A Review of the Mesochrysinae and Nothochrysinae (Neuroptera: Chrysopidae) PHlbLIP A. ADAMS HARVARD UNIVERSITY VOLUME 135, NUMBER 4 CAMBRIDGE, MASSACHUSETTS, L1l.S.A. FEBRUARY 24, 1967 PUBLlCATlONS ISSUED OR DiSTRlBUTED BY THE MUSEUM OF COMPARATIVE ZOOLOGY HARVARD UNIVERSITY BULLETIN1863- BREVIORA1952- MEMOIRS18641938 JOHNSONIA,Department of Mollusks, 1941- OCCASIONALPAPERS ON MOLLUSKS,1945- Other Publications. Bigelow, H. B. and W. C. Schroeder, 1953. Fishes of the Gulf of Maine. Reprint, $6.50 cloth. Brues, C. T., A. L. Melander, and F. M. Carpenter, 1954. Classification of In- sects. $9.00 cloth. Creighton, W. S., 1950. The Ants of North America. Reprint, $10.00 cloth. Lyman, C. P. and A. R. Dawe (eds.), 1960. Symposium on Natural Mam- malian Hibernation. $3.00 paper, $4.50 cloth. Peters' Check-list of Birds of the World, vols. 2-7, 9, 10, 15. (Price list on request. ) Turner, R. D., 1966. A Survey and Illustrated Catalogue of the Teredinidae ( Molluscs: Bivalvia) . $8.00 cloth. Whittington, H. B. and W. D. I. Rolfe (eds.), 1963. Phylogeny and Evolution of Crustacea. $6.75 cloth. Proceedings of the New England Zoological Club 1899-1948. (Complete sets only. ) Publications of the Boston Society of Natural History. Publications Office Museum of Comparative Zoology Harvord University Cambridge, Massachusetts 021 38, U. S. A. @ The President and Fellows of Horvard College 1967. A REVIEW OF THE MESOCHRYSINAE AND NOTHOCHRYSINAE (NEUROPTERA: CHRYSOPIDAE) ABSTRACT tirely new, necessitating a preliminary re- classification of the family." In this preliminary subfamilial classifica- There appears no justification for rctain- tion of the Chrysopidae, Mesochrysopidae ing in a separate family the Mesozoic forms, is reduced to subfamilial rank. Keys to sub- which are placed in the family Meso- families and to genera of Nothochrysinae chrysopidae. These already have achieved are presented. The Nothochrysinac, as the alignment of the inner gradatc veins newly constituted, is characterized by re- which precedes the pseudomedia of the tention of jugum and frenulum, lack of alar later forms. In addition, they show the fun- tympana1 organ, archaic pseudomedia (ex- damental chrysopid characteristics: many cept in Nothochrysa and Dyspetochrysa) straight, unforked branches of Ks, which and little sclerotized prosternum. It includes diverge from 13 at a wide angle; two reg- Paleochrysa, Archaeochrysa, Dyspetochrysa, ular gradate scries; MP1 and MP2 few- Tribochrysa, Dictyochrysa, Triplochrysa, branched, intersecting the wing margin IIypochrysa, Kimochr!jsa, Pamochrysa, Pi- ncar the base, so that most of the discal machrysa, and Nothochrysa. The living area is occupied by the K-Ks-MA system. species of Nothochrysinac are cataloged, These basic features of proportion, while difficult to define, readily separate the and the North American species described Chrysopidae from all other families. and illustrated. The Apochrysinae, a distinctive and ho- New fossil taxa are: Archaeochrysa, new mogeneous group, has recently been revised genus for Paleochrysa creedi Carpenter, by Kimmins ( 1952113 ) . The Chrysopinae fracta (Cockerell) and paranervis n. sp. constitutes a receptacle for the remaining (Florissant, Colo. ) ; Dyspetochrysa, n. &en. vast and various assemblage of forms, for Tribochrysa vetuscula Scudder. New doubtless requiring subdivision when bet- Recent species arc: Pi~nachrysaalbicostales, ter understood. Baja Calif., and Arizona; fusca, intermedia All described genera of Mesochrysinae and nigra, southern Calif. and Nothochrysinae are included in the keys, and all species have been cataloged. INTRODUCTION Descriptions and illustrations have been This study comprises the first section of included for all living North American a taxonomic revision of the North American chrysopidae. ~h~ concept of the subfamily ' Although this paper was submitted for publica- tion prior to the appearance of Tjeder's paper, Nothochrysinae as herein is en- 1966, it has been possible to include his new taxa. IIe deli~nitsthe Nothochrysinae (=Dictyochrysinae) ' California State College, Fullerton. similarly. Hull. Mus. Comp. Zool., 135(4): 215-238, February, 1967 215 216 Bt~lle~inMuseum of Comparative Zoology, Vol. 135, No. 4 spccies, and ior such fossil or Old World ences, and the University of California, species as are of particular intcrest. I have Santa Barbara. not examined material of Dictl\ochrysa or Triplochr~~sa,which were reviewed by DlSCUSSlON OF CHARACTERS Kimmins ( 1952a). The wing-coupling apparatus consists of It is interesting that thc southwestern a large jugal lobe on the fore wing, and United States should have such a rich rep- a frenulum, bearing several long setae, on resentation (two genera, six species) of the hind wing; this is essentially the same this archaic: group, which has changed but apl'aratus as in FIemerobiidac, etc. The littlc since the Miocene (Adams, 1957). Chrysopinae may have a wealc frenulum, This concentration of thesc relics is rivaled 1)ut the jugal vein is thin, and therc is no by South Africa, with two genera and four jugal lobe. Loss of the wing-coupling ap- species, and Australia and Tasmania, also paratus appears associated with narrowing with two gcnera and four species. Such a and strengthening of the wing base, and distribution contributcs to the mounting is probably OF great adaptive significancc. cvidence that thc southwestern United A similiar modification has occurred in States has served as an evolutionary rc- the evolution of the Myrmeleontidae and fugium for the Neuroptera. Ascalaphidae from an osmyloid ancestor (Adams, 1958), and in the.Mantispidae. ACKNOWLEDGMENTS In the Nothochrysinae, there is no obvi- Grateful acknowledgment is made of the ous tympanal organ (Friedrich, 1953; Er- help and encouragement of F. M. Carpenter hardt, 1916) in the base of 11 in the fore and P. J. Darlington, Jr., of thc Museum wing, and the stem of M is easily visible of Comparative Zoology ( MCZ ) , Harvard. extcnding in a straight line adjacent to 8. Ellis MacLeod, of the Biological Labora- In the Chrysopinac, the tympanal organ tories, I-larvard, has made many valuablc forms a conspicuous bulgc in R, at the criticisms, and has given generously of his point where Cu diverges; the base of M time during the preparation of the manu- usually is co~ilescedwith R, but if visible, script. D. E. Kimmins has kindly examined makes a dctour posteriorly around the tym- type material and has made useful sugges- panal organ. Probably this is an auditory tions. Material has also bccn made avail- organ ( Adams, 1962). able through the courtesy oi C. D. MacNcil, In Nobilinus (Apochrysinae), the tym- California Academy of Scienccs (CAS), J. panal organ involves a large area betwecn D. Powell, University oi California, Berke- K and M, but is longcr and does not form Icy, California Insect Survey (CIS), J. N. a bulgc on the underside of 11, as in Helkin, University of California, Los Angelcs Chrysopinae. (UCLA), P. 11. Timberlake, University of Another character, probably of great California, Kivcrside ( UCII ) , L. Stange, adaptive significance, is the pseudomedia, University of Califorilia, Davis ( UCU ) , J. which diffcrs fundamentally ill the more Lattin, Oregon State College (OSC), E. I. primitive chrysopids, and in the Chryso- Sleeper, Long Reach State College, J. E. H. pinae-Apochrysinae. In Pimachrysa (Figs. Martin, Entomology Research Branch, Cnn- 1, 2), Hypochrysa (Fig. S), and in most ada Eepartment ot Agriculture, Ottawa of the fossil genera, Psm is composed of (CNC), Hugo Rodeck, University of Colo- crossveins alternating with the branches rado Museum, and Floyd Werner, Univer- of Rs $- MA, and is merely a basad exten- sity of Arizona. sion of the inner gradate serics. The coursc Financial assistance has been provided of the longitudinal veins and compositio~i through grants-in-aid from Sigma Xi-KESA, of the primitive pseudomedia is particularly the American Academy of Art5 and Sci- clear in Mypochrysa. In the Chrysopinae (Fig. 45),and Apochrysiilac (Fig. 44),ancl of MP1 with Rs -1- MA. This evolutioilary in Nothochrysa (Fig. 3), Psm, at least sequence can be seen by coinparing I"&' 1 rures basally, is composed of overlapping zig- 41, 5, 4, and 2. zagged branches of Rs and M, with no In Chrysopinae and Apochrysinae, the crossveins between them. ( Some specimens wing flexes along a line immediately an- of A7. fulviccps show no overlap.) In a few terior to Psm (dotted line, Figs. 44, 45). genera of Chrysopinae, such as Yumachrysn To facilitate this flexion in the fore wing, and Chrysopiella, there is n tramition trom the first sectorial crossvein (first crossvein the primitive arrangement apically, with distal to thc base of Rs + MA) and the no o~~erlap,to the more advanccd arrange- branches of Rs + MA are interrupted or ment basally, with overlapping veins. articulatect at the point of intersection with Primitively, the longitudinal veins forked the pseudomedia. In the Nothochrysinae, at the posterior wing margin, except for the sectorial crossvein is always interrupted, the aids in both wings, ant1 CUP in the but the branches of Rs + MA never are so hind wing. This condition may be seen in ( except in Nothochrysa ) . Flexion along the the hind wing of Archaeocl;zry.sa (Figs. 40, pxudomcdia is accomplished, in Hypo- 41). There is a tendency for the point of chrysu, by folds traversing the crossveins furcation to move proximally until it reaches of Psm (Fig. 5). In Pimachrysa, there the outer gradate crossveins; when this appears to be no distinct line of flexion, ex- occurs, the pscudomcdial cells appear each cept perhaps in P. nigra (Fig. 4). In this to give rise to two marginal veinlets. This spccies the veins show no obvious wcaken- process bcgins at the wing base, and pro- ing or articulation, but most specimens ceeds apically; the sequence is best secn have a slight wrinkle in the membrane in the fore wing of Archaeochrysa para- par:tllel and anterior to Psm. A similar line newis (Fig. 40). Frequently the longi- of flexion occurs between MA and MP in tudinal veins fail to fork at all.