Ceraeochrysa, a New Genus of Chrysopinae (Neuroptera) (Studies in New World Chrysopidae, Part Ii)

NEUROPTERA INTERNATIONAL 11 (2) - 1982. p. 69-75

CERAEOCHRYSA, A NEW GENUS OF CHRYSOPINAE (NEUROPTERA) (STUDIES IN NEW WORLD CHRYSOPIDAE, PART II)

Phillip A. ADAMS Department of Biol'ogical Science California State University, Fullerton Fullerton, California 92634

Summary

Ceraeochrysa (type species Chrysopa cincta Schneider) is distinpuished by absence of tignum, presence of horn-like projections on the gonarcus, mediuncus with apical hook often flanked by projections, longitudinal gonapsis, elongate spermatheca, lack of microtholi or micropoculae. To this neotropical genus 24 species are referred, and 51 new synonymies are reported; a neotype is designated for C. arioles Banks, and a lectotype for C. claveri Navas.

Résumé

Ceraeochrysa (espèce type : Chrysopa cincta Schneider) se distingue par l'absence de tignum, par la présence de saillies en forme de corne sur le gonarcus, par un mediuncus avec un crochet apical souvent encadré de saillies, un gonapsis longitudinal, une spermathèque allongée et par une absence complète de microtholi ou de micropoculae. Sont ici données les 24 espèces relevant de ce genre néotropical, ainsi Que 51 nou- velles synonymies. Un neotype est proposé pour C. arioles Banks, et un lectotype pour C. claveri Navas.

This is the second of a series of papers intended to define genera of New World Chrysopinae, assign species to genera, and to reduce the excessive accumulation of synonyms to a workable level. PENNY(1977) listed 237 species in the genus (( Chrysopa », for the New World south of the United States, with no attempt at subdivision of the genus. BICKLEYand MACLEOD(1956) arranged the 47 named species of North American Chrysopa into species groups on the basis of the then-current colorational characters, recognizing only one subgenus as distinct from Chrgsopa S. str., based upon a single venational character. TJEDER'S(1966) erection of several subgenera, based upon genitalic characters, has been generally accepted, but there has been reluctance on the part of many workers to recognize these taxa as full genera, despite its subsequently having been pointed out that genitalic differences exist in both sexes and that non-genitalic characters can be found in some cases (PRINCIPI1977), and that fundamental differences in biology may also occur (SÉMÉRIA,1977). It is my view that the venational pattern of Chrysopa represents the plesiomorphic condition in this subfamily. Most of the genera separated from Chrysopa by earlier systematists were based upon venational apomorphies, these being about the only conveniently observable external modifications in this morpho- logically conservative subfamily. There is however, no obvious reason, except for convenience in sorting specimens, why relatively trivial-appearing vena-

@ 1982 Association Mondiale des Névropteristes U.S. Copyright Clearance Center Code Staternent : 0223-5137/82/020069-07 $ 2.00/0 tional alterations, such as rearrangement of gradate crossveins, should be given more importance in classification than probably biologically more significant modifications of reproductive, pheromonal, and larval structures, as well as features of developmental biology. As a paraphyletic taxon, Chrysopa in the wide sense has been so broadly construed that one can not make meaningful generalizations concerning it, regarding phylogeny, zoogeographic relations or larval and adult biology. 1 therefore am in agreement with ASPOCKet al. (1980) regarding recognition of the former subgenera at the generic level. As they point out, it is possible, when dealing with a regional fauna, to recognize the majority of the genera on eidonomic characters.

In the first paper of this series, Plesiochrysa was described as a subgenus of Chrysopa, because it appears quite closely related to Chrysopa S. str., based upon genitalic characters. This is not the case with the genus described below.

Ceraeochvysa, new genus

Mandibles srnall, asymmetrical, left-toothed ; scape often striped or suffused with red or black, flagellum usually slightly longer than wings. Pronotum without micropoculae, often with lateral red spots or stripes. Wings (Fig. 1) : tips pointed, psm occasionally with included croasveins (see ADAMS1967 for explanation), gradate series regular, last gradate ce11 of forewing rectangular, six apparent pseudomedial crossveins dis'cal to intramedian cell. Male abdomen (Fig. 2) without microtholi, eight and ninth sternites fused, often border of ninth sternite, its apodemes, and/or apodemes of 9T+ ectoproct produced as lobes or hooks ; tignuin absent, gonarcus bearing two short upward-projecting horns (Fig. 4, h) : arcessus usually with median hook flanked by lateral processes (Fig. 5) much as in Leucochrysa,/Nodita (Fig. 6-7), gonocristae present or absent, gonapsis (Fig. 3) in most species elongate. Spermatheca (Fig. 8, 12) elongate, contorted, with ventral impression, connected to bursa by a long slit ; bursa often with enlarged lateral glands (Fig. 8) ; area anterior to subgenitale usually sclerotized in mature individuals. Larvae trash carriers ; adults, as indicated by crop contents, not carnivorous, mostly honeydew or nectar feeders.

Type species : Chrysopa cincta Schneider 1851.

The name comes from Greek keraio, meaning horn, referring to the processes on the gonarcus. In cincta, the arcessus is subdivided into two sclerotized regions, with additional horns on the basal part, and the lateral projections flanking the arcessus tip are not present; in teneral specimens, only the tip of the hook of the ninth tergite and ectoproct is sclerotized, and the gonapsis is sclerotized at the posterior end only. This species is wide ranging, from Florida to Argentins, westward to the Galopagos Islands. Detailed biological studies may demonstrate some sibling species, consequently synonymies of cincta must at present be regarded as somewhat provisional.

Genitalia in this genus often exhibit striking maturational changes, but offer excellent characters in both sexes. It is usually not possible reliably to identify specimens on eidonomic features alone. TJEDER(1966) referred C. bimaculata (i. e., C. valicla) to the Old World genus Glenochrysa, which shares with Ceraeochr~salack of tignum, presence of gonapsis, and enlarged spiracles in males of some species. Enlarged spiracles also occur in some species of Suarius as yet undentified, and do not necessarily indicate relationship. Glenochrysa differs in lacking the gonarcus horns and lateral processes of arcessus tip, in having a transverse gonapsis, pillboxed shaped spermatheca, and in modified wing venation. In Ceraeochrysa the absence of tignum, pseudopenis, micropoculae, and microtholi indicate at best a distant affinity with the Chrysopa- Meleoma-Plesiochrysa complex. The gonarcus + arcessus of Ceraeochrysa (Pig. 4-5) closely resembles that of various species of Leucochrysa and Nodita (Fig. 6-7), both in the hook-notch arrangement of the arcessus tip, and in presence of horns on the gonarcus ; furthermore, females of these genera have similarly shaped spermathecae, and similarly enlarged bursal glands commonly occur in each of these genera. They share a pattern of pigmentation, commonly including red or black scape stripes and lateral pronotal stripes (perhaps indi- cative of participation in a mimetic complex rather than genetic affinity). These similarities point to a probable sister-group relationship with Leuco- chrysa/Nodita rather than with Chrysopa in the strict sense.

If this relationship in fact exists, one must account for the presence of the gonapsis in Ceraeochrysa, while it is absent as a plesiomorphy in Leucochrysal Nodita and related genera. 1 do not consider it improbable that in Ceraeo- chrysa the gonapsis may have originated independently of that in Mallada, etc. In this regard, 1 am in agreement with PRINCIPI(1977) concerning the secondary origin of this structure.

Ceraeochrysa is the dominant neotropical chrysopid genus, in terms of numbers of individuals and is high in diversity ; specimens of numerous undescribed species exist in collections. A few wide-ranging species are commonly encountered in neotropical agricultural habitats (MUMA1959, ADAMS unpub.) and will perhaps be important in biocontrol efforts. Only two species, C. placita and C. lineaticornis, are broadly distributed in the United States and Canada, C. lineaticornis extending south to Central America. The gulf states of the U. S. are included in the ranges of several additional tropical species. The number of synonyms given for each species seems to be related only to abundance and extent of geographic distribution, not to variability. Unless otherwise noted, al1 synonymies given below are new, and based upon examination of type specimens. Al1 species previously were in Chrysopa unless otherwise noted.

Ceraeochrysa 1. adornata Lacroix 1925, Brasil (Paris). 2. anceps Navas 1926, Brasil ( Q , Paris). 3. angulata Navas 1929 Colombia ( 9 , Paris) = josephina Navas 1926, Costa Rica (not found). 4. arioles Banks 1945, Costa Rica, [for binaria Navas 1928 (Hamburg, destroyed)] ; Neotype male : " Vergel, Chiapas / 13-V-35 1 light, " " Mexi- CO, A. DAMPF,"" 9 ", " Chrysopa arioles" [in BANKS'Shand]. (MCZ, Harvard). 5. berlandi Navas 1923, Costa Rica, (9,Paris). 6. castilloi Navas 1913, Paraguay, ( Q Munich). 72 7. cincta Schneider 1851, Brasil (Berlin). = advenu Navas 1922, Uruguay (cf, Paris). = alternans Navas 1933. Peru (cf, Berlin). = bessona Navas 1922, Argentina (Q, La Plata) [provisional, not dissected]. = bicarnea Banks 1920, Florida (cf, MCZ). = bilineata Navas 1914, Guatemala ( Q , BMNH). = bina Navas 1924, Cuba ($, MCZ). = caligata Banks 1924, Panama (cf MCZ). = cornuta Navas 1925 (not 1926) (Bracelona ?, not found) [provisionall. = incalis Banks 1915, Peru (cf, MCZ). = habana Navas 1922, Cuba (not found) [provisionall. = iona Banks 1944, Surinam (cf MCZ). = lafonei Navas 1921, Argentiana ($, Paris). = mestiza Navas 1923, Cuba ( Q , MCZ) Alayo 1968. = Chrysopodes sallei Banks 1945, Mexico (cf, MCZ). = villosula Navas, Cuba (MCZ, no abdomen - another in series dissected). = wollebaeki Esben Petersen 1934, Galopagos (Oslo, not seen; other Galopagos specimens dissected) . 8. claveri Navas, 1911, Colombia (lectotype cf, " Museum Paris / Colombie / C. PARZUDAKI1840 " " Chrysopa / claveri / Nav " " Chrysopa / haitiensis / cf SMITH/ compared / with type / det. E. MACLEOD' 68 »). = adoina Banks 1945, Mexico (cf, USNM). = deficiens Navas 1930, Siberia [several Siberias are in Hispano- america] (cf, Barcelona) . = haitiensis Smith 1931 (cf, MCZ). = inexpectata Alayo 1968, Cuba (Inst. of Biol, Havana, not seen). = siberica Navas 1930, " Siberia " ( Q , Barcelona, not seen) [provisionall. = silvana Navas 1913, Brasil (8, Paris). 9. cubana Hagen 1861, Cuba (Not found). = albatala Banks 1945 Brit. Guiana (MCZ). = damiensis jamaicensis Banks, Jamaica (9, MCZ). = epheba Navas 1924 Cuba (not found) (Alayo 1968). = freemani Smith, 1931, Haiti (d<, MCZ). = imbecilla Navas 1935, Barbados (d,BMNH). = scapularis Navas 1914, Brasil (BMNH). = seminole Banks 1924, Florida (9, MCZ). = tolteca Banks 1901, Mexico ( Q , MCZ). = venularis Navas 1913, Jamaica ( Q , Munich). 10. discolor Navas 1914, Guatemala (cf, BMNH). 11. effusa Navas 1911, Mexico ( Q , Paris). 12. fairchildi Banks 1945, Panama (Q, MCZ - not dissected). 13. fiebrigi Navas 1913, Paraguay ( Q , Munich). 14. gradata Navas 1913, Mexico (Munich) [probably = lineaticornis]. = guatemalteca Navas 1914, Guatemala (cf, BMNH) [provisional]. 15. gundlachi Navas 1924 Cuba (cf NICZ). = everes Banks 1920, Fr. Guiana ( Q , MCZ) [provisional]. = furculata Navas 1924, Cuba (cf, Paris). = instabilis Navas 1925, Brasil (cf, Paris). = jacobea Navas 1925, Cuba (9,MCZ). = petersenia Navas 1931, petersoni Navas 1929, Colombia (type destroyed, Hamburg) . 16. infausta Banks 1945, Costa Rica (cf, MCZ, not dissected). 17. lineaticornis Fitch 1856 (cf MCZ). = Allochrysa parvula Banks 1903 (8,MCZ). = columbiana Banks (cf MCZ) (Bram-Bickley 1963) 18. montoyana Navas 1913, Paraguay ( Q , Munich). 19. placita Banks 1908, Colorado (8,MCZ). = forreri Navas 1913, Mexico (cf, BMNH). 20. reducta Banks 1945, Colombia (9, MCZ). 21. rochina Navas 1915, Brasil ( (9, Paris). 22. sanchezi Navas 1924, Cuba ($ MCZ) 23. smithi Navas 1914, W. Indies ($, BMNH). = neotropica Navas 1929, C. and S. Amer. (Hamburg, destroyed). = poeyi Navas 1923, Cuba ($, MCZ). 24. valida Banks 1895, Mexico ( Q , MCZ). = bimaculata McClendon 1901, Texas (type not found). = breviata Banks 1914, Ecuador (cf, MCZ). = composana Navas (cf Paris, description not found). = damiensis Smith 1931, Haiti (cf, MCZ). = limitata Navas 1913, Brasil, Brit. Guiana (BMNH, Barcelona). = lioni Navas 1927, Haiti ( Q , Paris). = longicella Navas 1914, Guatemala (cf, BMNH - has enlarged spiracles). = seminole Banks 1924, Florida (MCZ). = wolcotti Smith 1931, Haiti (Q ?, MCZ).

Manuscrit reçu le 19 mars 1982.

1 am grateful to the following curators for making it possible to examine type material: John LAWRENCE(Museum of Comparative Zoology, Harvard), O. FLINT (U. S. National Museum), Peter BARNARD(British Museum (Natural History) , S. KEZNER-PILLAULT(Paris), K. GÜNTHER(Berlin), W. DIERL (Munich), F. ESPANOL(Barcelona) .

ADAMSP. A., 1967. - A review of the Mesochrysinae and Nothochrysinae (Neuroptera : Chrysopidae). Bull. Mus. Comp. 2001. Harv. 135 : 215-238.

ADAMSP. A., 1982. - Plesiochrysa, a new subgenus of Chrysopa (Neuroptera). Neuroptera international II (1) : 27-32.

ASPOCKH., U. ASPOCKand H. HOLZEL,1980. - Die Neuropteren Europas. Goecke and Evers, Krefeld.

HOLZEZH., 1970. - Zur generischen Klassifikation der palaarktischen Chrysopinae. Eine neue Gattung und Zwei neue Untergattungen der Chrysopidae (Planipennia). Zeitschr. der Arbeitgemeinschaft Osterr. Entomologen 22 : 43-52. MUMAM., 1959. - Chrysopidae associated with citrus in Florida. The Florida Entomolo- gist 42 : 21-29.

PENNYN., 1977. - Lista de Megaloptera, Neuroptera e Raphidio~terado Mexico, América Central, ilhas Caraibas e America do Sul. Acta Amazonica 7 (4) supplemento : 1-61.

PRINCIPINI., 1977. - La morfologia addominale ed il suo valore per la discriminazione generica nell'ambito delle Chrysopinae. Boll. Ist Entom. Univ. Bologna 31 : 325-360.

SÉMÉRIAY., 1977. - Discussion de la validité taxonomique du sous-genre Chrysoperla Steinmann Nouv. Rev. Ent. 7 : 235-238.

TJEDERB., 1966. - Neuroptera-Planipennia. 5. Family Chrysopidae. S. African Atzimal Life 12 : 228-534. Fig. 1-4, 8-12 : Ceraeochrysa cincta. Fig. 1, wings (8 Honduras). Fig. 2, abdominal apex, gonapsis indicated by dotted lines. (Figs. 2-4: mature 8, Venezuela) ; Fig. 3: gonapsis, dorsal view. Fig. 4 : gonarcus complex, oblique view, showing horn-like processes (hl and field of gonocrista-like scales on membrane (g). Fig. 5 : C'. lineaticornis (Florida), apex of arcessus, showing median claw-like hook and lateral processes. Fig. 6-7 : Nodita nictheroyana Navas (holotype), gonarcus complex, showing horns (h), arcessus hook, and lateral processes. Fig. 8-12: Ç. cincta. Fig. 8 : copulatory bursa, spermatheca and bursal glands, left lateral view. Fig. 9, 10, subgenitale, ventral and lateral views, entire area between subgenitale and 7th sternite sclerotized (mature 9). Fig. 11 : subgenitale (teneral). Fig. 12 : spermatheca, ventral, showin- ventral impression (teneral). Bibliography of the Neuropterida

Bibliography of the Neuropterida Reference number (r#): 1304

Reference Citation: Adams, P. A. 1982 [1982.12.30]. Ceraeochrysa, a new genus of Chrysopinae (Neuroptera) (Studies in New World Chrysopidae, Part II). Neuroptera International 2:69-75.

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