizona, California, Colorado, Missouri, verse bands across the frontoclypeal suture, lateral New Hampshire, North Carolina, New Mexico, New marks on the clypeus and broad genal stripes. Both York, Oregon, West Virginia, Tennessee, Texas, Wy- sexes are typiÞed by a relatively large, robust body, oming, Utah; Canada: British Columbia, Ontario, Que- short prothorax, yellow pronotal stripe, yellowish an- bec; Mexico: Michoaca´n (Penny et al. 1997, Tauber et tennae, and distinct genital structures (Figs. 13Ð16; al. 1998, 2000; J. D. Oswald, personal communication). descriptions below and in Adams and Penny 1987). Habitat. Ch. placita has been collected in temperate The wing venation is mostly green, with some black forests up to 2,800 m. Eggs and larvae occur on the veins in the forewings; the costal area is not widened trunks of intermediate-sized to mature deciduous and (see Adams and Penny 1987). evergreen trees, especially those with moss or lichen In the male, the arms and bridge of the gonarcus are growth. broad and heavily sclerotized. The triangular arcessus Biology. Ch. placita is largely a late-summer species; is supported basally by a convex, “H”-shaped internal most records for adults and larvae are from August to brace; it terminates in a setose, hook-shaped apical October (a few from June, in AZ). Larvae probably projection (Fig. 14). A gonapsis is absent. The female feed on psocids and other soft-bodied that terminalia are typiÞed by a relatively wide sper- are associated with lichens and moss on tree trunks. matheca with a broad ventral invagination and an Short daylengths retard development during the Þrst elongate, curved velum (Fig. 16A). The bursa is large; instar and induce hibernal diapause in second instars. the spermathecal duct is long and coiled (Fig. 16A). Eggs are attacked by Telenomus and Trichogramma The subgenitale is short, cordate (Fig. 16B). spp. Details on the life history were described by Larva. Second and third instars of Ch. collaris differ Tauber et al. (1998). from those of Ch. placita by: their relatively long man- dibles, thorny cranial setae (L2), dark brown head marks, jagged (not straight) submedian head marks Chrysopodes (Neosuarius) collaris (Schneider, 1851) [in which the anterior, postfrontal mark (post) is Ch. collaris is the only species of Chrysopodes pre- fused with, but distinct from the posterior, submedian viously recorded from the U.S.A. (Penny et al. 1997). epicranial mark (epi-m)], broken (not contiguous) July 2003 GENETIC CHARACTERISTICS OF Chrysopodes 487

Fig. 14. Chrysopodes collaris male genital structures (redrawn from Adams and Penny 1987). (A) Gonarcus complex (expanded), dorsal view. (B) Gonarcus complex (expanded), lateral view. arc ϭ arcessus; gsac ϭ gonosaccus; la ϭ lateral arm of gonarcus. lateral epicranial mark (epi-l) (Fig. 5B). Prothoracic Central America, and westward in northern Mexico to and mesothoracic setae and setal bases are cream- Baja California (Adams and Penny 1987). Records colored or golden (not dark brown). Head widths from South America include Colombia (Cali, 1,000 m), range from 0.94 to 1.01 mm (L3), 0.61Ð0.65 mm (L2), Venezuela and Guyana (Adams and Penny 1987). The and 0.41Ð0.46 mm (L1). The ratios of mandible to head larval specimens described here were reared from width are large: 1.0Ð1.1 : 1.0 (L3), 0.9Ð1.1 : 1.0 (L2), adults collected in Tampico, Tamaulipas, Mexico, and 0.9 : 1.0 (L1). December 1963 (M. J. Tauber and C. A. Toschi, In C. collaris Þrst instars, the posterior row of setae collectors, Tauber lot 63:86), from Ciudad Victoria, on the metathorax (T3ÐR1) has only two (not three) Tamaulipas, Mexico, February 1999 (E. Ya. Chouva- pairs of thorny setae, and abdominal segments 1Ð5 khina, collector and rearing), and from the southern each have seven (not 15) pairs of long, smooth-hooked end of Jost Van Dyke, in the British Virgin Islands, setae (SMS, SSp). November/December 1994 (L. M. Wilson and J. B. Egg. Ovoid, stalked, laid singly without a discernible Johnson, collectors; reared by J. B. Johnson). Field- pattern; stalk with oily droplets; stalk length 3.2Ð3.9 collected larvae were from Laguna Santa Maria, 20 air mm. miles SSE Tepic, Nayarit, Mexico, December 1963 Distribution. In the United States, Ch. collaris is (M. J. Tauber and C. A. Toschi, collectors). reported only from Texas and Florida, but the species Habitat. Ch. collaris has been collected at light and is also known from throughout the Caribbean region, by beating trees and shrubs. In Haiti the species was

Fig. 15. Chrysopodes collaris female terminalia (redrawn from Adams and Penny 1987). b.d. ϭ bursal duct; coll. g. ϭ colleterial gland; cop.b. ϭ copulatory bursa; g.l. ϭ gonopophysis lateralis; sg ϭ subgenitale; sp ϭ spermatheca; 7S ϭ seventh sternite; 8T ϭ eighth tergite; 9Tϩect ϭ ninth tergite and ectroproct (fused). 488 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA Vol. 96, no. 4

Fig. 16. Chrysopodes collaris female genital structures (redrawn from Adams and Penny 1987). (A) Spermatheca, lateral view. (B) Subgenitale, ventral view. b.g. ϭ bursal gland; b.d. ϭ bursal duct; cop.b. ϭ copulatory bursa; sp ϭ spermatheca; sp.d. ϭ spermathecal duct; v.i. ϭ ventral invagination. found on bamboo (Smith 1931). In Florida it was bifurcated distally (at least for one half its length). S1 collected from Hibiscus, a large ornamental shrub smooth to slightly thorny, other cranial setae smooth; (H. W. Browning, collector) and Eugenia, another patch of Ϸsix small secondary setae between S1and shrub (Eisner and Silberglied 1988). S2. Labial palp with distal annulation of second seg- Biology. The larvae on Eugenia in Florida were ment bearing one long lateral, two medial setae, with associated with colonies of mealybugs (Plotococcus remaining annulations bearing ϷÞve dorsal setae. eugeniae Miller and Denno) that produce white, waxy Scape light brown; pedicel and ßagellum amber. Ven- coverings (Eisner and Silberglied 1988). Presumably, ter of head white, except lateral margins of cranial the Ch. collaris larvae transfer wax from the mealybugs capsule amber; cardo and base of stipes dark brown. to their dorsa. Cervix cream-colored dorsally, with two longitudi- Developmental rates appear to be variable. In our nal bands of light tan spinules, one to three pairs of rearings (Ϸ21ЊC, LD 16:8, aphid prey): egg: 7Ð10 d; microsetae dorsolaterally; venter with pair of brown Þrst instar: 9Ð10 d; second instar: 7Ð10 d; third instar marks laterally; pleuron with dark brown spinules, Ϸ14 d; no diapause observed. In earlier rearings (con- amber marking. ditions unspeciÞed): egg stage: 4 d; larval stage: 14 d; Thorax (Fig. 6). White to cream-colored, with dor- period within cocoon: 11 d (Smith 1931). sal sclerites brown to light brown. Lateral setae (LS) white, stemming from white to amber bases. Lateral tubercles (LT) with Ϸ18 (prothorax), Ϸ15 (mesotho- Descriptions of Ch. collaris Larvae rax), Ϸ15 (metathorax) prominent, thorny setae (LS). Semaphoront B (L3). Body. White to cream-col- Venter cream-colored mesally, becoming light brown ored, without dorsal stripe; 5.2Ð7.7 mm long. Setae near coxae. Legs white to cream-colored, except: base white to pale amber as noted below. of coxae with irregularly-shaped brown marks dor- Head (Fig. 5B). Cream-colored; 0.94Ð1.01 mm sally; tips of femora with very small dark brown marks; wide. Mandible long, ratio of mandible length : head tarsi and tarsal claws, empodia brown distally. width ϭ 1.0Ð1.1: 1.0. Dorsal marks brown. Epicranial T1: Paired notal sclerites (Sc1) brown laterally, mark divided into two sections: mesal epicranial mark white interiorly, with anterior margin extending for- (epi-m) narrow, extending longitudinally from base of ward almost to base of LT. Median sclerite (Sc2) very cranium, becoming conßuent with postfrontal mark small, light brown. (post); lateral epicranial mark (epi-l) consisting of T2: Spiracular tubercle raised; anterior sclerites triangular marks basally (at lateral margin of cranial (Sc1) prominent, brown. Posterior subsegment with base), scattered light brown marks distally (approach- anterior sclerites (Sc2) small, light brown; Sc3 dark ing eyes). Postfrontal mark (post) dark brown medi- brown. Pleuron with small dark brown markings pos- ally, becoming light brown distally, conßuent medially teroventral to LTs, patches of light brown spinules with mesal epicranial mark. Frontal mark (fr) light mesally. brown, extending anteriorly from center of cranium to T3: Sclerite Sc2 small, dark brown; R1with ten to level of antennal bases. Area between eye and lateral eleven pairs of long, thorny-pointed setae extending base of mandible with small dark brown marks. Cly- from brown chalazae in two rows Ð Þve anterior pairs peal region cream-colored, unmarked. Genal mark with elongate brown marks extending anteroventrally. brown, extending from cervix almost to eye, deeply Pleuron with amber markings below tubercles. July 2003 GENETIC CHARACTERISTICS OF Chrysopodes 489

Abdomen (Fig. 8). White to cream-colored, with hooked, smooth LS. A2: four pairs of hooked, smooth dense covering of amber to light brown spinules. Setae SSp, Ϸ55Ð60 hooked, smooth SMS. A5: three pairs of generally brown, extending from brown setal bases; SSp, 20Ð25 hooked, smooth SMS. spiracles brown. Venter cream-colored, with dense A6: LTs with Ϸsix pairs of long, thorny LS and one covering of light brown spinules. hooked, smooth LS apically. Each LDT with one long, A1: Ϸ150 hooked submedian setae (SMS) in two serrated LDS, four to Þve pointed, smooth LDS; an- double or triple rows. terior row of Ϸ14Ð15 hooked, smooth SMS, pair of A2Ð5: LTs with apical three to four LS thick, short SMS between LDTs. hooked, thorny; subapical LS thinner, hooked, A7: LTs with Ϸsix pairs of long, serrated LS apically; smooth, some straight, thorny; each LT with small, two to three pairs of short, pointed, smooth LS sub- dark brown basal mark anteriorly, another posteriorly; apically. patch of short, hooked, smooth setae anterobasal to A8: LTs each bearing three short, one long LS; two LT. Patch of eight (A5) to 16 (A2) long, hooked pairs of pointed, smooth setae mesally between LTs. spiracular setae (SSp) anterior to spiracles; Ϸ140 A9: Row of six setae mesally; row with eight setae (A2)Ð100 (A5) hooked, smooth SMS in three double posteromesally. or triple rows; pleural region with two to three A10: Forked marking dorsally, numerous pointed, pointed, smooth setae. A3Ð5: venter with pair of very smooth setae ventrally. large white chalazae each bearing long seta stemming Semaphoront A (L1). Body. Very gibbous, with from brown setal base; on A4Ð5 each chalaza sur- abdomen strongly bent ventrally; 2.3Ð3.1mm long. rounded with brown. White, with light brown to brown marks. Venter A6: Anterior area with Ϸ35 smooth, hooked SMS in white, without marks except ventrolateral tubercles two irregular rows. LDTs with dark brown mark ba- (VLTs) of A2ÐA7 brown. solaterally; each LDT bearing one long, thorny LDS, Head (Fig. 5A). Amber; 0.41Ð0.46 mm wide. Ratio three pointed, smooth LDS; two pairs of short SMS of mandible length : head width ϭ 0.90Ð0.94 : 1.0. between LDTs. SSp small. Venter with small laterov- Head marks not distinct. Epicranial marking divided entral tubercle (LVT) bearing two setae stemming longitudinally; lateral epicranial mark (epi-l) extend- from brown bases, with small brown marks mesally. ing from base of head around eyes; mesal epicranial A7: LTs with brown mark ventrolaterally. LDTs mark (epi-m) conßuent with postfrontal (post) and prominent, white, surrounded by dark brown. One to frontal (fr) marks; entire submedial marking (includ- two pairs of small SMS between LDTs; one pair very ing postfrontal marking) extending from base of head small setae anteriorly. VLTs each bearing two setae to tip of labrum. S1, S4, S11 long, robust, thorny. Eyes from brown setal bases; venter with two transverse black. Cervix white, with light brown marking later- rows of setae extending from brown setal bases, two ally. pairs of setae in each row. Thorax. White, with light brown marks. T1-S3 A8: Two pairs of very small setae anteromesally; row thorny. T2-S1, T2-S1Sc3 robust, thorny. T3-R1 with of ϷÞve pairs of small setae posteromedially. LTs each two pairs long, thorny setae (Fig. 12B). bearing ϷÞve short LS. Venter with two pairs of small Abdomen. A1ÐA5 (Fig. 12B): anterior row with two setae anteriorly, Ϸthree pairs of short setae mesally, pairs SMS, posterior row with one pair SSp, four pairs Ϸtwo pairs of longer setae posteriorly. of long smooth, hooked SMS. Lateral-most SMS in A9 (Fig. 7): Dorsum with four large brown marks: posterior row located very close to each other, form- anterior one oblong, mesal pair circular, posterior one ing an LDT. A2ÐA5: Pleuron with long smooth, hooked irregular-shaped. Three pairs of small setae mesally; seta stemming from chalaza ventral to LT (seta small posterior row of Ϸsix pairs of longer setae. Pleuron on A2, A3, larger on A4, A5). A8: short, with acute with pair of long setae posteriorly. Venter with pair of ventral bend. A10: two pairs of lateral setae. very small setae anteriorly. Variation. Smith (1931) noted considerable vari- A10 (Fig. 7): Dorsum and pleuron with two pairs of ation in the red coloration on the head and body long setae. among his adult specimens from Haiti. In the current Semaphoront B (L2). Similar to L3, with the fol- study, specimens (adults and larvae) from the Carib- lowing exceptions: bean Islands (British Virgin Islands) had considerably Body. 3.8Ð4.7 mm long. paler markings than the continental specimens Head. 0.61Ð0.65 mm wide. Ratio of mandible (United States (Florida), Mexico, and Central Amer- length : head width ϭ 0.9Ð1.1 : 1.0. Frontal marks ica). joined anteromedially by pale subcuticular marking; S1, S4 thorny; second segment of labial palpus with four to Þve setae (two subapical, two to three mesal). Acknowledgments Thorax. LTs with 13 (T1), 11 (T2), 12 (T3) thorny LS; pair of short smooth setae anterior to each LT. I thank G. S. Albuquerque (GSA), V. Becker, H. W. Browning, E. Ya. Chouvakhina, E. M. G. Fontes, F. Franca, Metathoracic R1with Þve pairs long, thorny setae. J. Gavila´nez, P. D. Hurd (deceased), J. B. Johnson, S. L. Abdomen. A1: ten hooked, smooth SMS anteriorly, Ϸ Lapointe, M. A. de Medeiros, A. Nasciamento, C. S. S. Pires, 28Ð30 SMS in double row posteriorly. E. R. Sujii, Agatha J. Tauber, Michael J. Tauber, and Paul J. A2ÐA5: two (A2) or three (A3-A5) pairs of long Tauber for help in obtaining specimens; P. A. Adams, E. G. thorny LS; six (A3ÐA5) or seven (A2) pairs of shorter, MacLeod (both deceased) and N. D. Penny for assistance 490 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA Vol. 96, no. 4 with some of the identiÞcations; J. I. Lo´pez-Arroyo for the larval feeding and defensive behavior. Ecology 74: 1384Ð Spanish translation of the abstract; J. D. Oswald for providing 1393. comments on the manuscript and unpublished information Nava´s, L. 1913. Les chrysopides (Ins. Ne´vr.) du Muse´ede on the occurrence of Ch. placita in TX and NC. I also thank Londres. Ann. Soc. Sci. Bruxelles. 37: 292Ð330. L. E. Ehler, R. E. Page, D. E. Ullman, S. Kelley, and the Penny, N. D. 1997. Four new species of Costa Rican Ceraeo- Department of Entomology, University of California, Davis chrysa (Neuroptera: Chrysopidae). Pan-PaciÞc Entomol. for their cooperation in a variety of ways, and M. J. Tauber 73: 61Ð69. for collaboration on all aspects of the project and comments Penny, N. D. 1998. New Chrysopinae from Costa Rica on the manuscript. The work was supported, in part, by (Neuroptera: Chrysopidae). J. Neuropterol. 1: 55Ð78. USDA/NRI Competitive Grants 9802447 and 2002-35316- Penny, N. D. 2001. New species of Chrysopinae (Neurop- 12231 (M.J.T., C.A.T.), the National Geographic Society tera: Chrysopidae) from Costa Rica, with selected taxo- (M.J.T., C.A.T., G.S.A.), National Science Foundation Grant nomic notes and a neotype designation. Entomol. News. INT-9817231 (M.J.T., C.A.T., G.S.A.), Regional Project 112: 1Ð14. W-185, and Cornell University (M.J.T., C.A.T.). Penny, N. D., P. A. Adams, and L. A. Stange. 1997. Species catalog of the Neuroptera, Megaloptera, and Raphid- References Cited ioptera of America north of Mexico. Proc. Calif. Acad. Sci. 50: 39Ð114. Adams, P. A. 1982. Ceraeochrysa, a new genus of Chrysopi- Principi, M. M. 1977. Contributi allo studio dei Neurotteri nae (Neuroptera). (Studies in New World Chrysopidae, italiani. XXI. La morfologia addominale ed il suo valore part II). Neuroptera International 2: 69Ð75. per la discriminazione generica nellÕambito delle Chry- Adams, P. A. 1985. Notes on Chrysopodes of the M.N.H.N. in sopinae. Boll. Inst. Entomol. Univ. Bologna. 31: 325Ð360. Paris [Neuroptera, Chrysopidae]. Rev. Fr. Entomol., Rousset, A. 1966. Morphologie ce´phalique des larves de Pla- (N. S.). 7: 5Ð8. nipennes (Insectes Ne´vropte´roõ¨des). Me´m. Mus. Nat. Adams, P. A., and N. D. Penny. 1987. Neuroptera of the Hist. Ser. A. 42: 1Ð199. Amazon Basin, Part 11a. Introduction and Chrysopini. Smith, R. C. 1931. The Neuroptera of Haiti, West Indies. Acta Amazonica. 15: 413Ð479. Ann. Entomol. Soc. Am. 24: 798Ð823. Aspo¨ck, U. 2002. Male genital sclerites of Neuropterida: an Tauber, C. A., and T. de Leo´n. 2001. Systematics of green attempt at homologisation (Insecta Holometabola). Zool. lacewings (Neuroptera: Chrysopidae): larvae of Ceraeo- Anz. 241: 161Ð171. chrysa from Mexico. Ann. Entomol. Soc. Am. 94: 197Ð209. Banks, N. 1908. Neuropteroid Ð notes and descrip- Tauber, C. A., T. de Leo´n, J. I. Lo´pez-Arroyo, and M. J. tions. Trans. Am. Enomol. Soc. 34: 255Ð267. Tauber. 1998. Ceraeochrysa placita (Neuroptera: Chry- Brooks, S. J., and P. C. Barnard. 1990. The green lacewings sopidae): generic characteristics of larvae, larval descrip- of the world: a generic review (Neuroptera: Chrysopi- tions, and life cycle. Ann. Entomol. Soc. Am. 91: 608Ð618. dae). Bull. Br. Mus. nat. Hist. (Entomol.). 59: 117Ð286. Tauber, C. A., T. de Leo´n, J. I. Lo´pez-Arroyo, and M. J. de Freitas, S., and N. D. Penny. 2001. The green lacewings Tauber. 2000. The genus Ceraeochrysa (Neuroptera: (Neuroptera: Chrysopidae) of Brazilian agro-ecosystems. Chrysopidae) of America North of Mexico: larvae, adults Proc. Calif. Acad. Sci. 52: 245Ð395. and comparative biology. Ann. Entomol. Soc. Am. 93: Dı´az-Aranda, L. M., and V. J. Monserrat. 1995. Aphidopha- 1195Ð1221. gous predator diagnosis: key to genera of European chry- Tauber, C. A., M. J. Tauber, and G. S. Albuquerque. 2001. sopid larvae (Neur.: Chrysopidae). Entomophaga 40: Plesiochrysa brasiliensis (Neuroptera: Chrysopidae): lar- 169Ð181. val stages, biology, and taxonomic relationships. Ann. Dı´az-Aranda, L. M., V. J. Monserrat, and C. A. Tauber. 2001. Entomol. Soc. Am. 94: 858Ð865. Recognition of early stages of Chrysopidae, pp. 60Ð81. In Tjeder, B. 1970. Neuroptera, pp. 89Ð99. In S. L. Tuxen [ed.], P. McEwen, T. R. New and A. E. Whittington [eds.], TaxonomistÕs glossary of genitalia in insects, 2nd ed. Lacewings in the crop environment. Cambridge Univer- Munksgaard, Copenhagen. sity Press, Cambridge, United Kingdom. Tsukaguchi, S. 1995. Chrysopidae of Japan (Insecta, Neu- Eisner, T., and R. E. Silberglied. 1988. A chrysopid larva that roptera). S. Tsukaguchi, Aioi-cho 6Ð14-102. Nishinomiya- cloaks itself in mealybug wax. Psyche 95: 15Ð19. shi, Hyogo, 662 Japan. Eisner, T., A. B. Attygalle, W. E. Conner, M. Eisner, and Wheeler, Q. D. 1990. Ontogeny and character phylogeny. E. MacLeod. 1996. Chemical egg defense in a green lace- Cladistics 6: 225Ð268. wing (Ceraeochrysa smithi). Proc. Natl. Acad. Sci. U.S.A. 93: 3280Ð3283. Milbrath, L. M., M. J. Tauber, and C. A. Tauber. 1993. Prey Received for publication 4 October 2002; accepted 5 March speciÞcity in Chrysopa: an interspeciÞc comparison of 2003.