Larvae of five horticulturally important species of Chrysopodes (Neuroptera, Chrysopidae): shared generic features, descriptions and keys

Abstract An expanded list of generic level larval characteristics is presented for Chrysopodes; it includes a reinterpretation of the mesothoracic and metathoracic structure and setation. Keys, descriptions and images of Semaphoront A (first instar) and Semaphoront B (second and third instars) are offered for identifying five species of Chrysopodes (Chrysopodes) that are commonly reported from horticultural habitats in the Neotropical region.


introduction
Chrysopodes is one of the main groups of predaceous insects that have value in the biological control of arthropod pests in Neotropical agriculture (Albuquerque et al. 2001, Freitas and Penny 2001, Silva et al. 2007, Gonzalez Olazo and Heredia 2010. The genus is widespread and relatively large; it occurs throughout all of tropical and subtropical America and presently it consists of 47 species (Tauber et al. 2012). Many species in the genus are commonly found in disturbed habitats, most often in orchards and plantations. One species is reported from the United States (see Tauber 2003, Tauber andFlint 2010), substantially more from Mexico and Central America, and many more from South America. Navás (1913) described Chrysopodes on the basis of the external adult features of a single species. Subsequent researchers included ~30 additional species in the genus, recognized generically distinctive genitalic characteristics, and divided the group into two subgenera: Chrysopodes with sickle-shaped mandibles and Neosuarius with broadly-tipped mandibles (Adams andPenny 1985, Brooks andBarnard 1990; also see Banks 1945). Thereafter, other species of Chrysopodes (Chrysopodes) were described (Penny 1998, 2001, Freitas and Penny 2001, Tauber et al. 2012, and the subgenus Chrysopodes (Neosuarius) was revised (Tauber 2010). The subgenus Chrysopodes (Chrysopodes) is currently under revision (Tauber, C. A. in preparation).
Although most of the taxonomic work on the genus Chrysopodes has focused on the adult stage, an extensive suite of morphological traits was shown to distinguish Chrysopodes larvae from those in other genera of Chrysopini (Tauber 2003). To date, species-specific larval characteristics have been described for two Chrysopodes (Neosuarius) species: Chrysopodes (Neosuarius) collaris (Schneider) and Chrysopodes (Neosuarius) porterinus (Navás) (Tauber 2003, Monserrat andFreitas 2005). It is reasonable to expect that further comparative study of the larvae will provide important information for the systematics of Chrysopodes and increase the value of this group of natural enemies for ecological investigations and agricultural use.
With the above in mind, we describe and provide images of the larvae of five additional species of Chrysopodes. All five species are in the subgenus Chrysopodes (Chryso- Also, we present keys for identifying the larvae (all instars) of the five species. Prior to doing so, we make some minor corrections and important additions to an earlier list (Tauber 2003) of larval features that distinguish Chrysopodes. The genus-level features (i.e., those that are shared by larvae of all Chrysopodes species studied to date) are listed on the Appendix.

Methods
The specimens used in our study were reared from field-collected females. The rearing, preservation, descriptive procedures, and terminology are identical to those pub-lished previously (Tauber 2003, http://esa.publisher.ingentaconnect.com/content/esa/ aesa/2003/00000096/00000004/art00008). We suggest that readers refer to the illustrations and explanatory material in that paper when using the keys, descriptions and images here. Voucher specimens (adult females with their laboratory-reared offspring, and the larval specimens used in the study) are deposited in the Essig Museum, University of California, Berkeley, the insect collection at the Universidade Estadual do Norte Fluminense, Campos dos Goytacazes, and the research collections of the authors.
It is noteworthy that bilateral asymmetry in setal numbers is common, and that specimens occasionally exhibit variation in the numbers and sizes of setae (especially in the dorsal thoracic setae and the submedian abdominal setae) of all instars. The numbers presented here reflect this asymmetry. Also, in our descriptions, unless stated otherwise, all setae, other than the submedian setae, are smooth and pointed (not thorny, hooked or blunt).

shared Generic Characteristics
The chrysopid life cycle includes a larval stage with three instars. The first instar differs markedly in structure, setation and often coloration from the other two instars, which differ from each other only in minor ways, largely related to size. Thus, for taxonomic purposes, the first instar constitutes Semaphoront A, whereas Semaphoront B includes both the second and third instars and Semaphoront C encompasses all instars (= the larva) (See Wheeler 1990). We use the term "semaphoront" in our descriptions because of its systematic and phylogenetic value. Specifically: (i) the term highlights the relative degree of morphological change that occurs with each instar during metamorphosis, (ii) it reflects accurately the relative value of the three chrysopid instars to phylogenetic analysis, and (iii) the commonality of the pattern of variation among semaphoronts across chrysopid taxa, and indeed taxa in other insect orders (see Wheeler 1990), itself is of considerable biological interest.
Chrysopid genera fall into two general categories: those with "naked" larvae and those with "trash-carrying" larvae. Chrysopodes larvae are typical examples of the lat-ter -that is, they have compact, globose bodies, hooked abdominal setae, and well developed thoracic and abdominal tubercles that bear elongate setae adapted for carrying small pieces of plant or animal debris. Furthermore, Chrysopodes larvae express a unique set of morphological and setal characters that distinguishes them from the larvae of other trash-carrying genera (Tauber 2003; for additional comparisons, see Díaz-Aranda and Monserrat 1995, Tsukaguchi 1995, Monserrat and Díaz-Aranda 2012. In general, the species studied here exhibit all of the larval characteristics proposed earlier to typify Chrysopodes (Tauber 2003); moreover, several additional characteristic features were discovered during the current study. Thus, we provide an up-dated list of shared Chrysopodes generic-level characteristics (Appendix); those that are new as a result of the current study are marked with an asterisk. Among the most distinctive features that were previously unreported are the uniquely shaped submedian setae (SMS) on the anterior abdominal segments of all instars (Fig. 1C). These long, smooth, hooked setae are slender and bent throughout the midregion, but their hooked tips are robust, rigid and laterally compressed. We have not seen this type of seta on larvae of other neotropical Chrysopini.

Keys to larvae of five Chrysopodes (Chrysopodes) spp. commonly found in Brazilian fruit orchards
Note: To identify cephalic setae, see Fig. 5  Cranium with two dorsal setae (S1 and S11) thorny (Fig. 5A in Tauber 2003); dorsum of abdominal segments A1-A5 each with total of more than 30 elongate, smooth, hooked setae (submedian and spiracular setae) (Fig. 5C)  More than three (usually six) cranial setae (S1, S3, S4, S5, S6, S11) thorny (Fig. 5A in Tauber 2003); large lateral tubercles on thorax and abdomen with setae (LS) light amber to light brown (Fig. 23C)  Only two or three cranial setae (S1, S11, sometimes S4) thorny; large lateral tubercles on thorax and abdomen with setae (LS) dark brown (Fig. 14C)  Discussion. Chrysopodes divisus is probably one of the most common and widespread of the Chrysopodes species. It has a large number of synonyms (see Adams and Penny  1985), and originally it was placed in the subgenus C. (Neosuarius). Recently it was moved to C. (Chrysopodes) on the basis of adult characteristics (Tauber 2010). Adults of C. (C.) divisus are recognized by their relatively narrow costal cells, dark gradate veins, facial markings and very distinctive male and female genitalia. They can be identified using current keys and redescriptions (Adams andPenny 1985, Freitas andPenny 2001).
Larval diagnosis. Chrysopodes (C.) divisus larvae (all instars) are relatively short, compact, and rotund. Their bodies are white to light cream-colored, with brown to light brown prothoracic sclerites; most of the long setae are cream-colored to light brown. The extensive brown head markings of C. (C.) divisus (all instars) are similar to those of C. (C.) lineafrons in that: (i) The confluent frontal markings, together with the dark brown intermandibular region, form an extensive dark brown, triangular to T-shaped mark on the anterior region of the head. (ii) The mesal and lateral arms of the epicranial markings are completely or partially confluent; together with the dark brown postfrontal markings, they cover most of the posterior and mesolateral regions of the head.
The first instar of C. (C.) divisus is distinguished from C. (C.) lineafrons by the lack of thorns on all cranial setae other than S1 and S11, metathorax with a large number of setae (n = 6-7) in row (R1) on posterior fold, and a large number of abdominal SMS on segments A1 to A5 (n > 30). Second and third instars (Semaphoront B) are characterized by cranial setae that are all without thorns, prothorax that is white to cream-colored and without a conspicuous covering of brown spinules, and metathorax that has a large number of setae (n = 14-15) in row (R1), which, in the L3, is flanked laterally by a pair of long, smooth setae.
Gena, ventral margin of cranium brown; genal marking with small white spot behind eye. Labial palpus tinged with light brown, slightly darker distally. Mandibles amber, with brown basolateral spot. Antenna with scape brownish, pedicel white basally, brown distally, flagellum light brown.
Second and third instars. (Semaphoront B). Body (Figs 6E, 7A-B) length 3.3-3.4 mm (L2), 5.7-6.3 mm (L3); surface white to cream-colored, with light to dark brown integumental spinules somewhat dense, dark on pronotum; primary pronotal, mesonotal sclerites brown to dark brown, other dorsal marks small, brown; sclerites around coxae dark brown, abdomen with light to very light brown stripe laterally. Head (Figs 3A, 4A, 6A-B, 7C-D) cream-colored, with extensive brown and dark brown markings. Epicranial marking undivided, with mesal and lateral arms distinguishable, but broadly connected throughout, both arms in contact with posterior margin of head; lateral arm dark brown basally, lighter distally, extending from distal ~one-fourth of posterior cranial suture to lateral base of mandible; mesal arm light brown, extending from base of head, becoming confluent with postfrontal marking. Postfrontal marking very dark brown, narrow throughout, extending to inner base of scape. Frontal marking dark brown, with left, right arms contiguous with each other and with intermandibular marking, forming broad, triangular, dark brown anterior marking. Clypeolabral region distal to anterior marking cream-colored. Gena creamcolored, with brown marking basally, becoming lighter brown, forking near midregion, extending to base of eye. Mandible, maxilla dark brown laterally and distally, amber mesally, base with dark brown mesal mark. Labial palpus: basal segment cream-colored with light brown mesally; mesal segment ringed with light brown laterally, creamcolored mesally, with terminal subsegment brown; terminal segment dark brown basally, light brown distally. Antenna: scape, basal, mesal sections of pedicel cream-colored to amber, distal section of pedicel very light brown, flagellum light brown. Venter cream-colored to white; margin of cranium brown to dark brown; cardo, stipes brown; base of mentum with light brown patch.
All cephalic setae smooth, pointed; S1 slightly robust, medium length, S11 long, S2-10, S12 short to medium length; Vx setae fairly long, robust; with one to two pairs of secondary setae. Anterior margin of head protruding, straight with angled lateral margins; mesal pair of anterior setae much longer than two lateral pairs. Cervix brownish dorsally, slightly darker laterally; venter cream-colored mesally, light brown laterally; white ventrally.
Abdomen (Figs 6D-F, 7A-B, 8B-C) white to cream-colored, tinged with light brown, with small light brown to brown spots anterior to bases of LTs; spots more diffuse on A5-A8; white fat-body visible beneath integument; setae mostly light brown to amber-colored. A6, A7 each with pair of large, dark brown marks surrounding LDTs. A8 with pair of dark brown marks mesal to spiracles; A9 with pair of large, dark brown marks anterolaterally, dark brown mark mesally; A10 with inverted U-shaped, dark brown mark. Venter white to cream-colored, unmarked, except with some light brown pigmentation ventrolaterally on S5-S9 (some specimens); tip of A10 with pair of small, abutting, triangular dark brown marks. A1: Dorsum with ~78-88 (L2), 142-172 (L3) SMS in two double-triple transverse bands between spiracles. A2-A5: Dorsum with 64-112 (L2), 120-196 (L3) SMS in two broad transverse bands. LTs each with 11-21 (L2), 29-40 (L3) LS: one to five robust, thorny, blunt to spatulate LS on distal surface; remaining LS long, smooth, hooked, in large patch on dorsal surface. A6: Dorsum with transverse band of 20-30 (L2), 56-66 (L3) SMS across anterior of segment; midsection with one to two pairs of smooth setae, mesal pair hooked (similar to SMS), lateral pair short, pointed. LT with nine to eleven (L2), ten to eleven (L3) LS of various sizes. A7: Dorsum with one to two pairs of very small setae (S1, S2) anteriorly, between spiracles; LDTs each with one medium-length, robust, thorny, blunt to spatulate LDS, one to two smaller, smooth, pointed LDS; pair of very small setae between LDTs. LT with ten to 13 (L2), ten to 14 (L3) LS of various sizes. A8: Anterior region with one to two pairs of very small setae (S1, S2). Venter with pair of medium-length setae between LTs, two to three smaller setae slightly anteroventral to LTs. A9: Dorsum with one pair of very small setae anteriorly. Middle and posterior regions with two transverse rings of setae extending around segment; each ring with ~14-16 short to medium-length setae, several in each ring robust. A10: Dorsum with several pairs of small setae on V-shaped anterior sclerites, one slightly anterior to terminus. Two pairs of robust lateral setae. Venter with ~five pairs of small setae in V-shaped pattern, posterior row of microsetae anterior to terminus.
Larval specimens examined. Numerous lots, each originating from a single gravid female collected in Brazil, Bahia: Cruz das Almas, VI-19-96 ( Tauber  Biology. Adults and larvae of this species were collected on shrubs in disturbed, dry forest habitats. Adults are agile; they exhibit fast, evasive flight, usually inward toward the interior of the bush or tree. Based on the following observations, we think that C. (C.) divisus adults may enter a diapause-mediated dormancy. Adults collected during the spring in Minas Gerais (Parque Ecológico Cachoeiras do Rio Bonito) were yellow to yellowish brown, and they had a greasy appearance. In addition, their prothoracic stripes were pronounced, broad and relatively dark reddish brown. In the lab, reproduction by these adults did not oc-cur until after they had been held under warm, long-day conditions with ample food and water for over a week. Moreover, reproduction was correlated with the assumption of bright green coloration, the loss of some of the reddish brown coloration on the thorax, and a narrowing of the prothoracic stripes. In other Chrysopini adults, e.g., species of Chrysoperla, changing behavioral and color patterns like those described for C. (C.) divisus have been shown to be part of the diapause syndrome (Tauber et al. 1986).
In the lab, eggs from all the lots listed above were deposited separately (with isolated stalks), in small groups with no particular pattern; the stalks were sticky, but without droplets. During the first 24 hours after oviposition, the eggs were bright yellowish green to green, without spots. On the second day, they began to develop a bluish brown color, with grey or brownish mottling which became more pronounced as hatching approached (Fig. 9). At 24 ± 1°C, hatching occurred within six to eight days (lots from three females collected in Cruz das Almas, n = 17 -48 eggs/female).
In one case, a small proportion (n = 2 of 28) of the eggs laid by a female from Minas Gerais had a prolonged incubation period (approximately one week to ten days longer than the usual six-to eight-day incubation period). They were a dark bluish brown color during the period of delay. The cause of the prolongation is unknown; however, the resulting larvae developed normally and appeared healthy.
Larvae of C. (C.) divisus carry pieces of woody plant material and other dry debris on their backs. In the rearings (24±1°C) from three females collected at Cruz das Almas, development of the various stages required: L1, 5-8 days; L2, 5-7 days; L3, 5-9 days; cocoon, 16-20 days; complete development from oviposition to adult emergence, 40-48 days. Among the offspring of each of the three females, the sex ratio was approximately 1 : 1 (n = 19-33 individuals / female). The developmental and reproductive responses of C. (C.) divisus to a broad range of temperatures are reported elsewhere (Silva et al. in prep.). Tauber   cies in that their posterior ends are straight and do not curve or connect mesally. C. (C.) fumosus Semaphoront A also differs from those of the other two species in that it has only two thorny cranial setae (S1, S11). And, C. (C.) fumosus Semaphoront B is distinguished from C. (C.) geayi and C. (C.) spinellus in that all of its cranial setae (including S1) are without thorns.
Head (Figs 2B, 10B) 0.39-0.41 mm wide; mandibles 0.36-0.38 mm long (ratio, mandible length : head width = 0.86-0.96 : 1). Cranium mostly white to cream colored, with elongate, narrow, light brown markings. Epicranial marking consisting of two, unconnected, narrow, longitudinally elongated stripes (arms); mesal arm contiguous with postfrontal marking, extending from base of cranium to anteromesal margin of antennal socket; lateral arm lighter brown, more diffuse than mesal arm, extending from posterolateral margin of cranial suture approximately to distal base of mandible. Postfrontal marking indistinguishable, fused with mesal arm of epicranial marking. Frontal marking narrow, extending from midregion of cranium anteriorly to level of antennal socket, then curving laterally toward lighter brown mark at mesal margin of mandibular base; posteromesal ends of marking curving inward, fusing narrowly at tip. Intermandibular, clypeal areas white. Cranial setae amber to light brown; S1, S11 long, thorny, robust; others shorter, smooth; Vx setae small.
Gena, ventral region of head capsule brown, with white spot in anterior region of genal mark. Labial palpus white, marked with light brown on basal segment, distal two annuli of middle segment, base of terminal segment. Mandibles amber to light brown. Antenna with light brown scape, pedicel white basally, light brown distally; flagellum light brown.
Thorax (Figs 2B, 10B) mostly white, with sclerotized structures light brown to brown, small patches of light brown; episternum light brown. Legs white; base of coxa brown; tibia, tarsus tinged with very light brown; tarsal claws, empodia, brown. LS light brown to brown; other setae amber to light brown.
Abdomen (Figs 10C-D) white to cream-colored, with light dusting of brown, especially on A4-A6; sides and venter of these segments also with diffuse brown tinge. LTs, LDTs white with brown chalazae; chalazae of most dorsal setae brown. A1-A5: Dorsum with 12 SMS in two rows, with four mesal pairs equally spaced, two lateral-most pairs juxtaposed; spiracle with SSp elongate, robust, mesal to spiracle; chalazae of all SMS and SSp large, robust. A6: Anterior region with two SMS. Spiracle with very small SSp mesally. A7: Anterior region without setae or microsetae. Spiracle without apparent SSp. A8: Venter with one pair of medium-length setae posteriorly.
Head (Figs 3B, 4B, 11A-B, 12C-D) cream-colored, with dark brown markings. Epicranial marking dark brown, consisting of two narrow, elongate arms, separate from each other, both in contact with posterior margin of head; lateral arm extending from distal ~one-fifth of posterior cranial suture to upper level of eye, not touching eye, becoming broader distally; mesal arm extending from base of head, becoming confluent with postfrontal marking, which extends to inner base of scape. Postfrontal marking very dark brown, narrow throughout. Frontal marking dark brown, with each arm narrow, separate, extending from midsection of head, beyond tentorial pits to inner base of mandibles; darkness extending onto mandibles. Intermandibular marking absent. Clypeolabral region cream-colored tinged with light brown. Gena creamcolored, with large, brown marking from base of eye almost to posterior margin of cranium, with cream-colored mesal spot. Mandible, maxilla amber basally, mesally, becoming dark brown distally. Labial palpus: basal segment cream-colored with very slight tinge of brown; mesal segment ringed with light brown laterally, cream-colored mesally, with terminal subsegment brown; terminal segment dark brown basally, light brown distally. Antenna: scape and pedicel (basal, mesal sections) cream-colored, distal section of pedicel light brown, flagellum brown. Venter cream-colored, except margin of cranium, cardo with longitudinal brown marks; mentum unmarked.
Cervix cream-colored, tinged with light brown dorsally; sides with pair of broad brown patches; venter cream-colored mesally, light brown laterally.
Thorax (Figs 3B, 4B, 11C, 12A-B, 13A) cream-colored, tinged with light brown, with sclerites, markings brown; LTs white to cream-colored, with LS brown to dark  Abdomen (Figs 11D-F, 12A-B, 13B-C) largely cream-colored, tinged with light brown, with light brown to brown markings surrounding bases of LTs; markings becoming darker posteriorly; white fat-body visible beneath midline section (especially posteriorly); robust, thorny setae dark brown; other setae light brown to amber-colored. A5 with pair of light brown spots mesal to LTs. A6, A7 each with pair of large, diffuse, dark brown marks surrounding LDTs. A8 with pair of dark brown marks mesal to spiracles. A9 with U-shaped dorsal mark containing darker longitudinal mark along midline, with pair of lighter brown marks extending onto top of LTs. A10 dark brown distally. Venter white along midsection, cream-colored to light brown laterally; areas mesal to LTs with extensive, diffuse brown marks. Midsection of A10 with pair of small, abutting, triangular dark brown marks.
Larval specimens examined. Several lots, each originating from a single gravid female collected in Brazil, Rio de Janeiro: Conceição de Macabu, Santo Agostinho, V-21-2002, V-2-2003(Albuquerque Lot 2002:012, Tauber Lots 2002:021, 2003; two field-collected L3, Conceição de Macabu, Santo Agostinho, V-2-2003. Biology. Adults and larvae of C. (C.) fumosus were collected on citrus and other fruit trees in mixed agricultural situations. The adults are very agile; they make short, fast, evasive flights, usually downward and toward the interior of the tree. In the lab, eggs were deposited separately (with isolated stalks), in small groups with no particular pattern; the stalks were sticky, but without droplets. The larvae carry pieces of woody plant material and other debris; they are agile, but not particularly fast moving.
Gena brown with small central white spot, ventral region of head capsule white to cream-colored, with ventral margin of head capsule brown. Labial palpus white, with tinge of light brown on basal segment, lateral side of middle segment, darker brown on distal annulation, terminal segment. Mandibles amber to light brown, with dark brown basally. Antenna with light brown scape, pedicel light brown basally, brown distally; flagellum brown. Cervix with pair of light brown sublateral spots.
Thorax (Figs 2C, 14B) mostly white, with small to large patches of brown forming pair of submedian longitudinal, brown bands; sclerotized structures mostly light brown; pleural region with distinct brown band running through episterna, epimera, membranes between; episternum, epimeron dark brown. Legs white; base of coxa dark brown, tibia, tarsus tinged with very light brown; tarsal claws, empodia, brown. LS dark brown; other setae amber to dark brown.
Second and third instars. (Semaphoront B). Body (Figs 15E, 16A-B) length 4.6-5.1 mm (L2), 7.5-7.9 mm (L3); surface white to cream-colored, with light to dark brown integumental spinules especially dense, dark on pronotum; primary pronotal, mesonotal sclerites brown to dark brown, other dorsal marks brown; sclerites around coxae, base of coxae dark brown, abdomen (lateral) with brownish hue interrupted by white lateral stripe through abdominal lateral tubercles. Head (Figs 3C, 4C, 15A-B, 16C-D) cream-colored, with brown to dark brown markings. Epicranial marking brown, with two elongate arms, separate from each other, both in contact with posterior margin of head; lateral arm extending from distolateral margin of cranium to upper level of eye, touching eye, tapering distally; mesal arm extending from base of head, becoming confluent with postfrontal marking, which extends to inner margin of scape. Postfrontal marking dark brown, narrow throughout. Frontal marking dark brown, with each arm narrow, separate (except at basal tip), extending from midsection of head, beyond tentorial pit to inner base of mandible; base of each arm tapering, turning mesally, contacting tip of other arm. Intermandibular marking absent. Clypeolabral region cream-colored, tinged with light brown. Gena cream-colored, with large, brown marking from base of eye to posterior margin of cranium, with small, cream-colored mesal spot. Mandible, maxilla amber basally, mesally, dark brown laterally, distally. Labial palpus: basal segment cream-colored with very  Cephalic seta S1 moderately long, thorny, S2-S12 smooth, S11 long; Vx setae relatively long.
Cervix: dorsum cream-colored, tinged with light brown; sides with pair of broad brown patches; venter brown laterally, becoming cream-colored mesally.
Abdomen (Figs 15D-F, 16A-B, 17B-C) white to cream-colored, with light brown chalazae and setae; white fat-body visible beneath integument; setae mostly ambercolored. A6, A7 each with pair of large, dark brown marks dorsal to LTs; LDTs white with dark brown chalazae, brown marks anterior and posterior to LDTs. A8 with pair of dark brown marks mesal to spiracles. A9 with three elongate brown marks dorsally. A10 with inverted V-shaped, dark brown dorsal mark; light brown laterally. Venter white to cream-colored, without marks, except with some light brown pigmentation ventrolaterally; tip of A10 with pair of small, triangular dark brown marks. pairs of very small setae between spiracles; four pairs of small setae in transverse row between LTs. Venter with two transverse rows of setae, each with three to four smooth, small, pointed setae. A9: Dorsum with one pair of very small setae anteriorly. Middle and posterior regions with two transverse rings of setae extending around segment; each ring with ~14-16 short to medium-length setae, several in each ring robust. A10: Dorsum with two pairs of small setae: one posterior to V-shaped anterior sclerites, one slightly anterior to terminus. Two pairs of lateral setae, robust. Venter with ~five pairs of small setae, posterior row of microsetae anterior to terminus.
Larval specimens examined. Several lots, each originating from a single gravid female collected in Brazil, Rio de Janeiro: Campos dos Goytacazes, Parque Estadual do Desengano, Babilônia III- 27-2001, X-26-2003(Tauber Lot 2001 Biology. Eggs of C. (C.) geayi are deposited separately, with isolated stalks, in no particular pattern. Adults were collected in disturbed, forested areas of coastal Brazil. Occasionally, we encountered fairly substantial numbers in moist habitats near streams. The two larvae (L2, L3) from small trees in the field at Babilônia and Santo Antônio do Imbé (see data above) were both carrying relatively sparse coverings of plant material (brownish leaflets or bracts) and fibers loosely attached to the dorsal setae. On the two specimens from different localities, the leaflets or bracts appeared to be from the same or similar type of plant (Fig. 18).

Chrysopodes (Chrysopodes) lineafrons Adams & Penny, 1987
http://species-id.net/wiki/ Chrysopodes_lineafrons  Figs 2-4, 19-22 Discussion. Chrysopodes (C.) lineafrons occurs widely throughout South America where it frequently has been reported from cropping systems, especially tropical fruit orchards (Adams and Penny 1985, González Olazo et al. 1999, Freitas and Penny 2001, Silva et al. 2007). It is a relatively small-bodied species that can be recognized by its facial markings, parallel and dark gradate veins, and distinctive genitalia (male and female). Adults can be identified using the keys in Adams and Penny (1985) and Freitas and Penny (2001).
Larval diagnosis. The dorsal head markings of Chrysopodes (C.) lineafrons larvae are similar to those of C. (C.) divisus (Figs 2-4). However, C. (C.) lineafrons (Semaphoront A) can be differentiated from C. (C.) divisus by the smaller number of robust, thorny setae on the posterior fold (R1) of the metathorax (n = 4) and the smaller number of smooth, hooked SMS on abdominal segments A1-A5 (n = 16). A dense coating of spinules on the dorsal integument gives the C. (C.) lineafrons larval body (Semaphoront B) a distinctive dark brown hue (most notable on the thorax and abdominal segments A1-A6, including the pleural and ventral regions). Other distinguishing characteristics of C. (C.) lineafrons Semaphoront B are the thorny cranial setae S1, S4, S6, S11, the large, dark brown pronotal plates (Sc1), and the presence of secondary sclerites on the pronotal midline.
Gena, ventral margin of head capsule brown, with small clear patch within genal mark. Labial palpus tinged with light brown, slightly darker distally. Mandibles light brown. Antenna with light brown scape, pedicel white basally, brown distally; flagellum brown.
Abdomen  white to cream-colored, with LTs, LDTs tinged with light brown, chalazae of most dorsal setae brown. A1-A2: Dorsum with four to six SMS in anterior row, with 12 to 16 SMS in posterior row. Spiracles far lateral to anterior row, with SSp mesally. A3-A5: Anterior and posterior rows of SMS largely coalesced, with four to six SMS in anterior row, ten to 14 SMS in curved, posterior row. Spiracle with SSp near anteromesal margin. A6: Anterior region with row of two to six SMS, pair of small, straight setae lateral to SMS. Spiracle with pair of small SSp mesally. A7: Anterior region without setae or microsetae. Spiracle with SSp mesally. A8: Venter with two pairs of medium-length setae posteriorly, one pair of short setae anteriorly.
Head (Figs 3D, 4D, 20A-B, 21C-D) dorsum cream-colored, with brown to dark brown markings. Epicranial marking brown, with mesal and lateral arms in broad contact mesally; both arms in contact with posterior margin of head; lateral arm extending from distolateral margin of cranium to upper level of eye, distal part tapering , broken, almost surrounding dorsal margin of eye; mesal arm extending from base of head, almost confluent with postfrontal marking. Postfrontal marking dark brown, narrow throughout, extending toward inner margin of scape. Frontal marking dark brown, basal arms confluent with each other, forming broad central mark, extending from midsection of head, beyond tentorial pits to intermandibular marking; base of each arm rounded. Intermandibular marking dark brown, broad mesally, tapering laterally, at base of mandibles. Clypeolabral region cream to white. Gena cream-colored, with large, brown, forked marking near base of cranium, with tips of fork reaching approximately 3/4th distance to eye. Mandible, maxilla amber basally, mesally, dark brown laterally, distally. Labial palpus: basal section cream-colored with very slight tinge of brown laterally; mesal segment tinged with brown laterally, cream-colored mesally, with terminal subsegment brown; terminal segment brown. Antenna: scape amber; basal section of flagellum cream-colored with slight tinge of brown; distal 1/4th of pedicel, flagellum, darker brown. Venter amber, with large, white central area; margin of cranium, cardo dark brown; mentum with rectangular, brown spot basally.
Cervix brownish, with dense covering of spinules; sides with pair of broad brown patches; venter brown throughout, darker laterally.
Thorax (Figs 3D, 4D, 20C, 20E, 21A-B, 22A) brown dorsally, laterally, ventrolaterally, tinged by dense covering of brown spinules; sclerites, chalazae dark brown; LTs white, with LS white to light brown; chalazae cream-colored. Venter white to creamcolored mesally, with tinge of brown anteriorly, laterally, where spinules extend to ventral surface, without marks. Pleural region with small brown marks near the base of LTs. Legs: coxa white, with dark brown on dorsal surface; trochanter cream-colored to white, femur brown basally, dark brown distally; tibia white to tinged with very light brown, with light brown setae; tarsus tinged with very light brown; empodium, base brown; claws amber. Episternum with large brown mark; epimeron with small brown mark(s).
Abdomen (Figs 20D-F, 21A-B, 22B-C) dorsum light brown to brown throughout, LTs of A2-A5 with dark brown anterior spots, sometimes posterior spots; LS white to amber, with dark brown chalazae; chalazae of dorsal setae dark brown. A6, A7 each with pair of large, dark brown marks dorsal to LTs, pair ventral to LTs; dark brown marks surrounding anterior lateral margins of LDTs; LDTs white with dark brown chalazae; SMS amber-colored. A8 with pair of dark brown marks mesal to spiracles. A9 with dorsum almost entirely dark brown. A10 with dark brown, inverted V-shaped mark; light brown laterally. Sides of A2-A6 with large, diffuse, brown patch below each LT; venter lightly tinged with brown, light brown mesally, darker brown laterally; A7-A8 mostly white ventrally; A9 light brown ventrally; venter of A10 with pair of small, contiguous, triangular dark brown marks. A1: Dorsum with 40-76 (L2), ~128-150 (L3) SMS in two double-triple transverse bands between spiracles; bands not extending laterally beyond spiracles. A2-A5: Dorsum with 41-102 (L2), 119-173 (L3) SMS in two broad transverse bands. LT with 10-15 (L2), 18-30 (L3) LS: five to seven robust, thorny, blunt LS of various lengths, on distal surface; remaining LS less robust, smooth, hooked in patch on dorsal surface. A6: Dorsum with transverse band of 16-28 (L2), 28-38 (L3) SMS across anterior of segment; midsection with two pairs of smooth setae, mesal pair hooked, long, lateral pair pointed, shorter. LT with 9-11 (L2), 11-17 (L3) robust, thorny, blunt LS of various sizes. A7: Dorsum with two pairs of short setae between spiracles. LT with 8-10 (L2), 10-13 (L3) LS of various sizes. Venter with setal number and size variable, usually with two pairs of setae anteriorly (both small, smooth), three pairs of sublateral setae posteriorly (two lateral-most pairs robust, thorny, mesal pair small, smooth). A8: Dorsum with two pairs of small setae between spiracles; four pairs of small setae in transverse row between LTs. Venter with scattered, small setae anteriorly, two robust, thorny setae on small chalazae between LTs. A9: Dorsum with one pair of very small setae anteriorly. Middle and posterior regions with two transverse rings of setae extending around segment; each ring with ~14-16 setae of various sizes, several in each ring robust. A10: Dorsum with one pair of setae anteriorly, two pairs mesally, patch of several setae distally; one pair of small setae posterior to V-shaped anterior sclerites. Lateral region with two pairs of robust setae, two to three pairs of smaller setae. Venter with ~two pairs of robust setae, five pairs of small setae.
In the lab, eggs were deposited separately (with isolated stalks), in small groups with no particular pattern. During the first 24 hours of oviposition, the eggs were bright green, with dark green blotches. On the second day, they began to develop a bluish brown tone, with brownish mottling; by the third day the eggs were greyish blue to pinkish, with brown mottling. At 24 ± 1°C, hatching occurred within six days (n = 12).
Larvae of C. (C.) lineafrons carry dense packets of woody plant material and other dry debris; they exhibit a side-to-side rocking motion. Development of the various stages (population from the state of Bahia: Cruz das Almas, 24±1°C, n = 14) required: L1, 4-5 days; L2, 3 days; L3, 3-4 days; cocoon, 15 days; complete development from oviposition to adult emergence, 32 days. These data coincide well with the results from extensive rearings of C. (C.) lineafrons from the state of Rio de Janeiro (Campos dos Goytacazes) (see Silva et al. 2007).
Experimental life history studies of C. (C.) lineafrons in the laboratory and the field (southeastern Brazil) indicate: that the species can undergo development and reproduction all-year-round without interruption or dormancy; that during this time up to eight generations can be produced; and that temperature conditions play an important role in determining the rates of reproduction and development both in the lab and in the field (Silva et al. 2007). The species is considered to have excellent potential for mass rearing and for use in the biological control of pests in fruit orchards (Silva et al. 2007).

Chrysopodes (Chrysopodes) spinellus Adams & Penny, 1987
http://species-id.net/wiki/Chrysopodes_spinellus Figs 2-4, 23-26 Discussion. Chrysopodes (C.) spinellus was described from the Amazon region (Adams and Penny 1985); since then, it has not received particular attention. However, we, and others (e.g., Freitas and Penny 2001) have collected it in Brazilian agricultural habitats. We suspect that it is one of the more widespread and common species of Chrysopodes (Chrysopodes) in Brazilian agricultural settings.
Although the female and male genitalia of C. (C.) spinellus are distinctive, both sexes show considerable variation, and the species is not easily distinguished from other Chrysopodes (Chrysopodes) species. The species will be dealt with in an up-coming revision of the subgenus Chrysopodes (C. A. Tauber, in preparation). Meanwhile, the keys and information in Adams and Penny (1985) and Freitas and Penny (2001) are helpful for identification.
Chrysopodes (C.) spinellus was reported from Argentina (with larval description) (Gonzalez Olazo and Heredia 2010); however, the species identification in that report appears to be in error. The larva (L3) that was illustrated had a darkened head like both ) fumosus in that it usually has six thorny cranial setae (S1, S3, S4, S5, S6, S11), and the LS are amber to light brown (not dark brown or black). The C. (C.) spinellus Semaphoront B differs from both C. (C.) geayi and C. (C.) fumosus in having a thorny cranial seta S1, secondary cranial setae between S1 and S4, but no secondary sclerites on the pronotal midline.
Gena, ventral margin of head capsule mostly white, with light brown genal mark posterior to eye, with clear spot near anterior margin. Labial palpus mostly white, middle segment tinged with light brown laterally, distal segment mostly tinged with brown. Mandibles amber to light brown, with dark brown basolateral spot. Antenna with scape light brown, pedicel white basally, brown distally; flagellum light brown.
Thorax (Figs 2E, 23A) mostly white, with sclerotized structures light brown to brown; episternum brown. Legs white, with base of coxa brown, distal one-fourth of tibia, basal one-half of tarsus tinged with brown; tarsal claws, empodia, brown. LS brown; other setae amber to brown. T1: Row of three very small setae (R1) at anteromesal base of LTs. Sc1 with scattered brown areas, especially laterally; S2Sc1 small, immediately above S1Sc1. S1, S3 intermediate-length. S2, S3 thorny. T2: Spiracle with lip of atrium flush with level of integumental surface. Sc1, Sc2 transparent; Sc3 light brown; S2Sc3 medium-length to long, slender, S2 shorter. T3: S1Sc1, S2Sc1 usually present, S1Sc2 very small; S2Sc2 absent. Raised posterior fold with row of four robust, thorny, pointed setae on chalazae with ovate, light brown marks anteriorly. Abdomen (Figs 23C-D) white to cream-colored, with patch of brown on dorsolateral margin of A1; anterior regions of A7, A8 marked with diffuse, scattered brown patches. LTs, LDTs white; chalazae of most dorsal setae brown. A1-A5: Dorsum with 12 SMS in two rows, with four mesal pairs equally spaced, two lateral-most pairs juxtaposed; spiracle with SSp elongate, robust, mesal to spiracle; chalazae of all SMS and SSp not large. A6: Anterior row with two SMS, pair of small, straight setae lateral to SMS; spiracle at anterior base of LT, without apparent SSp. A7: Anterior region without setae or microsetae. Spiracle without apparent SSp. A8: Venter with two pairs of medium-length setae posteriorly.
Head (Figs 3E, 4E, 24A-B, 25C-D) cream-colored, with brown to dark brown markings. Epicranial marking brown, consisting of two elongate arms, separate from each other, both in contact with posterior margin of head; lateral arm extending from distolateral margin of cranium to lower level of eye, becoming narrow distally, extending to upper level of eye; mesal arm extending from base of head, contacting postfrontal marking near base of frontal marking. Postfrontal marking dark brown, robust throughout, extending to inner margin of antennal base. Frontal marking dark brown, with each arm narrow, separate (except at basal tip), extending from midsection of head, beyond tentorial pits to inner base of mandibles; base of each arm tapering, turning mesally, contacting tip of other arm. Intermandibular marking present as light brown connection between distal ends of frontal marking. Clypeolabral region beyond intermandibular marking cream-colored. Gena cream-colored, with large, brown marking from base of eye to posterior margin of cranium, with small, closed, cream-colored mesal spot distally. Mandible, maxilla amber basally, mesally, brown laterally, distally. Labial palpus: basal segment cream-colored with very slight tinge of brown; mesal segment ringed with brown laterally, cream-colored mesally, with terminal subsegment brown; terminal segment brown basally, cream-colored distally. Antenna: scape light brown, basal ~one third of pedicel cream colored, distal two-thirds of pedicel darker brown, flagellum cream-colored with slight tinge of brown. Venter cream-colored, with large, white central area; margin of cranium with light brown longitudinal marks; cardo marked with dark brown; mentum with very light brown spot basally.
Cervix cream-colored, tinged with light brown; sides with pair of broad brown patches; venter brown laterally, becoming cream-colored mesally; with three pairs of small setae ventrally.
Abdomen (Figs 24D-F, 25A-B, 26B-C) dorsum cream-colored to tan, with patches of white fat body visible beneath integument throughout; chalazae of dorsal setae amber to light brown; LTs white, LS cream-colored to amber. A6 with pair of brown marks anterodorsal to LTs; A6, A7 with brown marks anterior to LDTs. A8 with pair of small, light brown marks mesal to spiracles; A9 with dark brown mark mesal to spiracles. A10 with dark brown, inverted U-shaped mark distally; light brownish laterally. Sides of A2-A5 with large, diffuse, very light brown patch below each LT; venter mostly light brown laterally, white mesally; A6-A10 mostly white ventrally; venter of A10 with pair of small, dark brown marks.