An Unusual Chrysopid Larva: Identification, Description, and Taxonomic Implications Author(S): Catherine A

An Unusual Chrysopid Larva: Identification, Description, and Taxonomic Implications Author(s): Catherine A. Tauber and Maurice J. Tauber Source: Annals of the Entomological Society of America, 106(6):729-740. Published By: Entomological Society of America URL: http://www.bioone.org/doi/full/10.1603/AN13105

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BioOne sees sustainable scholarly publishing as an inherently collaborative enterprise connecting authors, nonprofit publishers, academic institutions, research libraries, and research funders in the common goal of maximizing access to critical research. SYSTEMATICS An Unusual Chrysopid Larva: Identiﬁcation, Description, and Taxonomic Implications

1,2,3 1,2 CATHERINE A. TAUBER AND MAURICE J. TAUBER

Ann. Entomol. Soc. Am. 106(6): 729Ð740 (2013); DOI: http://dx.doi.org/10.1603/AN13105 ABSTRACT Berchmansus is a small Neotropical genus in the green lacewing tribe Leucochrysini; its larvae and biology are unknown. Adults of Berchmansus adumbratus Nava´s were found in samples from the Smithsonian National Museum of Natural History Upper Amazonian forest canopy project; these samples came from palms at each of two widely separated Peruvian localities. The same samples also yielded specimens of an unusual leucochrysine larva. For a variety of reasons, we conclude that the larvae are likely conspeciÞc with the adults. If our reasoning proves correct, they would represent the Þrst reported larvae from the genus Berchmansus. Their anatomical features are consistent with either specialized trash-carrying or a naked lifestyle. Here we describe the larvae, and because they have many attributes not previously reported from leucochrysines, we reevaluate the suite of larval features that characterize the tribe. Our analysis illustrates that in both larval morphology and perhaps trash-carrying habits, the tribe Leucochrysini displays a much broader range of variation than previously recorded.

KEY WORDS Neuroptera, Chrysopidae, Leucochrysini, Berchmansus cinctipes, trash-carrying

Currently, the chrysopid tribe Leucochrysini contains lection and preservation, to do so would be unusual seven valid genera (Brooks and Barnard 1990, Tauber for leucochrysine larvae. 2007, Tauber et al. 2008b). Of these, larvae have been 2) Although the larvae display several of the traits that described for three: Leucochrysa (both subgenera), we had considered diagnostic of leucochrysine lar- Gonzaga, and Santocellus (Adams 1987; Tauber 2004; vae, they lack several others. For example, they Mantoanelli et al. 2006, 2011; Tauber et al. 2008a,b, have the elongate, digitiform thoracic tubercles 2011, 2013). Together, these larvae display a large set that are uniquely characteristic of all currently of distinctive features that heretofore we had consid- described leucochrysine larvae. However, they ered as characteristic of the tribe Leucochrysini also have a narrow, slightly ßattened, fusiform (Mantoanelli et al. 2011, Tauber et al. 2011). Larvae body, and all their dorsal setae are without hooks; are unknown for the remaining four leucochrysine both of these features are unknown for leuco- genera (Berchmansus, Cacarulla, Neula, and Nuvol). chrysines. During recent visits to the Smithsonian National 3) The larvae express an attribute that until now has Museum of Natural History (USNM), we sorted not been documented for any extant chrysopid through a large number of insecticidal fogging samples larvaÑa series of elongate lateral tubercles (LTs) (in alcohol) that T. L. Erwin and M. G. Pogue col- on abdominal segments A2 through A7. These tu- lected in the Upper Amazonian forest canopy. Among bercles resemble those on some myrmeleontiform the Neuroptera that we found were adults of Berch- larvae, except that the setae on the leucochrysine mansus adumbratus Nava´s and specimens of an un- larvae are much longer and more ßexible (e.g., usual chrysopid larva. We identiÞed the larvae as sec- Nymphidae: New 1991; extant Ascalaphidae: ond and third instars, belonging to the tribe Henry 1976; fossil Ascalaphidae: MacLeod 1970). Leucochrysini, but we did not recognize the genus or The only other chrysopid larva that is reported to species. have elongate abdominal tubercles is the fossil These larvae are exceptional among leucochrysines Hallucinochrysa (Pe´rezÐde la Fuente et al. 2012), in several respects: but its tubercles are very unlike those on the leucochrysine larva. 1) None had packets of trash on their dorsa. Although it is possible that they lost their trash during col- The presence of such exceptional larval traits raises some intriguing questions: What is the identity of the 1 Department of Entomology, Comstock Hall, Cornell University, larvae? Should they be classiÞed as trash-carriers or as Ithaca, NY 14853-2601. naked? Given the inclusion of these unusual larvae 2 Department of Entomology, University of California, Davis, CA 95616. within the Leucochrysini, how should the list of larval 3 Corresponding author, e-mail: [email protected]. features that currently characterize the tribe (Man-

0013-8746/13/0729Ð0740$04.00/0 ᭧ 2013 Entomological Society of America 730 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA Vol. 106, no. 6 toanelli et al. 2011, Tauber et al. 2011) be revised? scribed (see details later in the text). The remaining Moreover, how does the degree of larval variation possibilities among the leucochrysines are Berchman- among leucochrysine genera now compare with that sus, or one of the rare monotypic leucochrysine gen- among genera in the other tribes of Chrysopinae? era, or a previously undiscovered genus. To help provide a basis for answering these ques- Larval Size. The relatively small size of the second tions: we 1) provide evidence for a probable generic and third instars in our samples corresponds well to and speciÞc identiÞcation of the larvae, 2) describe the Berchmansus adults, which are among the smallest the larvae, 3) reevaluate each of the 12 larval features leucochrysines (for relative forewing sizes: see Brooks that currently characterize the tribe Leucochrysini, and Barnard 1990, Tauber 2007). Cacarulla (also re- and 4) brießy discuss aspects of the larval morphology ported from the Amazonian region of Peru) has a body of this leucochrysine in relation to the biology and that is much too large to correspond to the larvae in phylogeny of the genus. the samples. The adults of the other two monotypic genera are also reported to be large (Nava´s 1916, 1917); we have been unable to conÞrm these reports Materials and Methods because specimens from these genera are lacking. All terminology and methods for the preparation Thus, on the basis of body size, Berchmansus remains and measurement of the larvae are identical to those the closest Þt. we used previously (Mantoanelli et al. 2011, Tauber et Coincidence in Time, Locality, and Habitat of Lar- al. 2011). Ultimately, the specimens will remain in the vae and Adults. Among the samples we sorted, the USNM. larvae in question (n ϭ 11) consistently were associated with adult specimens of B. adumbratus (n ϭ 13). SpeciÞcally, the larval and adult specimens were re- Identification of the Larvae stricted to samples from two widely separated Peru- Normally, we are very hesitant to identify unknown vian localities, one from Loreto in northern Peru, the larvae based primarily on contemporaneously col- other in Madre de Dios in southeastern Peru. At each lected adults (without rearing), but, as outlined later of these localities, the adults and larvae were taken in the text, we conclude that the larvae in the USNM only from palm trees. We did not Þnd similar adults or samples are those of B. adumbratus Nava´s for a series larvae among the many other samples that we sorted of three independent reasons as follows: from nearby or distant localities, or from other types Taxonomic Considerations. In identifying the lar- of plants. Furthermore, at each locality, the larval and vae, we considered all three chrysopid subfamilies. adult specimens coincided in their collection dates. First, the larvae do not express the well-established This complete overlap in time, locality, and habitat distinguishing features of Nothochrysinae (cf DõázÐ constitutes substantial evidence for a close association Aranda et al. 2001, Monserrat and DõázÐAranda 2012). between the larvae and the adults. They also lack the large size and morphological fea- Conclusion From Evidence. Given the diverse tures of the known larvae of Apochrysinae (cf Tsu- sources of evidence mentioned earlier, it is reasonable kaguchi 1995; Fig. 129 in Aspo¨ck and Aspo¨ck 2007). to postulate that the Peruvian larvae are those of B. However, their larval morphology and body size fall adumbratus. We plan to test this hypothesis further by well within the range of variation reported for using molecular data from both adults and larvae. Chrysopinae (cf Tsukaguchi 1995, DõázÐAranda et al. Unfortunately, it has proved difÞcult to obtain a reli- 2001). able molecular signal from our relatively old speci- Three of the four tribes of Chrysopinae are reported mens, which were retrieved from jars containing from the New World: Belonopterygini, Chrysopini, much organic material collected in 1991. If our efforts and Leucochrysini. The larvae in our samples differ continue to be unsuccessful, we will attempt to obtain signiÞcantly from those in the Belonopterygini and fresh well-preserved material from either or both of Chrysopini (cf DõázÐAranda and Monserrat 1995, the Peruvian sites in the future. Tauber et al. 2006). Although they do not express all of In the meantime, the generic and speciÞc identiÞ- the features shared by the larvae known from three cations of the leucochrysine larvae remain uncorrob- genera in Leucochrysini, they do share many distinctly orated. Nevertheless, the extraordinary characteris- unique leucochrysine characteristics. For example, they tics of the larvae and the novel association of both the bear a pair of elongate, digitiform tubercles laterally on adults of B. adumbratus and the unusual larvae with each thoracic segment; these tubercles are longer than palm trees are of considerable interest and well worth one-half the width of the segment and the prothoracic reporting at this time. pair extends to the middle of the head. Larvae in no other chrysopid tribe are known to have such elongated tho- B. adumbratus Navas (tentative) racic tubercles, and the feature is considered a synapo- ´ Second and Third Instars (Semaphoront B) morphy of the tribe. Thus, it is unlikely that our larvae Figs. 1Ð5 represent a previously undescribed tribe, and we are conÞdent that they belong in Leucochrysini. Diagnosis. Among the leucochrysines, these larvae In several respects, the larvae differ markedly from are distinguished by a fusiform body with a pair of those of other leucochrysine genera (Leucochrysa, elongate, digitiform lateral tubercles (LTs) on each of Gonzaga, or Santocellus) whose larvae have been de- abdominal segments A2ÐA7; the absence of hooked November 2013 TAUBER AND TAUBER:UNUSUAL CHRYSOPID LARVA 731

Fig. 1. Berchmansus adumbratus third instar. (A) Head, dorsal. (B). Head, prothorax, dorsal. (C). Head, thorax, dorsal. (D, E). Head, ventral, showing variation in markings. (F). Head, lateral. (G). Body, lateral. (H). Body, dorsal. (All, Madre de Dios, Peru). (Online Þgure in color.) setae, but the presence of slender knobbed setae; and light brown mesal to eye. Postfrontal marking slightly distinctive head markings. Long slender setae extend darker brown than epicranial marking, elongate, ex- from all surfaces of the thoracic and abdominal LTs; tending from midregion of head to base of antenna. most of these setae (except the ventral ones) termi- Frontal marking brown; basal section, an unconnected nate in a small knob. The notal setae and the dorsal pair of small, oblong mesal spots; distal section fused abdominal setae are Þne, knobbed, and relatively short mesally into Y-shape, with distal region of arms fusing (compared with the submedian setae [SMS] of other with intermandibular marking on clypeal region. In- leucochrysine larvae); those on the abdomen are con- termandibular marking extending mesally from inner siderably more numerous than those on the notum. base of mandible, not fusing mesally. Gena without The tips of all the dorsal setae (notal and abdominal) marking; region around eyes and stemmata white. often appear to be acute, but relatively high magniÞ- Labial palpus: basal segment white to cream-colored; cation (Ͼ150ϫ) and oblique lighting show that they mesal segment white basally, with annulations indis- taper and extend distally as extremely Þne Þlaments tinct, distal regions suffused with light brown; terminal and they terminate in a small knob. segment light brown. Mandibles brown basally, me- Body. Dorsum white, with few light brown mark- sally, dark brown at tip. Antenna white to cream- ings; venter white to cream-colored, without mark- colored, with very light brown tinge on lateral sides ings. Dorsal integument with dense covering of of distal annulations. Ventrolateral cranial surface spinules. Length: 5.3Ð5.8 mm (L2); 6.4Ð7.2 mm (L3). cream-colored. Cardo, stipes cream-colored, with All setae smooth. brown marks basally. Mentum white to cream-col- Head. Cranial width: 0.51Ð0.57 mm (L2), 0.77Ð0.84 ored; palpiger, base of maxilla tinged with light brown. mm (L3); cranial length: 0.42Ð0.44 mm (L2), 0.58Ð Cervix cream-colored, unmarked. 0.69 mm (L3). Mandible length: 0.50Ð0.53 mm (L2), Anterior margin of head with slightly rough, obtuse 0.69Ð0.78 mm (L3). Epicranial marking paired, not lateral edges. All setae with acute tips; S1ÐS12 present; contiguous mesally, divided into mesal and lateral sec- S11 longest, S1, S12 longer than others; midregion of tions; mesal section light brown, narrow, short, ex- cranium with patch of ϳ20 small secondary cranial tending anteriorly approximately one-third distance setae. Labial palpus: basal segment with approximately to eye level; lateral arm reduced to very light haze of Þve to six setae, midsegment with distal annulation 732 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA Vol. 106, no. 6

Fig. 2. B. adumbratus third instar (cleared). (A). Head, dorsal. (B). Head, ventral. (C). Prothorax, dorsal. (D). Meso- thorax, dorsal. (E). Metathorax, dorsal. (F). Abdominal segments A1, A2, dorsal. (G). Abdominal segments A3, A4, dorsal. (H). Abdominal segments A5, A6, dorsal. (I). Abdominal segments A7ÐA10, dorsal. Scale bar ϭ 0.5 mm for all images. (Madre de Dios, Peru). (Online Þgure in color.) bearing three long setae, remaining annulations indis- than other setae, S2Sc1 not identiÞed. Mesal pri- tinct, with approximately 6 (L2) or 20 (L3) shorter mary sclerite (Sc2) transparent, oval to triangular, setae, one long basal seta ventrally. Mandible with two ϳone-sixth length of Sc1. LT (L2) with ϳ45Ð48 LS; short dorsal setae; maxillae with two basal setae. Pal- LT (L3) with ϳ80Ð90 LS. Integument mesal to Sc1 piger with two basal setae, one distal seta. Scape with with ϳ10 pairs of medium-length, knobbed setae; approximately Þve setae; terminus of pedicel with regions below, lateral to Sc1 with ϳ12 pairs of me- stout thorn-like projection; ßagellum with long ter- dium-length, knobbed setae; primary setae (S1ÐS5) minal seta. not distinguished. Thorax. LTs white to cream-colored, without T2: Anterior subsegment with transverse row of marks; three to four apical setae on the lateral tuber- ϳ14Ð20 medium-length, knobbed setae between spir- cles (LS) dark brown to black, others white to cream- acles; spiracles with margins sessile, walls of chamber colored; large sclerites (T1Sc1, T2Sc3, and T3Sc2) amber-colored, tapering internally, not bulbous; spi- shiny, very lightly tinged with brown. Spiracles light racular setae, SSp, not distinguished; small anterior amber; setae pale to very light brown. Legs cream- sclerite (Sc1) with three, small associated setae colored, without markings. (S1Sc1, S2Sc1, S3Sc1). Posterior subsegment with LTs digitiform, elongate, with LS extending from all small anterior sclerite (Sc2) bearing two very small surfaces of tubercle (including venter), most dense associated setae mesally (S1Sc2, S2Sc2); Sc3 relatively apically; LS straight to curved throughout, very Þne large, circular, approximate size of prothoracic Sc2, distally, usually tipped with small terminal knob; dark associated seta (S1Sc3) not identiÞed; notum with two apical LS, ventral LS somewhat more robust than broad, transverse rows of medium-length, knobbed lateral or dorsal LS, sometimes acute-tipped. Dorsal, setae; anterior region (anterior to Sc3) with ϳ24 setae lateral surfaces (distal half), ventral surface (entire) (L2), ϳ32 setae (L3); posterior region with ϳ24 setae with numerous microsetae, mostly on ventral surface. (L2, L3). LT bearing ϳ45Ð48 (L2), ϳ70 (L3) LS. Notal setae slender, tapering distally, tips with small to T3: Sclerites (Sc1, Sc2) each with one, small asso- minute knobs. ciated seta (S1Sc1, S1Sc2). Sc3 large, circular, similar T1: First primary sclerite (Sc1) large, elongate, in size to mesothoracic Sc3. Notum with three upward irregularly shaped, extending anterolaterally to base folds bearing medium-length, knobbed setae: anterior of LT, with one seta (possibly S1Sc1) slightly longer one along frontal border of segment, with transverse November 2013 TAUBER AND TAUBER:UNUSUAL CHRYSOPID LARVA 733

Fig. 3. B. adumbratus third instar. (A). Head, dorsal. (B). Head, ventral. Abbreviations: co, cardo; cr, ventral margin of cranium; cx, cervix; epi-l, lateral section of epicranial marking; epi-m, mesal section of epicranial marking; ß, ßagellum; fr, frontal marking; gen, gena; int, intermandibular marking; lp, labial palpus; mb, mandible; mx, maxilla; ped, pedicel; pg, palpiger; post, postfrontal marking; sc, scape; stp, stipes; S1ÐS12, primary cranial setae 1Ð12; Vx, three short posterior setae surrounding a pore. row of ϳ20 setae; middle fold anterior to Sc2, with ϳ48 racular setae, SSp, or the LDS of other chrysopids?). evenly spaced setae; posterior fold behind Sc2, with Dorsum with two rows of ϳ12Ð16 SMS (L2) or two ϳ42 setae. LT (L2) bearing ϳ43 LS; LT (L3) bearing bands ϳ20 SMS (L3). ϳ70 LS. A2ÐA3: L2: LTs with ϳ23 LS on tubercle, approx- Abdomen. LTs on A1ÐA7 digitiform, elongate imately two LS on sclerotized base. Anterior dorsal (length of each approximately one-third width of seg- fold with ϳ15 (A2, A3) SMS; posterior fold with ment); LS straight throughout, tips usually very Þne, ϳ33Ð35 (A2) or ϳ40 (A3) SMS in two bands. L3: LTs with small terminal knobs (acute on ventral LS, oc- with ϳ22 ϩ 8 LS; anterior dorsal fold with ϳ28 (A2) casionally on dorsal, apical LS); apical LS dark brown or ϳ14 (A3) SMS; posterior fold with ϳ34 (A2) or to brown, slightly thicker, more robust than lateral or ϳ44 (A3) SMS in two bands. basal LS. Laterodorsal tubercles (LDTs) absent; each A4ÐA5: LTs with ϳ18 ϩ 2 LS (L2) or ϳ20 ϩ 6LS segment with pair of slightly longer, more robust setae (L3). L2: Anterior dorsal fold with ϳ12 (A4) or ϳ10 (LDS?) in place of LDT. SMS medium-length, pale, (A5) SMS, posterior fold with ϳ40 (A4) or ϳ26 (A5) tapering distally, knobbed, distributed on the surface SMS in two bands. L3: Anterior dorsal fold with ϳ22 in bands as described below. Ventral setae with acute (A4) or ϳ10 (A5) SMS, posterior fold with ϳ40 (A4) tips. Spiracles sessile, unmodiÞed. or ϳ28 (A5) SMS in two bands, each posterior band A1: Lateral margins with ßeshy protuberances, each with one pair of setae longer, more robust than others bearing approximately four (L2) or six (L3) interme- (homologous with LDS of other chrysopids?). diate-length, knobbed setae. Lateral region with long, A6: LTs with ϳ18 ϩ 3 (L2) or ϳ20 ϩ 1 LS (L3). L2: knobbed seta below spiracle (homologous with the spi- Anterior fold with approximately six SMS; posterior 734 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA Vol. 106, no. 6

Fig. 4. B. adumbratus third instar. Three thoracic segments, dorsal. Abbreviations: LS, setae on lateral tubercles; LT, lateral tubercle; Sc1Ð3, primary dorsal thoracic sclerites; sp, spiracle. fold with ϳ24 SMS in two well-separated rows. L3: A10: Dorsum with approximately four short setae, pos- Anterior fold with ϳ10 SMS; posterior fold with ap- terior Þeld of microsetae; terminus membranous with proximately six SMS in each of two well-separated short setae. rows, posterior row with pair of setae longer, more Venter: A2 with two pairs of small setae anteriorly, robust than others, each arising from small, ßeshy one pair laterally. A3ÐA7 each with pair of LDTs bear- protuberances (homologous with LDT, LDS of other ing one relatively long seta, one short seta, three trans- chrysopids?). verse rows of four setae: anterior row with medium- A7: LT with ϳ18 ϩ 1 (L2) or ϳ15 ϩ 6 LS (L3). length setae, middle row with short setae, posterior Dorsum (L2, L3) with three rows each with approx- row with long setae. A8 with approximately four rows imately three to four pairs of SMS; posterior row with of setae, longest in posterior row. A9 with numerous pair of setae much longer, more robust than others, short setae in irregular pattern, longest near posterior arising from small, ßeshy protuberances (homologous margin. with LDT, LDS of other chrysopids?). A8: LT small, rounded protuberance with approximately Þve straight Specimens Examined (Larvae and Adults) LS, with acute tips (one intermediate length, others short). Dorsum with two rows of approximately three PERU: Loreto: one third instar, Cocho Shinguito, short setae each. A9: Dorsum with one pair of interme- 140 m, 05Њ 08Ј S, 74Њ 45Ј W, 27ÐIXÐ91, T.L. Erwin & diate-length setae mesally, approximately six short setae M.G. Pogue et al., Insecticidal fog at TSn 7 of Sheelea anteriorly, approximately seven setae along each side. palm with dry fronds, Lot 54; 1 &, idem, Lot 61; 1 (, November 2013 TAUBER AND TAUBER:UNUSUAL CHRYSOPID LARVA 735

Fig. 5. B. adumbratus third instar. Abdominal segments A1ÐA10, dorsal. Abbreviations: A1ÐA10, Þrst through tenth abdominal segments; LDS?, possible laterodorsal setae; LS, setae on lateral tubercles; LT, lateral tubercle; prot, ßeshy, lateral protuberance; SMS, submedian seta; sp, spiracle.

1 &, two third instars, idem, Lot 58, USNM. Madre de Features Retained on the List of Leucochrysine Dios: one third instar, Rio Manu, BIOLAT Biol. Sta. Larval Characteristics Pakitza, 356 m, 11Њ56Ј47Љ S, 71Њ17Ј00Љ W, 10ÐXÐ91, T.L. Erwin & M.G. Pogue, Insecticidal fog of Astrocaryum 1. **Thoracic LTs long: prothoracic LTs extending at dry leaves (nine fronds) 4Ð5 m height, Tr. Can˜ a least to the middle of the head, mesothoracic and Brava/7, Lot 215; 2 (,1&, two second instars, two metathoracic LTs longer than half the width of the third instars, idem, 10ÐXÐ91, Insecticidal fog of Jesse- corresponding segment (all instars). This appears nia dry leaves (12 fronds) 4 m height, Tr. Can˜ a Brava/ to be the most distinguishing synapomorphy of leu- 7.5, Lot 232; 1 (, idem, 10ÐXÐ91, Insecticidal fog of cochrysine larvae. It is expressed by all known leu- Jessenia dry leaves (16 fronds) 3Ð5 m height, Tr. Can˜ a cochrysine larvae (including the second and third Brava/7.5, Lot 237; 1 &, idem, 14ÐXÐ91, Insecticidal instars described in this study), and it is not known fog of Astrocaryum (13 fronds to 4 m) Tr. Pasal/24Ð25, to occur in larvae from any other chrysopid tribe Lot 267; one third instar, idem, Insecticidal fog of (for Ankylopterygini: see Tsukaguchi 1995; for Be- Astrocaryum (Þve fronds to 3.5 m) Tr. Pasal/24Ð25, lonopterygini: see Principi 1946, Monserrat and Lot 270; 2 (,1& with parasitoid, one third instar, DõázÐAranda 2012; for Chrysopini: see, Tauber idem, 14ÐXÐ91, Insecticidal fog of Astrocaryum (Þve 1974, 1975, 2003; DõázÐAranda and Monserrat 1995; fronds to 3.5 m) Tr. Pasal/24Ð25, Lot 273; 2 (, one Tsukaguchi 1995; Penny et al. 2000; Tauber et al. third instar, idem, 14ÐXÐ91, Insecticidal fog of Astro- 2000, 2001; Tauber and de Leon 2001; Monserrat caryum (seven fronds to 4 m) Tr. Pasal/24Ð25, Lot 274, and DõázÐAranda 2012; Silva et al. 2013). USNM. 2. **Terminal seta of the antenna at least half the length of the ßagellum (all instars). In the leucochrysine specimens dealt with here, the terminal Larval Characteristics of Leucochrysini—Reevaluated seta of the antenna ranges from 0.65 to 0.73ϫ the Given the unusual leucochrysine larva described length of the ßagellum. These Þgures fall well earlier, we reevaluate the 12 larval features previously within the range of other leucochrysines, and at this used in characterizing the tribe (for the most recent time, the characteristic remains typical of leuco- discussion, see Mantoanelli et al. 2011). Two of the 12 chrysine larvae. It is not known to occur in chryso- (#1 and #2) continue to be useful in assigning tribal pine or belonopterygine larvae (Principi 1946, afÞliation; they are marked with two asterisks (**). A DõázÐAranda and Monserrat 1995, Tsukaguchi third trait is retained until we can examine the Þrst 1995, Tauber et al. 2000, Tauber 2003, Tauber et al. instar, and a fourth is retained with modiÞcation. The 2006, Monserrat and DõázÐAranda 2012). latter two are marked with a single asterisk (*). The 3. *Prothorax with primary setae S1, S3, S4, and S5 other eight traits are removed from the list, and we present, S2 absent (all instars). This character state discuss their value as generic or species-level charac- is expressed by the larvae of all previously de- teristics. scribed leucochrysine larvae except the L2 and L3 736 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA Vol. 106, no. 6

of two species (Mantoanelli et al. 2006, 2011; the leucochrysine genera (Leucochrysa, Gonzaga, Tauber et al. 2008a,b, 2011, 2013); we have not seen Santocellus) also express this feature (Tauber et al. it noted for any other chrysopid tribe. In the second 2008a,b, 2011, 2013; Mantoanelli et al. 2011); the and third instars described here, the large number larva described here represents the Þrst leuco- of secondary setae on the prothorax, which are chrysine known to have a ßattened, fusiform larval indistinguishable from primary setae, makes this body. Thus, Leucochrysini joins the Chrysopini in character difÞcult to interpret. Thus, we retain the expressing generic-level variation in body shape, character, with a caveat that we await the discovery and the attribute is removed from the list of leu- of a Berchmansus Þrst instar for evaluation. cochrysine characteristics. However, the feature 4. *Abdominal segments A1ÐA5 each with a pair of remains very useful at the generic level in both LDTs, each LDT with one long, smooth, hooked Leucochrysini and Chrysopini. seta (LDS) and one long or short LDS. This char- 3. Abdominal segments A7ÐA10 small, curved ven- acter state is expressed by all previously described trally, and partially telescoped into the anterior leucochrysines; in contrast, the leucochrysine sec- segment (all instars). The terminal segments of ond and third instars described here do not have neither chrysopine nor ankylopterygine trash-car- distinct LDTs. However, each abdominal segment riers are curved and withdrawn, but they are in the (A1ÐA7) has one pair of knobbed setae that are belonopterygine Italochrysa (Principi 1946) and in longer and more robust than the other dorsal setae. the previously described trash-carrying leuco- These setae occur posteriorly and in the position of chrysine generaÑLeucochrysa, Gonzaga, and San- the LDTs; on segments A5ÐA7, they arise from tocellus (Tauber et al. 2008a,b, 2011, 2013; Man- small, ßeshy protuberances. It is possible that these toanelli et al. 2011). The fusiform leucochrysine protuberances and knobbed setae are homologous described here does not express the condition. to the LDTs and hooked LDS of other leuco- Thus, like the earlier feature, this characteristic is chrysines, and that the character remains useful at useful for differentiating leucochrysine genera, but the tribal level, if it were modiÞed as follows: not the tribe. Abdominal segments A1ÐA5 each with a pair of LDTs, 4. LTs on abdominal segments A2 and A3 papilliform, each LDT with one long, smooth, hooked seta (LDS) with long setae extending from the anterior, dorsal and one long or short LDS, or with LDT absent or and apical surfaces (L2, L3). This characteristic greatly reduced, but with one long, knobbed seta holds true for the larvae in the previously described (LDS?) in its place. leucochrysine genera. However, the digitiform abdominal tubercles of the leucochrysine described here provide a notable exception. We now consider Features Removed From the List of Leucochrysine abdominal tubercle length as a useful generic-level Larval Characteristics character within the Leucochrysini. 1. Thoracic tubercles with long, setae (LS), extending 5. LTs on abdominal segments A4ÐA7 spherical to in a fan-shape mainly from the apical and lateral papilliform, bearing long setae mainly on the apical surfaces (L2, L3). In the second and third instars surface (L2, L3). This feature characterizes the described here, like those of all previously de- larvae in all previously described leucochrysine scribed leucochrysine larvae, the thoracic LS are genera. Again, the larvae described here provide a long. Thus, this feature remains a consistent char- notable exception, and we now consider the feature acteristic of the leucochrysine larvae. However, to be useful for distinguishing leucochrysine gen- some genera (e.g., Ankylopteryx, Ceraeochrysa, era. Chrysopodes, Semachrysa) in other chrysopid tribes 6. Mesonotum and metanotum each with two trans- also have long LS that extend from the LTs in a verse rows of long, smooth, hooked (rarely similar pattern (Tsukaguchi 1995, Tauber et al. straight) setae arising from chalazae. This feature 2000, Silva et al. 2013). Thus, we remove this char- previously was shown to have two exceptions acteristic from the list of diagnostic leucochrysine within Leucochrysa (Leucochrysa); in these species, features with the note that all known leucochrysine the L1 lacks hooked setae on the thorax, and the L2 larvae express the trait. and L3 lack hooked setae in the anterior row of the 2. Anterior segments of the abdomen thicker than mesothorax and metathorax (Tauber et al. 2013). In those of the thorax, giving the larva a humpback the leucochrysine here, the mesonotal and met- appearance in lateral view (all instars). A thickened anotal setae are knobbed and distributed broadly body is typical of trash-carrying larvae in Belonop- across the dorsal surface. Because of these excep- terygini (Principi 1946) and Chrysopini (Dõ´azÐ tions, we now consider this character to be useful Aranda and Monserrat 1995, Tsukaguchi 1995, in differentiating leucochrysine genera and/or spe- Monserrat and Dõ´azÐAranda 2012; for neotropical cies, rather than as a diagnostic feature of the tribe. genera, see Tauber et al. 2000, 2012; Tauber 2003; 7. Abdominal segment A1 with a single row of smooth, Silva et al. 2013). The few known larvae of Anky- hooked SMS, between the LDTs (all instars; L1: lopterygini (from three genera) are typical trash- four setae in row; L2, L3: number of setae variable). carriers (Ho¨lzel et al. 1990, Tsukaguchi 1995, Unfortunately, the samples we sorted did not yield Aspo¨ck and Aspo¨ck 2007) and probably have thick- Þrst instars; but the second and third instars provide ened abdomens. All previously described larvae in an interesting situation. First, the LDTs are absent November 2013 TAUBER AND TAUBER:UNUSUAL CHRYSOPID LARVA 737

from A1 or they are greatly reduced. Moreover, the chrysine larva may not preclude the trash-carrying smooth and knobbed (not hooked) SMS are dis- habit. tributed across the dorsal surface of A1 in two broad Specialization. All of the adult B. adumbratus and bands (L3) or two rows (L2). Thus, this feature larval specimens that we retrieved from the samples remains typical of Leucochrysa, Gonzaga, and San- had been collected from the fronds of three genera of tocellus, but not all leucochrysines. It is noteworthy palms (Astrocaryum and Jessenia in Madre de Dios and that the margins of the knobs on the SMS and LS Sheelea in Loreto); notes accompanying the speci- appear well-sclerotized and slightly expanded; they mens indicate that dry fronds were present on at least may represent a hook, the center of which has been some of the palms trees that were fogged, and that the Þlled. However, the distal region of the shank is height of the palms extended to 5 m. From these data, extremely tapered and slender, unlike that on the we deduce that the larvae may have a speciÞc habitat more robust, hooked SMS of other leucochrysines. association with palms and/or a restricted prey associa- 8. Abdominal segments A2ÐA5 with two (L1) or three tion with an organism, or organisms, living on palms. (L3, L2) rows of smooth, hooked SMS, the poste- Specialized habitat and prey associations are known rior row of which lies between the LDTs (L1 with to occur in leucochrysines. For example, Leucochrysa four setae in anterior row, two in posterior row, all (Nodita) pavida (Hagen) inhabits tree trunks in east- between LDTs; L2, L3 with number of setae vari- ern United States and is known to consistently carry able). Here again, although we found no Þrst instars trunk-inhabiting lichens (Skorepa and Sharp 1971, in our samples, the second and third instars deviate Wilson and Methven 1997). Similarly, Leucochrysa from the previously proposed leucochrysine pat- (L.) insularis (Walker) larvae have only been re- tern. First, the SMS are knobbed, not hooked. ported to carry snails and/or snail shells in their trash Moreover, they are distributed across each segment packets; they also feed on small snails (Jones 1929). in three broad bandsÑone band on the anterior Many genera of palms, including those that yielded the fold and two bands across the posterior fold. This leucochrysine larvae described here, have hairs and pattern roughly resembles that of some other leu- other small structures on the lamina and/or leaf axis cochrysines (and chrysopines), but homology is (Tomlinson 1961). Perhaps, the leucochrysine larvae not clear. carry trash packets containing plant defensive structures for their own protection or camoußage, as do the trichome-carrying Ceraeochrysa lineaticornis (Fitch) Biological Notes and Hallucinochrysa (Eisner et al. 2002, Pe´rezÐde la Fuente et al. 2012). Trash-Carrying. On the basis of Þeld and laboratory Parasitization. One of the adult specimens in our observations, as well as morphological features, leu- samples (a female) contained an internal hymenop- cochrysine larvae have been classiÞed as trash-carriers teran parasitoid larva. The parasitoidÕs body occu- (as opposed to naked larvae) (Jones 1929; Adams 1987; pied most of the abdominal cavity of the host, and Tauber 2004; Mantoanelli et al. 2006, 2011; Mantoanelli it compressed the host gut against the ventral wall and Albuquerque 2007; Tauber et al. 2008a,b, 2011, of the abdomen (Fig. 6). The parasitoid appears 2013). In addition, the trash packets of most leuco- similar to the braconid parasitoid (Chrysopophtho- chrysines that we and others have observed are highly rus chrysopimaginis Goidanich) reported from the consolidated, not easily dislodged, and, in some cases, adults of a range of chrysopid species in Italy and interwoven with silk threads (Slocum and Lawrey France (Principi 1948, 1956; Principi et al. 1979; New 1976). However, the leucochrysines described here 1984). The parasitoid specimen now is preserved in did not have trash attached to their dorsal tubercles the abdomen of its host (in a genitalia vial inserted into when we sorted them. Although it is possible that any the larger vial containing the lacewing; USNM, Lot trash they carried was dislodged during collection and 273, cited earlier). preservation, we cannot conÞrm that the larvae should be classiÞed as trash-carriers. Given their structure and the absence of associated trash, we suggest that Larval Morphology, Trash-carrying, and either they are naked larvae or they carry a loosely Chrysopid Systematics constructed and/or loosely attached packet of spe- During the past several decades, larvae within the ciÞc, small debris. large tribes Chrysopini and Leucochrysini have re- It is noteworthy that our leucochrysine is similar in ceived considerable attention, and distinct patterns of general body form to an Early Cretaceous trash-car- generic variation in larval morphology and habits are rying chrysopid, Hallucinochrysa, recently described becoming apparent. The following developments led by Pe´rezÐde la Fuente et al. (2012). Although the us to conclude that larval morphology can and will larvae of the two species differ in many respects, both play a greater role in the systematics of the groups than have a slightly ßattened, fusiform body shape, and it has in the past. long, narrow thoracic and abdominal tubercles. The Of the two tribes, the Chrysopini includes many Hallucinochrysa specimen was embedded in amber more genera, and it also has the broader range of with a loosely constructed packet of fern trichomes, intergeneric variation in larval structure and known and therefore it was considered a trash-carrier. Thus, habits. For example, the tribe is documented to have the morphological features of our extant leuco- naked, fusiform larvae, trash-carrying, globose larvae, 738 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA Vol. 106, no. 6

Fig. 6. Mature internal parasitoid within B. adumbratus female abdomen (Madre de Dios, Peru). Abbreviations: body, body cavity of parasitoid; d.i., torn dorsal integument of lacewing abdomen; g.l., gonapophysis lateralis; gut, lacewing gut displaced to venter of body cavity; head, head of parasitoid; sp, lacewing spermatheca; S3, S7, lacewing third, seventh sternites. (Online Þgure in color.) as well as intermediate forms. Each of these larval been appreciated to date. Now, although we reiterate forms is associated with a large and diverse array of our earlier call for caution in selecting larval charac- anatomical structures, and also distinctive ecological ters for use in chrysopid phylogenetic studies (Tauber and behavioral traits. Importantly, these characteris- 1975), in doing so, we emphasize that the larval mor- tics vary markedly and distinctively among chrysopine phology and behavior related to trash-carrying offer a genera [for summaries, see: DõázÐAranda and Mon- rich source of valuable characters from which to serrat 1995, Monserrat and DõázÐAranda 2012 (Euro- choose. pean Chrysopini); Tsukaguchi 1995 (Japanese fauna); Tauber 1974, 1975, 2003; Penny et al. 2000; Tauber et al. 2000, 2001, 2012; Tauber and de Leon 2001; Silva et Acknowledgments al. 2013 (New World taxa)]. In contrast to the Chrysopini, and until now, it was We are pleased to thank O. S. Flint, T. L. Erwin, D. G. Furth, and W. E. Steiner (USNM) for facilitating our work at thought that the larvae of all leucochrysine genera the Museum and for providing access to the fogging samples were trash-carriers and that they exhibited a relatively and Steven A. Nadler (UC-Davis) for his cooperation. Our limited range of variation in larval morphology (Man- research received support from the National Science Foun- toanelli et al. 2011, Tauber et al. 2011; see earlier). dation (grants INT-9817231, DEB-0542373), the National Given the unusual leucochrysine larva described here, Geographic Society, the United States Department of Agri- it now appears that the tribe Leucochrysini also ex- culture/National Research Initiative (Competitive grant presses a broad range of variation in larval anatomical 9802447), Regional Project W-3185, and Cornell University. structures (and perhaps lifestyle)Ñmuch more than previously recognized. However, despite this back- ground of diversity, a suite of distinctive leucochrysine References Cited characteristics are evident. Adams, P. A. 1987. Studies in Neotropical Chrysopidae In our view, the large diversity and stable patterns (Neuroptera) III. Notes on Nodita amazonica Nava´s and. of morphological variation currently documented in N. oenops, n. sp. Neuroptera Int. 4: 287Ð294. extant and fossil chrysopid larvae illustrate two Aspo¨ck, U., and H. Aspo¨ck. 2007. Verbliebene Veilfalt ver- pointsÑone old, one new: 1) As stated earlier (Tauber gangener Blu¨ te. Zur Evolution, Phylogenie und Biodi- 1975, Monserrat and DõázÐAranda 2012, Pe´rezÐde la versita¨t der Neuropterida (Insecta: Endopterygota). De- Fuente et al. 2012), trash-carrying may have arisen and nisia 20: 451Ð516. Brooks, S. J., and P. C. Barnard. 1990. The green lacewings been lost numerous times in the Chrysopidae, and thus of the world: a generic review (Neuroptera: Chrysopi- the presence/absence of “trash-carrying,” and some of dae). Bull. Br. Mus. Nat. Hist. Entomol. 59: 117Ð286. the morphological adaptations associated with it, are Dıáz–Aranda, L. M., and V. J. Monserrat. 1995. Aphidopha- likely to express homoplasy. 2) However, we now gous predator diagnosis: key to genera of European know that trash-carrying (and/or the absence of it) is chrysopid larvae (Neur.: Chrysopidae). Entomophaga 40: based on large and diverse sets of distinct morpho- 169Ð181. logical features, the distribution of which is very stable Dıáz–Aranda, L. M., V. J. Monserrat, and C. A. Tauber. 2001. and systematically revealing at all taxonomic levelsÑ Recognition of larval Neuroptera. Recognition of early from species through subfamily (for some examples, stages of Chrysopidae, pp. 60Ð81. In P. K. McEwen, T. R. New, and A. E. Whittington (eds.), Lacewings in the see: DõázÐAranda and Monserrat 1995, DõázÐAranda et Crop Environment, p. 546. Cambridge University Press, al. 2001, Tauber et al. 2006, Monserrat and DõázÐ Cambridge, United Kingdom. Aranda 2012). Given this situation, it appears that Eisner, T., J. E. Carrel, E. van Tassell, E. R. Hoebeke, and M. comparative larval morphology constitutes a much Eisner. 2002. Construction of a defensive trash packet richer source of phylogenetic information than has from Sycamore leaf trichomes by a chrysopid larva (Neu- November 2013 TAUBER AND TAUBER:UNUSUAL CHRYSOPID LARVA 739

roptera: Chrysopidae). Proc. Entomol. Soc. Wash. 104: Leach (C. flavifrons Brauer, prasina Burm. e clathrata 437Ð446. Schn.). Bollettino dellÕIstituto di Entomologia della Uni- Henry, C. S. 1976. Some aspects of the external morphology versita` degli Studi di Bologna. 21: 319Ð410. of larval owlßies (Neuroptera: Ascalaphidae), with par- Principi, M. M., M. Memmi, and D. Sgobba. 1979. Reperti su ticular reference to Ululodes and Ascaloptynx. Psyche 83: Chrysopophthorus chrysopimaginis Goidanich, parassita 1Ð31. solitario delle immagini di neurotteri crisopidi. (Con- Ho¨lzel, H., M. Stelzl, and P. Ohm. 1990. Chrysopidae (Neu- tributi allo studio dei Neurotteri italiani, XXIII). Bollet- roptera) aus Senegal und Gambia. I: Ankylopterygini. tino dellÕIstituto di Entomologia della Universita` degli Neuroptera Int. 6: 63Ð73. Studi di Bologna 34: 247Ð273. Jones, D. T. 1929. A snail-collecting aphis-lion larva. A pre- Silva, P. S., C. A. Tauber, G. S. Albuquerque, and M. J. liminary study of external features. Marietta College Res. Tauber. 2013. Larvae of Þve horticulturally important spe- Publ. (Zool.). 1: 1Ð9. cies of Chrysopodes (Neuroptera, Chrysopidae): shared ge- MacLeod, E. G. 1970. The Neuroptera of the Baltic Amber. neric features, descriptions and keys. ZooKeys 262: 39Ð92. I. Ascalaphidae, Nymphidae, and Psychopsidae. Psyche (http://www.pensoft.net/J_FILES/1/articles/4119/ 77: 147Ð180. 4119-G-3-layout.pdf). Mantoanelli, E., and G. S. Albuquerque. 2007. Desenvolvi- Slocum, R. D., and J. D. Lawrey. 1976. Viability of the epiz- mento e comportamento larval de Leucochrysa (L.) varia oic lichen ßora carried and dispersed by green lacewing (Schneider) (Neuroptera, Chrysopidae) em laborato´rio. (Nodita pavida) larvae. Can. J. Bot. 54: 1827Ð1831. Revista Brasileira de Zoologia 24: 302Ð311. Skorepa, A. C., and A. J. Sharp. 1971. Lichens in “packets” of Mantoanelli E., G. S. Albuquerque, C. A. Tauber, and M. J. lacewing larvae (Chrysopidae). Bryologist 74: 363Ð364. Tauber. 2006. Leucochrysa (Leucochrysa) varia (Sch- Tauber, C. A. 1974. Systematics of North American neider) (Neuroptera: Chrysopidae): larval descriptions, de- chrysopid larvae: Chrysopa carnea group (Neuroptera). velopmental rates, and adult color variation. Ann. Entomol. Can. Entomol. 106: 1133Ð1153. Soc. Am. 98: 7Ð18. (http://esa.publisher.ingentaconnect. Tauber, C. A. 1975. Larval characteristics and taxonomic com/content/esa/aesa/2006/00000099/00000001/art00003). position of the lacewing genus Suarius. Ann. Entomol. Mantoanelli E., C. A. Tauber, G. S. Albuquerque, and M. J. Soc. Am. 68: 695Ð700. Tauber. 2011. Larvae of four Leucochrysa (Nodita) spe- Tauber, C. A. 2003. Generic characteristics of Chrysopodes cies (Neuroptera: Chrysopidae: Leucochrysini) from (Neuroptera: Chrysopidae), with new larval descriptions BrazilÕs Atlantic coast. Ann. Entomol. Soc. Am. 104: 1233Ð and a review of species from the United States and Can- 1259. (http://esa.publisher.ingentaconnect.com/content/ ada. Ann. Entomol. Soc. Am. 96: 472Ð490. (http://esa. esa/aesa/2011/00000104/00000006/art00014). publisher.ingentaconnect.com/content/esa/aesa/2003/ Monserrat, V. J., and L. M. Dıáz–Aranda. 2012. Los estadios 00000096/00000004/art00008). larvarios de los Criso´pidos ibe´ricos (Insecta, Neuroptera, Tauber, C. A. 2004. A systematic review of the genus Leu- Chrysopidae), nuevos elementos sobre la morfologõá lar- cochrysa (Neuroptera: Chrysopidae) in the United States. varia aplicables a la sistema´tica de la familia. Graellsia 68: Ann. Entomol. Soc. Am. 97: 1129Ð1158. 31Ð158. Tauber, C. A. 2007. Review of Berchmansus and Vieira and Nava´s, L. 1916. Neuro´pteros Sudamericanos. Tercera [III] description of two new species of Leucochrysa (Neurop- serie. Neuro´pteros del Brasil recogidos por el R. P. tera: Chrysopidae). Ann. Entomol. Soc. Am. 100: 110Ð138. Joaquõń da Silva Tavares S. J. Brote´ria (Zoolo´gica) 14: (http://esa.publisher.ingentaconnect.com/content/esa/ 14Ð35. aesa/2007/00000100/00000002/art00003). Nava´s, L. 1917. Neue Neuropteren. Dritte [III] serie. En- Tauber, C. A., and T. de Leoń. 2001. Systematics of green tomologische Mitteilungen 6: 274Ð282. lacewings (Neuroptera: Chrysopidae): larvae of Ceraeo- New, T. R. 1984. IdentiÞcation of hymenopterous parasites chrysa from Mexico. Ann. Entomol. Soc. Am. 94: 197Ð209. of Chrysopidae, pp. 193Ð204. In M. Canard, Y. Se´me´ria, Tauber, C. A., T. de Leoń, N. D. Penny, and M. J. Tauber. and T. R. New (eds.), Biology of Chrysopidae. Dr. W. 2000. The genus Ceraeochrysa (Neuroptera: Chrysopidae) Junk Publishers, The Hague, The Netherlands. of America north of Mexico: larvae, adults, and comparative New, T. R. 1991. Neuroptera (lacewings), pp. 525Ð542. In biology. Ann. Entomol. Soc. Am. 93: 1195Ð1221. I. D. Naumann (ed.), The Insects of Australia, 2nd ed., vol. Tauber, C. A., M. J. Tauber, and G. S. Albuquerque. 2001. Ple- 1. Melbourne University Press, Melbourne, Australia. siochrysa brasiliensis (Schneider) (Neuroptera: Chrysopi- Penny, N. D., C. A. Tauber, and T. de Leoń. 2000. A new dae): larval stages, biology, and taxonomic relationships. Ann. species of Chrysopa from western North America with a Entomol. Soc. Am. 94: 858Ð865. key to North American species (Neuroptera: Chrysopi- Tauber, C. A., M. J. Tauber, and G. S. Albuquerque. 2006 dae). Ann. Entomol. Soc. Am. 93: 776Ð784. Berchmansus elegans (Neuroptera: Chrysopidae): Larval Pe´rez–de la Fuente, R., X. Delclo`s, E. Pen˜ alver, M. Speranza, and adult characteristics and new tribal afÞliation. Eur. J. J. Wierzchos, C. Ascaso, and M. Engel. 2012. Early evo- Entomol. 103: 221Ð231. lution and ecology of camoußage in insects. Proc. Natl. Tauber, C. A., G. S. Albuquerque, and M. J. Tauber. 2008a Acad. Sci. U.S.A. 109: 21414Ð21419. Gonzaga nigriceps (McLachlan) (Neuroptera: Chrysopi- Principi, M. M. 1946. Contributi allo studio dei Neurotteri dae): descriptions of larvae and adults, biological notes, Italiani. IV. Nothochrysa italica Rossi. Bollettino dellÕIstituto and generic afÞliation. Proc. Entomol. Soc. Wash. 110: di Entomologia della Universita` degli Studi di Bologna 15: 417Ð438. 85Ð102. Tauber, C. A., M. J. Tauber, and G. S. Albuquerque. 2008b. Principi, M. M. 1948. Contributi allo studio dei Neurotteri A new genus and species of green lacewings from Brazil Italiani. VII. Osservazioni su alcuni parassiti di crisopidi. (Neuroptera: Chrysopidae: Leucochrysini). Ann. En- Bollettino dellÕIstituto di Entomologia della Universita` tomol. Soc. Am. 101: 314Ð326. (http://esa.publisher. degli Studi di Bologna 17: 93Ð121. ingentaconnect.com/content/esa/aesa/2008/00000101/ Principi, M. M. 1956. Contributi allo studio dei Neurotteri 00000002/art00006). Italiani. XIII. Studio morfologico, etologico e sistematico Tauber, C. A., E. Mantoanelli, G. S. Albuquerque, C. Reguiloń, di un gruppo omogeneo di specie del Gen. Chrysopa E. Gonza´lez Olazo, and M. J. Tauber. 2011. A taxonomi- 740 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA Vol. 106, no. 6

cally signiÞcant polymorphism in Leucochrysa (Neuroptera: Tomlinson, P. B. 1961. Anatomy of the Monocotyledons, II Chrysopidae): nomenclature, larval and adult descriptions, Palmae. Oxford, Clarendon Press, Oxford, United King- and biological notes. Zootaxa 3130: 1Ð29. (http://www. dom, 453 pp. mapress.com/zootaxa/2011/f/zt03130p029.pdf). Tsukaguchi, S. 1995. Chrysopidae of Japan (Insecta, Neu- Tauber, C. A., G. S. Albuquerque, and M. J. Tauber. 2012. roptera). S. Tsukaguchi, Aioi-cho 6Ð14-102, Nishinomiya- The neotropical genus Titanochrysa (Neuroptera, shi, Hyogo, Japan, 223 pp. Chrysopidae): larval descriptions, biological notes, a new Wilson, P. J., and A. S. Methven. 1997. Lichen use by larval species, and taxonomic changes. Zootaxa 3514: 1Ð26. Leucochrysa pavida (Neuroptera: Chrysopidae). Bryolo- Tauber, C. A., F. Sosa, G. S. Albuquerque, and M. J. Tauber. gist 100: 448Ð453. 2013. Adults and larvae of two Leucochrysa (Leuco- chrysa) species (Neuroptera: Chrysopidae) descriptions, biological notes, and relationships. Zootaxa (in press). Received 30 June 2013; accepted 26 August 2013.