Pure and Applied Research in Neuropterology. Proceedings of the Fifth International Symposium on Neuropterology. Cairo, Egypt, 1994. Canard, M., Aspock, H. & Mansell, M.W. (Eds). Toulouse. France, 1996. Pp. 19-30.

Venational homologies and nomenclature in , with comments on the Myrmeleontoidea (Insecta: )

Phillip A. ADAMS California State University, Fullerton, U.S.A.

ABSTRACT Fossil history and studies of comparative morphology have largely corroborated the interpretation of venational homologies in Chrysopidae advanced by TILLYARD (1916b), based on pupal tracheation. Despite difficulties in identifying terminal branches of the Posterior Median (MP) and Anterior Cubitus (CuA), it is shown that the latter is multibranched.

Venational nomenclature used by SCHNEIDER (1851), NAVAS (1923), and BANKS (1944) are reviewed. Differences in the venation of fore- and hindwings have led to mis- interpretations in both Chrysopidae and Myrmeleontoidea. In the hindwings of and Myrmeleontidae the second branch of MP is often mistaken for CuA.

Key words: Chrysopidae, Myrmeleontidae, wing venation, pseudomedial, pseudocubital.

REVIEW The first comprehensive treatment of chrysopid morphology, by SCHNEIDER (1851), with its detailed system of venational nomenclature, is the starting point of this study. Although the names of longitudinal veins were not yet standardized, nor was the convention of naming crossveins and cells for the vein preceding them, aspects of SCHNEIDER'Ssystem remained in use for nearly 100 years. Neither N. BANKS nor L. NAVAS recognized the existence of pseudomedia (Psm) or pseudocubitus (Psc), and persisted with the terminology they had used before the publication of TILLYARD'S (1916b) paper (Table 1). When these authors mention "cubital cells", or "cubital cells beyond the divisory", they refer to cells distal to the intramedian cell, between Psm and Psc. While this count of cells provides a useful taxonomic character, it may be misleading to the modem worker, who mistakenly might expect "cubital cells" to refer to cells posterior to the "cubitus", between Psc and the posterior margin of the wing. Ph.A. Adams

Table 1. Chrysopid venational nomenclature.

Modern TILLYARD, SCHNEIDER, NAVAS, BANKS, usage 1916 1851 1923 1944

Fore wing Costa Costa Costa Costa Costa Subcosta Subcosta Subcosta Subcosta Subcosta Radius Radius Radius Radius Radius Radial sector (+MA) Radial sector Second Rs R s R s Rs branches - ending on Psm Banksian sectors - Intermediates Intermediates Inner gradates Inner gradates Inner gradates Inner gradates Inner gradates Outer gradates Outer gradates Outer gradates Outer gradates Outer gradates Pseudomedia Pseudomedia First Rs Procubitus Medius Intramedian cell Intramedian cell Areole of cubitus Divisory cell Divisory cell Median cells Median cells - Cubital cells Cubital cells MP2 Median loop Divisory veinlet Divisory Divisory vein branch CuA and Psc Cubitus Cubitus Cubitus Cubitus Pseudocubitus CUP Cubitus 2 Cubital branch Cubital sector - 1A First anal Postcosta Postcubitus - 2A. 3A Anal 2, 3 Forked marginal Axillary veinlet - vein 1, 2 ...... Hindwing Psm Pseudomedia Procubitus - Psc Pseudocubitus Cubitus Cubitus - CuA Cubitus 1 Postcosta Postcubitus -

ABBREVIATIONS A1, A2 = first and second anal veins dcc = distal cubital cell acsx = apical subcostal cross vein i m c = intramedian cell b = Banksian cell (in Chrysopidae); M A = media anterior = basal piece of MA in forewing NZF = mf = medial fork of myrmeleontoids (in Fig. 5) o = oblique vein, MP2 crossing to b .1 . = Banksian line CuA in myrmeleontoid forewing Bn = Banksian sectors (TILLYARD) o .g . = outer gradate crossveins C uA = cubitus anterior rf = radial fork CuF = cuf = CF = cubitus fork R S = radial sector CUP = cubitus posterior s n = branches of RS (TILLYARD) Icu, 2cu = first, second cubital cell x = crossvein Chlysopid venation

COMSTOCK & NEEDHAM (1898-1899) emphasized tracheation as a basis for venational homologies, maintaining that the veins formed around the tracheae, which determined their course. Objections were raised against over-reliance on tracheal information, with WOODWORTH (1906) indicating that there is no apparent difference between veins that are traversed by tracheae and those that are distant from them, that there is great variation in paths taken by tracheae, and that the tracheae form sinuous paths, while the veins make abrupt turns. NEEDHAM (1935) introduced a discussion with "I begin where veins begin, with tracheae", indicating his firm belief in the role of tracheae. HOLDSWORTH (1942) by contrast, showed that in Pteronarcys Newman (Plecoptera), the vein channels appear before the tracheae penetrate the wing pad, demonstrating that the venational pattern is not primarily dependent upon tracheation.

TILLYARD (1916b) followed COMSTOCK & NEEDHAM (1898-1899) in the use of pupal tracheation as a basis of venational homologies. He was able to illustrate for the first time the tortuous paths of the veins in both wings of chrysopids (Fig. I), and his system of chrysopid venational terminology, with minor changes, is the basis of that in use today. Recognizing the structural similarity of the Psm and Psc to the undulation of veins giving rise to the Banksian lines (b.1.) of myrmeleontids (Fig. 2), he named the branches of the chrysopid Radial Sector (Rs) which intersect the Psm as "Banksian sectors", and several of the cells between the Psm and Psc as "Banksian cells". In addition, he accurately noted the extent of CuA and its branches, after its junction with Psc, and in the hindwing the origin of the Psc from the stem of the second branch of MP. Moreover, he correctly analyzed the venation of the Apochrysinae.

Fig. 1. Tracheation of pupal wings of Mallada signata (Walker). Note difference in paths of "M2" (MP2) and "Cul" (CuA) in fore- and hindwings, zig-zagging and overlap in tracheae forming Psm and Psc. From TILLYARD (1916b). Fig. 2. Wing venation of Nrsydrion nigriner.i#e Esben-Petersen (Nyrnphidae). The medial fork MF of the hindwing is configured similarly to the cubital fork CF of the forewing; the anterior cubitus CuA of the hindwing is reduced and concave. Arrows indicate the boundary between medial and cubital systems. From ADAMS (1958).

COMSTOCK (1918) reviewed the earlier literature on wing venation, and presented a standard system of venational nomenclature for all orders, based largely on studies of the development of wing tracheation. Despite its shortcomings, the COMSTOCK- NEEDHAM system of venational nomenclature is still in use. He largely followed TILLYARD'Snomenclature for the Chrysopinae, and presented a drawing of chrysopine pupal tracheation by R.C. SMITH. In the absence of pupal apochrysine material, he unsuccessfully attempted to analyse the complex venation of Apochrysa Croesus Gerstacker as a representative of the Apochrysinae. He noted the absence of the intramedian cell and, in the hindwing, the origin of the apparent pseudocubitus (Cul) from the posterior branch of the median (M), as in the myrmeleontoids. COMSTOCK (1918) failed to recognize the presence of a Psm and Psc, as well as not observing the similarity in origin of the Psc in the hindwings of Chrysopinae and Apochrysinae and, despite having seen TILLYARD's (1916b) paper, he erroneously concluded that the wings of "Apochrysidae" were closely analogous to those of Myrmeleontidae.

The work of ADOLPH (1879) was reviewed by COMSTOCK (1918) and will not be discussed here. LAMEERE (1923) and MARTYNOV (1930) revived ADOLPH'S emphasis on convexity and concavity of wing veins as an aid to determining homology, detecting the presence of an anterior branch of M, fused basally with Rs in living . In Neuroptera, MA is more obvious than in most orders, appearing as a convex branch from Rs and often, in and the myrmeleontoid families, MA is more extensively branched than the branches of Rs, occupying a triangular area termed the "radial cuneate area" (COMSTOCK 1918). In the hindwings of several families, excepting Chrysopidae, MA is connected to MP by a sinuous, nearly longitudinal vein, which is probably the basal piece of MA. Where venal fusion is more obvious, such a sinuous formation is quite usual (e.g. KUKALOVA-PECK 1991: fig. 6, 27 H, hindwing). In the forewing, various crossveins have unconvincingly been suggested as the basal piece of MA by CARPENTER (1940). Chrysopid venation

The LAMEERE-MARTYNOV approach has been extended and applied to fossil as well as extant insects by KUKALOVA-PECK (1991), who maintains that each of the veins consists of convex anterior, and concave posterior branches, and finds evidence of this condition in the precosta, costa, and subcosta, as well as the other main veins in the most archaic insects. In the hindwings of Neuroptera, convexity-concavity appears to be reversed in MP2 which is convex, and CuA, which becomes a concave vein (ADAMS 1958). KUKALOVA-PECK (1991: fig. 6,27 H) interprets the venation of Varnia Walker () as having undergone fusion of CuA and MP, CuA retaining its convexity after separating from MP distally. But, in that case, the entire CuA is fused with MP. In the ithonid Oliarces Banks, CuA is a concave vein which does not coalesce with MP, as in other Neuroptera.

Fossil evidence The well-preserved wings of the pre-nothochrysine Protochrysa aphrodite Willrnann & Brooks (Fig. 3), from the early Tertiary, afford the opportunity to examine a venational pattern which, while primitive, is sufficiently similar to that of recent chrysopids to allow reliable comparisons of venational homologies. Figure 3 has been modified by completion of veins across minor gaps in the original rendering where this is not controversial; the proximal branch of CuA in the forewing is discernible in published photographs and has been added. It will be noted that a series of inner gradate crossveins extends basally to MP1, but there is no zig-zagging of the branches of Rs + MA in either wing, and thus no trace of an incipient Psm. In the photographs (WILLMANN & BROOKS 1991: plate I, 2) the location of the median flexion line in the forewing (WOOTTON 1979) appears to be represented by a weakening of MP1 and the basal rs + ma crossvein near their intersection, as is the case in recent Nothochrysinae.

Fig. 3. Wing venation of Protochrysa aphrodite Willmann & Brooks. Note smoothly curving branches of radial and medial systems and nearly homonymous venation in fore- and hindwings. Modified from WILLMANN & BROOKS (1991). Some modem nothochrysines have additional points of flexion in the Rs area, as illustrated by ADAMS (1967): these are not indicated in WILLMANN & BROOKS' (1991) illustrations of P. aphrodite, and were probably absent. The Psc is weakly developed, but is manifest more strongly in the forewing by zig-zagging of the branches of MP and Rs + MA. There is no overlap of veins in the Psc, zigzagged veins alternating with crossveins. CuA vein is five-branched in the forewing, but in the hindwing it occupies a smaller expanse of the wing area, and its fusion with MP2 is slightly more extensive.

Subfamily Nothochrysinae In many respects, members of Nothochrysinae represent various intermediate conditions in the degree of zig-zagging of veins which ultimately gives rise to the straight Psm in both wings of Chrysopinae. Plmachrysa Adams (Fig. 4) exemplifies a stage where Psm is hardly developed. The pseudomedia may be quite well developed in this subfamily, e.g. Nothochrysa californica Banks. BROOKS & BARNARD (1990) list as a key character for this subfamily "in forewing, Psm continuous with inner row of gradates ..." but this character, while traditionally used, and valid for the European species of Nothochrysa McLachlan, does not apply to all of this subfamily, as a critical examination of their figures, and of ADAMS'(1967) figure of N. californica will show.

Pig. 4. Wing vendtion of P~n~acllrysafuscaAdams, exhibiting abrupt deflection of MA and branches of MP toward hind margin, rudimentary Psm and Psc with little overlap of veins. Arrows indicate probable boundary between branches of MP and CuA; dashed line indicates point of flexion. Modified from ADAMS (1967).

Subfamily Chrysopinae The extent of wing area occupied by branches of CuA in chrysopids has not generally been appreciated, despite its early documentation by TILLYARD (1916b) and COMSTOCK (1918). Their evidence from tracheation indicates that in modern Chrysopinae, this vein gives rise to branches, four in Chrysopa nigricornis Burmeister, Chrysopzd venation and three in Mallada signata (Walker) (Fig. 1). It is often difficult or impossible to make a reliable assessment of the extent of CuA based upon analysis of venation, but sometimes it is possible. In the plesiomorphic condition, the hind marginal forking is extended basally to include the ends of MP2, each vein terminating in a single fork as may be observed in the forewing of Protochrysa Kolbe (Fig. 3). As an apomorphy, the point of forking moves anteriorly, the branches of the fork appearing as crossveins arising from Psc. Often one or several branches of Rs, MA or MP fork more than once, thus giving rise to three or more of these transverse veins reaching the wing margin, or they may not fork at all. In such cases, it may not be possible to trace the course of each vein as it traverses Psm and Psc, and the extent of CuA is obscure.

Cubitus anterior, as is usual in neuropterous hindwings, occupies a slightly smaller area than in the forewing. In the hindwing, Psc is aligned with, and appears to originate from MP2, rather than with CuA as in the forewing. This difference in origin of Psc was illustrated by both TILLYARD (1916b) and COMSTOCK (1918), but was not discussed. At first glance, the Psc in fore- and hindwings appear homonymous, but careful analysis proves this is not the case. In the Apochrysinae, the condition of Psc in the hindwings was at least partially responsible for the misinterpretation by COMSTOCK (1918), referred to below.

Subfamily Apochrysinae TILLYARD (1916b: fig. 6) correctly interpreted the venational homologies of Apochrysinae, according to the COMSTOCK-NEEDHAMsystem. But COMSTOCK (19 IS), despite having seen TILLYARD'S (19 16b) paper, failed to recognize the presence of an obvious Psm and Psc. Citing the absence of the basal subcostal crossvein and intramedian cell in the forewing, as well as the presence of a hypostigmatic cell, he considered Psm and Psc to be homologues of M1 + 2 (MP1) and Mg + 4 (MP2 + CuA) in Myrmeleontidae. As final evidence, he mentioned the formation of Psc in the hindwing.

In his study of the Myrmeleontidae, COMSTOCK (1918) correctly observed, in contrast to the findings of TILLYARD (1916a), that the prominent convex forking vein in the hindwing of seems to be the homologue of "Mg + 4 + Cul" (MP2 + CuA) in the forewing, but is of entirely different origin, consisting only of "M3 + 4" (MP2), with no participation of CuA. The origin of "M3 + 4'' (Psc) as a branch of M rather than of C + M provided evidence to COMSTOCKof an apochrysine-myrmeleontid similarity, despite his observation of a comparable basis of Psc in the hindwing of Ch. nigricornis.

Myrmeleontoid venation It is of considerable interest that COMSTOCK (1918) and TILLYARD (1916a) came to diametrically different conclusions concerning the nature of hindwing venation in Myrmeleontidae, based upon studies of tracheation. The generally accepted interpretation at that time, and until quite recently, was that myrmeleontid fore- and hindwings are homonymous, the prominent convex forking vein lying more or less in the middle of the wing in both being considered CuA. This obvious interpretation was used by BANKS, NAVASand TJEDER, throughout their publications, and is still encountered. On PRA. Adams the other hand, COMSTOCK(1918) considered that the prominent convex forking hind- wing vein was "M3 + 4" (MP2).

ADAMS (1958) undertook a study to resolve this question of hindwing venational homologies in the myrmeleontoid families, by surveying venation in the extinct order Protoperlaria, in Megaloptera, and among Neuroptera in the families Polystoechotidae, Osmylidae, , , Myrmeleontidae, and Ascalaphidae. In all of these, he concluded that the usual convexity-concavity relationships were altered in the hindwing, MP2 becoming convex, and CuA concave. The wing articulations, basal patterns of venation and venation of entire wings were illustrated. These indicate a progressive diminution of the hindwing area occupied by Cu, in various Nymphidae, Myrmeleontidae and Ascalaphidae. The venation of the nymphid Nesydrion nigrinerve Esben-Petersen (Fig. 2) is shown here for comparison with the myrmeleontid Vella texana Banks (Fig. 5).

Fig. 5. Wing venation of Vella texana Banks (Myrmeleontidae). Similar venation is radically different in the fore- and hindwings. From ADAMS (1958).

Why did TILLYARD (1916a) reach an erroneous conclusion regarding the identity of the cubital veins? Perhaps the answer lies in the method of study. In clipping the wing pads from the pupa in preparing his figure 7, the basal connection between the tracheae of MP was destroyed, and perhaps he never observed it, as did COMSTOCK (1918: fig. 195). Furthermore, the interpretation of COMSTOCK differed from the conventional, and as it was suspiciously heterodox, it was subsequently largely ignored. Chrysopid venation

Taxonomic characters of chrysopid wings The number of costal and radial crossveins are easy to determine and useful. Gradate crossveins can usually be counted easily: if the inner gradate series extends basally, paralleling Psm, one should differentiate the basal series, but this may prove difficult or impossible to distinguish from the distal series, and an illustration should be included. Figure 6 depicts a species of Leucochrysa McLachlan with such a basal series of gradates, as well as a supplemental intermediate gradate series. Occasionally, instead of a single crossvein terminating the series of apparent pseudomedial crossveins, there may be a short series, resembling a gradate series. In Fig. 6, two may be seen. Length of the gradate cells may be expressed as a lengthlwidth ratio of the third cell anterior to Psm, in species where the inner gradates do not extend far basally. NAVAS, and occasionally BANKS, referred to the branches of Rs + MA intersecting Psrn (plus the basal rs crossvein) as "intermediates". Obviously, this number is directly related to the apparent number of "crossveins" or pseudomedials between Psrn and Psc, distal to the intrarnedian cell, comprising MP1, MP2, MA, branches of Rs and one or more distal true crossvein(s). In Chrysopodes frguralis (Banks) (Fig. 7), six pseudomedials are normally present, and four intermediates, exemplifying the usual relationship, where the number of intermediates plus two equals the number of pseudomedials. If the venation is complex, and the number of veins multiplied, this relationship may not be valid. The number of pseudomedials can usually be made accurately, with no need for interpretation, in contrast to intermediates when the gradate series extends basally. The count of intermediates as a practical character has fallen into disuse for a good reason.

Fig. 6. Wing venation of Leucochrysa McLachlan sp. Specialized wing showing Psm curving anteriorly to join outer gradates, intermediate gradates between inner and outer series, and short series (2) of external gradates. The linear Psc is unusual.

ADAMS (1967) suggested the degree of overlap of veins comprising Psm, as a possible taxonomic character. This is most easily determined by tracing the course of these veins, starting basally, as they pass along Psrn and cross to Psc. In some genera these veins may alternate with crossveins (plesiomorphic) while in others, Psrn characteristically consists of overlapping veins for its entire length. An example may be seen in Fig. 7, where a short crossvein (x) connects RslO and Rsll at Psm. Unfortunately, this character has proven too variable to be of use. Ph.A. Adams

The widths of the pseudomedial and pseudocubital areas may be compared as a ratio, measured at the distal end of the third medial cell, where MP2 crosses the pseudomedial area. This is particularly helpful in genera such as Nacarina Navhs where the numbers of veins and crossveins is extremely variable, but proportions of wing areas are specifically characteristic.

Fig. 7. Wing venation of Chrysopodss frguralis (Banks). A generalized chrysopine with extensive overlap in veins comprising Psc, whereas in Psm only MP2 overlaps MP1.

BROOKS & BARNARD (1990: fig. 1) denote the veins passing from Psc to the posterior wing margin as "posterior medial crossveins". While the vein indicated by the tip of the pointer in their figure is indeed the termination of one of the branches of MP, it is not a true crossvein. Nor is it clear whether it is intended to apply this terminology to all the veins passing from Psc to the posterior margin. To include them all under this term is inaccurate, and it is impractical and unnecessary to attempt to identify them all as branches of specific veins. I prefer the term pseudocubital "crossveins" (in quotes), or simply pseudocubitals.

DISCUSSION Despite its recent fall from prominence, evidence provided by tracheae was responsible for unravelling the unrecognized complexity of chrysopid venation, and for recognition of forewing fusion of CuA and MP2 in the forewing of myrmeleontids. Chrysopid venation

There is an element of conservatism regarding branching patterns of tracheae which may on occasion be useful in understanding venational complexities. It is now, however, recognized that there is no evidence that the veins form around the tracheae. Excessive reliance on tracheation patterns alone may lead to quite erroneous conclusions, as for example, in the case of the suggested "crossover" of Rs and MP1 at the nodus of Anisoptera by COMSTOCK & NEEDHAM (1898-1899), recently disproved by RIEK & KUKALOVA-PECK (1984). Evidence from comparative anatomy and the fossil record is far more convincing. In the case of the Chrysopidae, the fossil record validates conclusively the interpretation which resulted from the tracheational evidence.

In the myrmeleontoid families, taxonomists should be aware of the two different interpretations of hindwing venation, and which of the two is being applied in a particular publication. Despite its obvious appeal, the interpretation of TILLYARD (1916a) and most other previous workers on the group, should not be followed.

In Chrysopidae evidence from tracheation, comparative anatomy and fossils indicate that CuA in both wings gives off more branches than one would expect.

REFERENCES ADAMS, P.A. 1958. Studies in the Neuroptera, with special reference to wing structure and evolution in the Osmyloidea. Ph.D. Thesis, Harvard University, Massachusetts, U.S.A. 120 pp., 16 pl.

ADAMS, P.A. 1967. A review of the Mesochrysinae (sic) and Nothochrysinae (Neuroptera: Chrysopidae). Bulletin of the Museum of Comparative Zoology, Harvard University 135: 215-238.

ADOLPH, G.E. 1879. ~berInsektenflugel. Nova Acta. Leopoldinisch-Carolinische Deutsche Akademie der Naturforscher 41: 215-291.

BANKS, N. 1944. Neuroptera of Northern South America. Part 111. Boleti'n de Entornologfa Venezolana 3: 1-34.

BROOKS, S.J. & BARNARD, P.C. 1990. The green lacewings of the world: a generic review (Neuroptera: Chrysopidae). Bulletin of the British Museum (Natural History). Entomology Series 59: 117-286.

CARPENTER, F.M. 1940. A revision of the Nearctic , , Sisyridae, Polystoechotidae and . Proceedings of the American Academy of Arts and Sciences, Boston 74: 159-280.

COMSTOCK, J.H. 1918. The Wings of Insects. 430 pp. Comstock Publ. Co., Ithaca, New York, U.S.A.

COMSTOCK, J.H. & NEEDHAM. J.G. 1898-1899. The Wings of Insects. 124 pp. Comstock Publ. Co., Ithaca, New York, U.S.A.

HOLDSWORTH, R. 1942. The wing development of Pteronarcys proteus Newport (Pteronarcidae: Plecoptera). Journal of Morphology 70: 431-462. Ph. A. Adams

KUKALOVA-PECK, J. 1991. Fossil history and the evolution of insect structures. In: Insects of Australia: a Textbook for Students and Research Workers. Second edition, Chapter 6: 141-179. Commonwealth Scientific & Industrial Research Organization, 1 500 pp.

LAMEERE, A. 1923. On the wing-venation of insects. Psyche 30: 123-132. (Translation from "Sur la nervation alaire des insectes". Bulletin, Classe des Sciences, Acadkmie Royale de Belgique 1922: 138-145).

MARTYNOV, A.V. 1930. The interpretation of the wing venation and tracheation of the Odonata and the Agnatha. Psyche 37: 245-279. (Translation from Russian. Entomo- logichescogo Obozrenie 18: 145-174, 1924).

NAVAS, L. 1923. Entomologfa de Catalunya. Neurdpters. Fasc. I. Neurbpters propis. 271 pp., 1 pl. Institut d'Estudis Catalans, Secci6 de Ciences, Barcelona, Espaiia.

NEEDHAM, J.G. 1935. Some basic principles of insect wing venation. Journal of the New York Entomological Society 43: 113-127.

RIEK, E.F. & KUKALOVA-PECK, J. 1984. A new interpretation of dragonfly wing venation based upon Early Upper Carboniferous fossils from Argentina (Insecta: Odonatoidea) and basic character states in pterygote wings. Canadian Journal of Zoology 62: 1 150-1 166.

SCHNEIDER, G.T. 1851. Symbolae ad Monographiam Generis Chrysopae, Leach. 177 pp., 60 pl. Vratislaviae.

TILLYARD, R.J. 1916a. Studies in Australian Neuroptera. No. 1. The wing venation of the Myrmeleonidae. Proceedings of the Linnean Society of New South Wales (1915) 40: 734- 751, pl. LVIII.

TILLYARD, R.J. 1916b. Studies in Australian Neuroptera. No. 3. The wing venation of the Chrysopidae. Proceedings of the Linnean Society of New South Wales 41: 221-248, pl. X & XI.

WILLMANN, R. & BROOKS, S.J. 1991. Insekten aus der Fur-Formation von Danemark (Moler, ob. Paleozan/ unt. Eozan?) 6. Chrysopidae (Neuroptera). Meyniana 43: 125-135.

WOODWORTH, C.W. 1906. The wing veins of insects. University of California Publications, Technical Bulletin, Agricultural Experimental Station, Entomology 1: 1-152.

WOOTTON, R.J. 1979. Function, homology and terminology in insect wings. Systematic Entomology 4: 81-93.

Address of author: Prof. Phillip A. Adams Department of Biology California State University Fullerton, California 92834-6850 U.S.A. Bibliography of the

Bibliography of the Neuropterida Reference number (r#): 8999

Reference Citation: Adams, P. A. 1996 [1996.??.??]. Venational homologies and nomenclature in Chrysopidae, with comments on the Myrmeleontoidea (Insecta: Neuroptera). Pp. 19-30 in Canard, M.; Aspöck, H.; Mansell, M. W. (eds.). Pure and Applied Research in Neuropterology. Proceedings of the Fifth International Symposium on Neuropterology (2-6 May 1994, Cairo, Egypt). Privately printed, Toulouse, France. 341 pp.

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